THE CARNIVORES OF WEST AFRICA BY D. R. ROSEVEAR \ .— cSt-T BRITISH MUSEUM (NATURAL HISTORY) z o-]^ /^ A a. O Price £18. so THE CARNIVORES OF WEST AFRICA «AT. HISf. '\ -4FEB1975J ft ^*^ Publication No. 723 THE CARNIVORES OF WEST AFRICA BY D. R ROSEVEAR with II plates in colour by Rita Parsons and 172 line drawings by Patricia Wolseley, Monika Shaffer, Rita Parsons and the author TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1974 -'•^'.^c First published 1974 © Trustees ot the British Museum (Natural History) 1974 Pubhcatioii number 723 ISBN 0565 00723 X Primed in drcit Brilain by Sraplts Printers Limned at 'Hie George Press, Kerterinir Nortiidmptimsliire NOTE Definition of the area taken herein as representing West Africa and of other topographical terms will be found on page 518. "... I should have much stronger expectations than I dare yet entertain, to see philosophy solidly established, if men would more carefully distinguish those things that they know from those that they ignore or do but think, and then explicate clearly the things they conceive they understand, acknowledge ingen- uously what it is they ignore, and profess so candidly their doubts, that the industry of intelligent persons might be set on work to make further enquiries and the easiness of less discerning men might not be imposed on". ROBERT BOYLE The Sceptical Chymisi 1661 "Ce que nous connaissons est peu de choses, ce que nous ignorons est immense". 'iltc Inst words of PIERRE SIMON, MARQUIS DE LAPLACE 1S27 PREFACE The research for this, the third volume on the West African mammals, was made possible by the generous support for three years of the Wellcome Trust, to whose Trustees, Director and staff I am most sincerely grateful. I must also once more express my thanks to the Trustees of the British Museum (Natural History) for the facilities again accorded to me. This present work to all intents and purposes brings to a close an association with the Mammal Room extending over almost forty years, and I cannot too strongly express my deep appreciation of the ever-friendly help and encouragement 1 have received throughout from the members of this section and most particularly in recent times from Dr. Gordon Corbet and all his subordinate staff in their various capacities. I also gladly acknowledge my indebtedness to the Museum's Librarian, Mr. M. J. Rowlands, and his assistants who have repeatedly and readily gone out of their way to give me their help. I have been much encouraged by various correspondents who have found the previous volumes useful, and by the help I have received from collectors and field and museum workers who have kindly furnished me with specimens or items of informa- tion. These are more fully acknowledged in the appropriate places; but I would here mention A.J. Hopson, G. S. Child, R. H. Kemp, Sonia Jeffreys and Gerald Durrell for notes concerning distribution or behaviour ; Dr. P. H. J. van Bree of Amsterdam and Dr. Renate Angcrmann of Berlin for the loan of specimens; and J. J. C. Mallinson for photographs. I owe a very special debt to Theo S. Jones for his unflagging interest in this entire project, for his continual encouragement and for his practical help, despite an over-busy existence, in reading through the typescript and supplying numerous valuable notes from his wide experience in the field. The typescript has been read, too, by Robert Hayman who, although now in well-earned retirement, has once again as so often before kindly offered me helpful suggestions, thus adding yet more to my long- standing indebtedness to his wide knowledge of African mammalogy. No one who studies the morphology and taxonomy of the Carnivora can fail to be deeply impressed with the encyclopaedic knowledge of the Order possessed by the late R. I. Pocock, the outcome of a lifetime's acquaintance as much with living or recently dead specimens in zoos as with mere skulls or dried skins. I should be truly wanting in gratitude if I failed to acknowledge how very much tliis present work owes to the innumerable clearly illustrated papers of this remarkable and indefatigably inquisitive student. All who use this book will agree that it owes a great deal of its value to the painstaking work of the artists who have assisted in its production, Rita Parsons, Patricia Wolseley and Monika Shaffer. I have been fortunate in fmding such willing helpers and I am indeed grateful for their cheerful co-operation in the exacting task of achievingaccuracy and clarity. I must add m)' thanks also to Sylvia Oliver who has almost faultlessly typed the whole of my lengthy manuscript, skilfully and imcomplainingly taking in her stride a variety of unfamiliar languages, technical terms, columns of figures, niceties of spacing and abbreviations, often bedevilled with maddening complexities of punctuation. Though less immediately concerned with the day-to-day production of the vii work, I cannot overlook how much I owe initially to Lord Grey of Naunton whose foresight and faith originally got the whole of this project off the ground; and to Sir Terence Morrison-Scott whose friendly help and persistence in the face of financial difficulties kept it airborne over a period of years. All, however, would have been useless without the never-failing support and encouragement of my wife and her forbearance through several decades of collecting, skull cleaning and, ultimately, printer's litter. Lastly I make a wholeheartedly sincere acknowledgment of my indebted- ness to Mr. Stanley Raw, F.R.C.S., but for whose surgical skill this work would never have reached its conclusion. To all of these, as well as to many other unnamed friends and helpers, I am more deeply grateful than words can properly convey. It is only to be hoped that this account of a truly fascinating group of animals goes someway to justifying their faith, and that, in a region not ver\^ rich in local literature, the work will be found to fill a gap and continue to serve a useful purpose for some years to come. The subject is a complex one, far more so than is commonly imagined even by those who justifiably claim a considerable knowledge of the animal world. Those who study the often involved taxonomic sections will gather something of this author's disquiet regarding mammal- ogical taxonomy in general, with its insistence on reference to anciently named types often chiefly notable for their ridiculously sketchy diagnoses, and with its placid acceptance into an already encumbered nomenclature of new names — without question, until it is too late and synonymy is, under the present system, burdened with them for eternity. Turning from the museum to the field it is interesting to note the remarkable change which the last two decades have witnessed, in that the habits and behaviour of African mammals, once the recognized speciality of the professional hunter, have now become a fashionable field of intensive scientific study supported by large fimds and expensive equipment. This, as the exact niches in nature tilled by each species are more clearly appreciated, must result in benefit to the animals themselves — as, indeed, it has already done for some at least of those dealt with in this volume, which not so very long ago v/ere almost universally regarded as harmful and expendable predators. D. R. ROSEVEAR Hartley Wintney Hampshire April, 1972 CONTENTS Page Preface ... List of Illustrations Checklist General Introduction Canoidea Canidae. Caninae Simocyoninae mustelidae mustblinae . Mellivorinae lutrinae Feloidea. ViVERRIDAE Viverrinae . Paradoxurinae . Herpestinae. Hyaenidae Felidae: Felinae Glossary of terms . Note on the area taken Note on vegetation References Index AS West Africa Vll X I 5 29 30 34 75 92 93 no 128 160 161 164 229 239 341 373 513 518 519 5^4 538 ILLUSTRATIONS Fig. I. 3- 4- 5- 6. 7- 8. 9- 10. II. 13- 14- 15- i6. 17- i8. 19- 20. ai. .11.26, (J, X I : i; TypRal carnivorous teeth, showing the position of closure: a. right upper jaw, palatal view above, lateral view below; b. right lower jaw, lateral view above, dorsal view below. Feiiiicais zcida, B.M. No. 25.5.12.22, $, x 2 . Carnassial teeth, lateral and surface views; a. right upper; b. right lower. Fclis iihir^driici aircmis, B.M. No. 67.1429, ,^, x 3 . Golden lackal [Cnnis aureus) .... Ciiiiis aureus: skull, B.M. No. 21. 2. 11. 28, 5, x § . Side-striped Jackal {Catiis adustus) Civus athistus: skull, B.M. No. 28.6.3.8, sex ?, x | Fenneais zerda: skull, B.M. No. 1939. 1746, ?, x i Vulpi's rueppelli: skull, Type ot caesia. B.M. No. 21 Hunting Dog [Lycaoii pictus) .... Lycaoii pictus: skull. Type o( sharicus, B.M. No. 7.7.8.74, view ......... Lycaoii pictus: skull. Type of sharicus, B.M. No. 7.7.8.74, & dorsal views. ....... African Striped Weasel {Pcecilictis lihyca) and Striped Polecat {Ictony: striatus) .......... Ictcnyx striatus seiicgakusis: skull, Type, B.M. No. 50.7.8.38, $, lateral view ......... Ictonyx striatus seiiegalensis: skull. Type, B.M. No. 50.7.8.38, ?, palatal & dorsal views .... Poecilictus lihyca: skull. Type of rotlischiUi, B.M. No. 25.5.12.26, cJ, Mellivora capctisis: skull, B.M. No. 26.8.7.1, J, x |; lateral view Mellivora capciisis: skull, B.M. No. 26.8.7.1, cj, ^ |; palatal & views ....... Ratel or Honey Badger [Mellivora capeusis) . Mellivora capensis: dorsal coat patterns, showing the extent of white in the various forms: a. leucouota, b. buchauaui, c. concisa, d. signata, e. cottoni Lutra uiaculicollis: skull, B.M. No. 23.1.22.44, sex ?, x i ; lateral view Lutra iiwculicollis: skull, B.M. No. 23.1.22.44, sex ?, .• i; palatal & dorsal views ............ Lutrinae: posterior cheekteeth; top row, right upper p'^ and m^; bottom , ■ 2: a. Lii/cd omnf/ico///.';, B.M. No. 23.1.22.44; No. 10. II. 25. 2; c. Acnyx [Paraonyx) cougica. lateral palatal X 2; > 2; ^3- 24. row, left lower hii and m; b. Aoiiyx capeusis, B.M. B.M. No. 1938.9.29.8 Aoiiyx [Paraoiiyx) cougica: view .-iouyx [Paraonyx) cougica: d< dorsal views. skull, B.M. No. 1938.9.29.4, sex ?, x |; lateral skull, B.M. No, 1938.9.29.4, sex ?, x |; palatal Page 16 17 37 45 SI 53 59 68 77 80 81 97 98 99 107 114 115 119 1^5 144 145 148 156 157 ILLUSTRATIONS African Civet {Civettictis civetta) ...... Civettictis civetta: skull, B.M. No. 59.941, $, x f ; lateral view Civettictis civetta: skull, B.M. No. 59.941, ?, x f ; palatal & dorsal views Genetta: left bullae, x 3: a. G. genetta (? ajra), B.M. No. 9.11.2.34, S b. G. geneltoides, B.M. No. 46.397, (J; c. G. cristata, B.M. No. 39.323, J Geueffa iCHc^a/eo^-w: skull, B.M. No. 1939. 1766, c?, X i. Pdirtiia nVZ/rtrJic'/ii; skull, B.M. No. 98.3. 19.11, cJ, X I . Naiidinia binotata: skull, B.M. No, 48.814, sex ?, X I ; lateral view . Naiidinia hinotata: skull, B.M. No. 48.814, sex ?, x i ; palatal x dorsal views Herpestiuae: illustrating the two contrasting positions of the cheekteeth a. p^ well anterior to the root of the zygoma {Ichneumia albicauda, B.M No. 25. 5. 12. 13); b. posterior corner oi p* about level with the zygoma [Xeiwgale tiaso, B.M. No. 10.6. 1. 14) Miingos gainbiamis: skull, B.M. No. 36.10.30.12, cJ, x | Miingos gambianus: bulla, x 2 . Herpestes ichneumon: skull, B.M. No. 9.1 1.2. 10, sex ?, x i Crossarchus obsciirus: skull, B.M. No. 48.841, sex ?, x f. Atilax palndinosHs: skull, B.M. No. 13.4.1 i, (J, x i Iclmetwiia albicauda: skull, B.M. No. 12.4.3.3, (Ji X I Galerella sanguinea melanura: skull, B.M. No. 35.1.30.53, cJ, Galerella sanguinea: bulla, x 2 . Galeriscus nigripes: skull, B.M. No. 20.10.26.2, (J, X i; lateral view Galerisctis nigripes: skull, B.M. No. 20.10.26.2, i'.'; (Pucheran, 1855) Typical Black-footed Mongoose Genus XENOGALEj. A. Allen, 1919 Greater Long-nosed Mon- gooses X. luisc (de Winton, 1901) Greater Long-nosed Mon- goose A'. II. iiaso (de Winton, 1901) Typical Greater Long- nosed Mongoose Genus LIBERIICTIS Hayman, 1958 -Liljcriau Mongoose L. Liiliiii Haynian, 1958 Kuhn's Mongoose Family HYAENIDAE Gray, 1869 Hyaenas Subfamily HYAENINAE Mivart, 1882 Hyaenas Genus HYAENA Brisson, 1762 Striped and Brown Hyaenas H. hychiui (Linnaeus, 1758) Striped Hyaena Genus CROCVTA Kaup, 1828 Spotted Hyaenas C. ciocHta (Erxleben, 1777) Spotted Hyaena Fanul)' FELIDAE Gray, 1821 Recent and Fossil Cats Subfamily FELINAE Trouessart, 1885 Cats Genus FEL/S' Linnaeus, 1758 Cats Subgenus FELIS Linnaeus, 1758 True Cats F. Uhyca Forster, 1780 African Wild Cat F. I. haussiJ Thomas & Hiiiton, 1921 Hausa Wild Cat F. /. foxi Pocock, 1944 Mid-belt Wild Cat F. iiiargarita Loche, 1858 Scind Cat F. //;. iiiiriisis Pocock, 1938 Air Sand Cat Subgenus CARACAL Gn^y, 1843 Caracals F. ctiiciail Schreber, 1776 Caracal; Desert Lynx Subgenus LEPTAILURUS Severtzov, 1858 Serval Cats F. scnnil Schreber, 1776 Serval Cat Subgenus PROFELIS Severtzov, 1858 Golden Cats F. iiiiiiitii Temminck, 1824 African Golden Cat Genus PANTHERA Oken, 1816 Great Cats P. /»iii(///.< (Linnaeus, 1758) Leopard P. p. paiihis (Linnaeus, 1758) W. African Open-country Leopard P. p. Icopdidiis (Schreber, 1775) W. African Forest Leopard P. Ico (Linnaeus, 1758) Lion Genus ACINONYX Brookes, 1828 Cheetahs .4. yi(/iii/ir.< (Schreber, 1775) Cheetah GENERAL INTRODUCTION The system of classification adopted throughout this work is in the main that of Simpson (1945) with relatively minor amendments in the lower taxonomic ranks. That author divides the Euthcrian (placental) mammals into four cohorts, the Carnivora forming part of the cohort Ferungulata, the other three cohorts being the Unguiculata, comprising the insectivores, chiroptera and primates; the Glircs, consisting chiefly of rodents; and the Mutica, the whales and dolphins. Cohort FERUNGULATA Simpson, 1945 As far as recent, living, mammals arc concerned the Cohort Ferungulata covers the following Orders: Carnivora (Flesh-eating mammals), Tubulidcntata (Aardvarks), Proboscidca (Elephants), Hyracoidca (Hyraxcs, known also as Dassies or Conies), Sircnia (Manatees, Dugongs), Perissodactyla (Odd-toed Ungulates) and Artiodactyla (Even-toed Ungulates). From the point of view only of existing mammals some of these seem to make strange bed-fellows; especially in that the carnivores, that is to sav the wild cats, dogs, genets, mongooses and so forth, should be regarded as having any kind of near relation- ship with the artiodactyls or antelopes and their close kin — in relation to which, both superficially and in general mode of life, they appear not only to have nothing in common but to stand, rather, at opposite poles. The explanation of this seeming paradox lies, of course, in the fossil record, with the complications of which this present account cannot involve itself beyond the brief statement that differentiation from a common ancestor into what are now broadly the hunter and the hunted took place in early Tertiary times. It must be further added that Simpson's views on this matter are not those of all palaeontologists. The Ferungulata are regarded as comprising five Superorders, of which we are here concerned only with the Ferae. These latter may be broadly regarded as covering beasts of prey, the remaining Superorders consisting, by and large as far as living mammals are concerned, of herbivorous, or occasionally insectivorous, species. Superorder FERAE Linnaeus, 1758 Order CARNFVORA Bowdich, 1821 On the other hand, the Ferae comprise a single Order, named by reason ot the predominantly predacious habit of its constituent members the Carnivora, from the Latin auo, cainis flesh, and vow to devour. The animals included under this heading are divided into two Suborders, the Fissipeda or land-based carnivores, and the Pmnipedia, which spend the greater part ot their active lives in the sea, upon which they are wholly dependent for their food. The land carnivores are to be found everywhere with the exception of the soutliem polar regions and a few islands. The pinnipedcs, that is to say the seals and their kin, are of wide distribution in the oceans of the world but onlv one Tlir CAUNIVdlllA (H WIST M-IilrA genus, Moiuwhns, is partly tropical, and tliat is not known to occur withiji the Innits chosen for this book, the monk seal [M. iiioiiachiis) being recorded no further south along the western coast ot Africa than Cape Blanco (Rio dc Oro). This present account, therefore, is in effect concerned only with the Fissipeda. It must be added that there is an increasing tendency amongst systematists to accord to these two divisions full ordinal, rather than subordinal, rank; but though there are plausible arguments to support this such a classification does, in fact, tend to obscure the close relationship that exists between the two sections. Suborder FISSIPEDA Blumcnbach, 1 791 The two Suborders of Carnivora are named from, and to some extent founded on, the general build of the feet. The Fissipeda are characterized by having the digits more or less clearly independent of each other, even though in some cases connected by interdigital webs. The name is, tor this reason, derived from the Latin words fissuiii cleft and pes, pedes foot. In the Pinnipcdia, on the other hand, the digits, notably those of the forclimbs, in conjunction with the contiguous bones fiuiction as a single unit adapted as a paddle for swimming, encased for this purpose in a common integument — though there are, of course, exceptions of greater or lesser degree to this broad plan. The name, thus, is built from the Latin word piiiiui a feather because the structure, in general shape, and in the unitary way it functions as a means of propulsion, recalls a bird's wing rather than the normal run of mammalian foot. The fissipede carnivores consist, in West Africa, of wild dogs (i.e. jackals and foxes), the hunting-dog, striped weasel, striped polecat, honey badger, otters, civet, palm- civet, genets, mongooses, hyaenas and wild cats of various kinds including the lion, leopard, cheetah and others less well-known. These are disposed in j families, 27 genera and 45 species embracing about 59 different forms. Of the two families which are lacking entirely from the African famia the more notable is the Ursidae or bears. The other, the Procyonidae, though of considerable importance in both northern and southern America and in eastern Asia, comprises less widelv known animals such as the raccoons, coatis, kinkajou and pandas. General description. The carnivores, especially the fissipede carnivores, are an extremely interesting group. They are certainly more intimately associated with man, in the capacity of close friend or bitter enemy, than any other Order of mammals. They have fairly big brains and are more often than not highly intelligent; they live by their wits; and under domestication, therefore, they are quick to grasp a situation and to learn. Without question, both the domestic cat and the domestic dog have, each in its own special way and ni return for assured food and snug shelter, turned themselves into man's most intimate and seemingly luidcrstanding associates. Further, many ot the normally wild species take iniexpectcdly kindly to captivity and become amazingly responsive to close association with human beings. Yet in the wild state there are no more unremitting foes to man's cattle, sheep and poultry, as well as to wild game, than the majority of fissipedes. The suborder exhibits a wide range of size, varying in West Africa from a GENERAL INTRODUCTION little weasel with a head & body length of only a few centimetres and weighing about 200 grammes (i lb) to an animal of the vast bulk of a lion which may weigh 200 kg (4 cwt) or more. Between the different families there is a good deal of variety of shape; but in a general sense the bodies throughout are mostly relatively slender, long, of subcylindrical form, frequently without any very marked constriction at the neck, highly muscular and often, especially in the cats, of extreme suppleness and agility. Variety of external appearance is brought about by differences in the shapes and relative proportions of the head, the tail, the legs, and the feet, apart from any question of texture and pattern in the pelage, hi much of the suborder the common form of head is long, tapering m a greater or lesser degree to a pointed muzzle; but the cats differ markedly from this general shape in having a much rounder head with a notably short muzzle. Ears. Ears often form a very conspicuous feature. They are long and upstanding in the dogs and hyaenas, sometimes sharply pointed; but in the scrval cat they are very rounded besides being tall and broad. In other cats they may be rounded but of much less size. They reach their greatest reduction in the ratel, otters and mongooses. The backs are clad with short hairs, the inside of the pinna with long hairs, often densely; and in the cats the backs mostly bear a black or black and white patch, quite charac- teristic of the species. An apical tuft exists in the caracal. A curious aural character occurring in large sections of the Carnivora is a doubling of the rim of the pinna near the base of the outer margin, forming a little pocket, known as the bursa, with an anterior and a posterior flap. The latter is usually semi-lunar; the former emarginate. A bursa is always present in the Canidae, Felidac, Viverrinac and Paradoxurinae ; it is completely absent from the Hyacnidae, the West African Mustclidae and the Hcrpest- inac. Near the base of the inner margin of the pinna is a small process known as the tragus; and opposite this on the outer margin is another process, the antitragus, having a notch into which, in some cases, the tragus can fit tightly and help to cficct closure of the entrance to the ear passage. On the inner face of the pinna are three or four shallow folds or ridges crossing it in various directions. These also, in those species which fmd it necessary, aid in closing the ear when the piima is collapsed and folded. The termin- ology for these is not well fixed. Rhinarium. The rhinarium in this suborder, that is to say the region at the extreme anterior end of the muzzle surroiuiding the nostrils — the "nose" in common parlance — is naked over a larger or smaller area and, in the Canidae at least, has a slow secretion of mucus. The nostrils themselves may be narrow and slit-like or rather more open and comma-like; and the whole form of the rhinarium is gcncrically, or even specifically, characteristic. This has been studied and illustrated by Pocock in a number of published papers; and there are figures also in J. A. Allen (1924). Eyes. The eyes of the carnivores are mostly of a moderate, or sometimes relatively small, size; and they are situated well to the fore on the head, vision being thus effectively binocular, permitting that accuracy in the judgment of distance necessary to the successful striking of prey. Acuity of vision is in general good but in fact varies quite appreciably not only between species but amongst individuals too. It is a matter of common observation that some breeds of dog, or different members of a single 8 TUP, CARNMVOKrS OF WHST AIIUCA breed, can see much better than others; and tliis apphes as much to wild species ot carnivores as it does to domestic animals. There is Httlc doubt that cheetahs liave con- siderable sharpness ot sight over long distances, whereas otters are in a different category and, indeed, have eyes probably far better adapted to seeing under water than on dry land. Yet however acute the power of vision may be, it is in many, if not all, cases aided to a great extent by the faculties of smell and hearing, both of which are better developed than m human beings, sometimes to an nicomparably greater extent. Equipped with a highly sensitive sense of smell, a carnivore can build up a mental picture of its surromid- ings almost beyond the comprehension of man. Sound frequently has no very great range, may be easily obstructed or blurred, and is often transitory; but scent sometimes lasts for long periods and may carry fir, and it enables a suitably equipped animal to "see round a corner" where sight is of no avail. Its chief drawback lies in the degree to which it is affected by air currents; for whereas it may lie for long in sheltered under- growth it will in open cotmtry certainly be carried considerable distances by the wind. Carnivores, therefore, if circumstances permit choice, approach their prey from down- wind, securing the double advantage of having the scent of their proposed victim brought to thcni whilst their own is borne away. The canidae are notable possessors of a very highly developed olfictory sense which they generally rely on in preference to sight and, where possible, use to confirm what their eyes appear to have told them. Sensitivity of nasal perception and analysis amongst some of the Canidae has been shown to be of such delicacy that one second's holding in a man's hand of a billet of wood suffices for a trained dog to be able to select it from a pile of numerous similar pieces. Cats are more dependent upon hearing as an auxiliary to vision. Many carnivores are able to sec unusually well in poor crepuscular or nocturnal light conditions, a power very necessary to the securing of prey in these circumstances. Some night-active species have irises that arc highly and rapidly responsive to variation in light intensity. Accommodation in this respect is achieved in the most typical cats by a pupil which in response to a lessening intensity can change from a narrow vertical slit, customary for a bright light, to a wide circular opening. By no means all felines have this power; and in some the pupil remains, within narrower limits, broadly elliptical. It is interesting to note that eyes of a similar type with a vertical slit are found amongst the owls, also nocturnal hunters; but the horizontal pupils of ungulates and kangaroos do not serve the same function. The effect of this enlargement of the pupillary aperture in dim conditions is aug- mented in the carnivores by the possession of a second means of increasing the efficacy of weak light. Anyone who has observed cats or dogs at night is acquainted with the remarkable way in which the eyes assume a green or yellow phosphorescent luminosity. Since mediaeval times this somewhat ghostly glowing ot the eyes has been popularly held to be something to do with "seeing in the dark" by the mysterious projection of beams as from a lamp to illuminate the object looked at. This last, of coiu'se, is not so. The effect is due to a subcircular area of tissue at the back of the eye situated immediately behind the retina around the optic nerve, having no common name but known in anatomy as the tapctum hicidiiiu. This possesses the property of virtually doubling the effective intensity of such poor light as may exist by reflecting it back again through the GENERAL INTRODUCTION receptive cells ot the retina instead of its being at once absorbed in the posterior layers of the eye as it is in organs unprovided with this structure. It does thus enable an animal to see virtually twice as well in the near-dark as it might otherwise do, but not by the generation of light in its own eyes. It follows that it is quite inoperative in absolute dark- ness— a condition which rarely exists in nature — and the eyes would emit no glow in such circumstances. This dual use of feeble illumination is, of course, of considerable advantage but nevertheless has the drawback ot some loss of sharpness of vision (Tansley, 1965). Eyes of this kind arc found also amongst whales and ungulates. A nictitating membrane, or so-called third eyelid, is fairly well developed in some of the carnivores, especially in the cats, though it is never wholly and constantly functional as it is in reptiles and birds. Vision varies markedly in its sensitivity to colour. Diicker (1957, 1964) found Genella and Fclis to be totally colourblind, but the mongoose Hcrpcstcs ichneumon to be relatively well equipped, having a positive colour sense though exliibiting difficulty in distinguish- ing between nearly related hues such as red and orange or green and blue. Legs and feet. Legs are possibly responsible for a greater variety of fissipede form than heads. They are in some cases long or very long, adapted almost wholly to running, as in the dogs or that in some ways rather dog-like feline the cheetah. In these cases the feet are digitigrade and carry moderately straight, non-retractile claws. In the typical cats the feet are also digitigrade but the legs are relatively shorter and a little stouter with greater flexibility of movement. The feet are armed, with strongly curved, very sharp claws which are retractile within sheaths to preserve them from abrasion in rmuiing and walking but extensible when a furm anchorage is required, either in climbing, taking off for a spring, or in clinging to prey. Such limbs are adapted rather to a bounding than to a rurunng gait, to slinking with lowered body, to climbing trees and other rough surfaces, and to performing powerful leaps. Between these two extremes lie a number of forms, mostly with short legs suited to a trotting gait and sometimes to climbing, either with fixed or, in the genets, partly retractile claws, digitigrade, semi-plantigrade or, in the ratel, nearly fully plantigrade. Bears, which do not occur as wild species anywhere in Africa, provide the best example of carnivores that are fully plantigrade, that is adapted to walking with the whole of the foot from toe to heel in contact with the ground. There may be four digits on each foot, or five, or a combination of the two. Where the ist digit is present it is mostly widely separated from the rest, more particularly on the hindfoot, where it is far removed from any possible contact with the ground. In the Canidae this is popularly known as the "dew claw", possibly because of its ephemeral nature since it often disappears in adolescence. Although the digits, unlike those of the pinnipcdes, remain virtually independent they are nevertheless often joined by webs at least basally though in semi-aquatic species they may be extensive. A notable feature of the fissipede foot is its sole, consisting largely of rather rubbery, mostly independent cusliions, sometimes collectively referred to, for the fore and hindfeet respectively, as the palmar and plantar pads, though these terms have also a more restricted application as explained shortly. These pedal pads, in general, fall into three categories: those situated below the distal portions of the digits, which may be THE CARNIVORES OF WEST AFRICA conveniently referred to as the digital or apical pads; those lying in the middle ot'thc foot, basically comprising tive pads surrounding a central depression but often reduced in number or indistinguishably merged, termed by J. A. Allen (1924) interdigital pads but more often called the palmar (forefoot) and plantar (hindfoot) pads. Finally, posterior to these, there is often a narrow, longer or shorter metacarpal or metatarsal pad, frequently compound. Pads follow set patterns in ditferent species or genera and are often wholly diagnostic; but family resemblance is more unitonn in the Felidae and Canidae than in the \'iverridae and Mustelidac (Pocock, I9r4b). Tails. The tail in the fissipedes is a structure of considerable variet\' and is, with few exceptions, usually a showy as well as a characteristically formed appendage. In the cats and the genets it is for the most part long and of considerable suppleness of move- ment; and it is clad with hairs of subequal, moderate length resulting in a more or less cylindrical structure. In the dogs it is far less flexible, capable of little more than stiff side to side or up and down motion from the root only. It is sometimes uniformly long haired and of striking bushiness. hi the mongooses the tail is tapering, in some species markedly so; in some it is rclativelv short-coated, in others of somewhat asymmetrical coutour owing to the drooping carriage of its long hairs. The lion's tail is tuiusual amongst the African fissipedes in not only being close-haired but in carrying also a promir.cnt. blackish, long-haired terminal tuft — hidden amongst which is the famous "thorn" of hard skin with which in ancient times it was reputed to lash itself into fury before attacking its prey. On a far smaller scale, a terminal tuft, sometimes black but much less bushy than in the lion, occurs also in some of the mongooses. Apart from variety of shape, tails in this suborder may be unicoloured. bicoloured, annulated, spotted or speckled. Scent glands. One of the most notable features of the carnivores is the possession by many members of the Order of scent glands. These arc mostly situated in the region of the sexual organs or the anal area, and their functions are recognition, sexual attraction and stimulation, demarcation of territory, warning or defence. As an example of the last in West Africa the striped polecat {Ictoiiyx) can, if alarmed, by the deliberate and abrupt contraction of the circumscribing muscles emit to a surprising distance a spray of fluid with an extremely disagreeable and long-lasting stink. Despite the existence of glands, in many cases, including that of this striped polecat, little or no offensive odour is under normal circumstances apparent to human beings; but in some animals, notably some of the dogs and h)-aenas, a fetid stench unremittingly pervades the whole animal bv a continual imcontroUed oozing of the offensive secretion. Involuntary emission of odour also in many species accompanies the acts of defaecation and micturition, both of which functions, but especially the latter, are in consequence used to indicate the boundaries of a territory and publish a warning against trespass by prospective intruders. In other cases, establishment of ownership, or other scent-borne message, is conveyed by the rubbing of circumanal glands against trees and rocks or by dragging the anus across the ground. Scent-marking in the Canidae, especiallv in respect of postures adopted, has been dealt with by Kleiman (1966). The precise siting of fissipede scent glands is, within a relatively limited bodily reginrt vcr)' varied. They may be associated with the anus, either within the rectum oronexe,- GENERAL INTRODUCTION II ally aroimd the orifice (circumanal); between the anus and the scrotum or vulva (perineal) ; in association with the prepuce (preputial) ; between the scrotum and the penis (prcscrotal) ; on the tail, above or below, (caudal); in hyaenas, between the root of the tail and the anus (supra-anal); or, in the palm civet, anterior to the vulva and the penis (pregenital). Usually, but not invariably, the position follows a family pattern. The secretion is sometimes liquid, sometimes of a waxy consistency as in the case of the civet. It is interesting to note that this latter secretion, while nauseating to humans when in concentration, is in dilution pleasing; and "civet", extracted with a spoon from the glandular sacs of captive animals kept for the purpose, has for many centuries commonly been one of the ingredients of a number of commercially successful per- fumes in Africa, Asia and Europe. In connexion with the wide occurrence of scent glands it must be remembered that many carnivores, except at breeding time, lead a more or less solitary existence and some more reliable means than chance encounter is needed to enable the sexes to find one another at considerable distances. The emission of powerful and significant odours that may cling for long to undergrowth or be carried far on the wind, in conjunction with a higlily sensitive and critically analytical sense ot smell, is an ideal way of sur- mounting difficulties of time, distance, intervening obstructions or darkness that niay render sight useless in bringing the sexes together. Sound, though sometimes similarly used as a remote inter-sexual signal, is often not so effective in overcoming difficulties of time and space as well as having a less emotive impact. Mammae. The mammary glands are, so far as known, mostly purely abdominal and number between one and five pairs except in the cheetah, where they are far more numerous. Pelage. The pelage, by its length, texture, colouring and pattern, accounts for very marked differences of appearance not only between families but often between genera and species. There is, unfortunately, no clearly defined and commonly accepted terminology in English for the various types of hair found in mammalian coats. At one extreme there are bristles, or vibrissae as they are more technically termed, which are of circular section and considerably stouter than the greater part of the pelage. These occur most noticeably as the "whiskers", or mystacial vibrissae, on the upper lip; but also, in the carnivores, singly, in pairs or limited groups in other set positions, of which there are five on the head besides the mystacial area. These are submental, on the anterior part of the chin; intcrramal, on the posterior part of the chin; genal, on the cheeks; superciliary, above the eyes; and subocular, below the eyes. Apart trom the mystacial vibrissae not all these are always present; in the fissipedes the interramal vibrissae are lacking in the Felidae alone. Vibrissae occur, mostly singly, on other parts of the body, in some cases on the legs, but more commonly at wide intervals over the back and flanks. The function of these stout and sensitive hairs, wherever they are situated, is to give tactile warning or information. Those on the face and on the body, for example, furnish vital information in the dark when creeping into a narrowing hole or through a restricted rock fissure; the whiskers and possibly other facial vibrissae clearly tell the direction of the wind; and those of the otters may play a part in the detection of prey in ill-lit or muddy waters by revealing currents set up by the powerful 12 THE CARNIVORES OE WEST AERKA swisli ot .1 fish's tail. Tlic mystacial vibrissac arc generally stouter and longer than the others, and sometimes very stitf — as thev are in tlic leopard. Indeed, in some parts of West Africa in the past the whiskers of this animal were looked upon as a certain means of causing death if chopped up and mixed with anyone's food, being reputed to penetrate, or at least cause fatal inflammation of, the stomach wall. At the other extreme ot liair is the underfur, so called because in many mammals it torms the lowest, and often rather insignificant, constituent of the pelage. But this is not always so; and in some species it is the major element — as, for example, the wool ot sheep. For this reason it is not a good term but it is in general use since there is nothing else with which to replace it. Undertur is always of fmc diameter, sometimes extremely fuie; in the carnivores it may be short or moderately long and is otten slightly sinuous. Occasionally it is almost completely lacking, as in the Banded and Gambian mongooses of the genus Miin(;os; but in the majority of cases it is dense; sometimes exceedingly so, as tor example in the otters where it forms, on submergence, a waterproof coat aniongst which air is trapped, thus keeping the body both dry and warm. It is also remarkably dense and, amongst West African carnivores, at its longest in the striped polecat {Icloiiyx) in which it in etiect torms the entire pelage apart from a few widely dispersed lengthy bristles. In this case it probably serves to shield the body from the ettccts ot direct tropical siuishine and hot dry winds. Above the undertur lies the longer outer fur, which may be so abiuidaiit as to conceal the underfur completely or, as in the case just mentioned, it may be merely a widely scattered and comparatively unimportant element. The tormcr is by far the commoner. This part of the pelage is, in a greater or lesser degree, harsher than the undertur, being composed ot stouter hairs though by no means so stout and strong as the vibrissae — to which the unqualified term bristles is best confined. More often than not there arc two distinct elements in this outer coat. The more important and abundant consists of stoutish hairs of round or sometimes slightly flattened section, tapering distally to a slender point but not decreasing much in diameter towards the base. When the fur is turned back they therefore usually stand out against the background ot undertur by size and sometimes by colour. In the carnivores they never exhibit a concavo-convex section commonly found in the Rodentia (Roscvear, 1969). These hairs are in this present work, for want ot any better recognised term, referred to as bristle-hairs. They are often broadly aniiulated with different colours, the tips being almost invariably black, the basal portion very often pale or even white. In many species there is a second element in the outer fur, termed herein sub-bristlc-hairs, shortened to sub-bristles. These, like the bristle-hairs, are longer than the undertur and play their part, too, in overlying and concealing it; but they are of different form, consisting of a long, slender, pale stalk (the petiole) which passes into a broader, terete or slightly flattened coloured blade ending in a darker, pointed tip. This distal part, except under magnification, resembles the bnstle-hair, and the fundamental difference ot form has therefore often been overlooked; the slender proximal halt though ot somewhat greater diameter than the underfur is nevertheless not so disparate as to stand out clearly from it when t!ie pelage is turned back as the bristle-hairs do. This gross morphology of the pelage can be seen w ith the naked e\e or iiiuler low GENERAl. INTItODUCTION 13 magiiificatioii. No reference is made here to the surface patterns occurring on hairs since this is a matter for the microscope or cicctroscan and beyond the scope of this present account. It need only be said of tlicse that, though little or no research has yet been carried out into African carnivores, it is possible that they may follow at least generic patterns and might be of use for identification, as between other mammalian groups. It is the surface scales giving rise to these patterns that underlie the property ot close adherence together known as "felting" which takes place when a mass of hairs is suitably beaten. The coat is nearly always plentifully developed to afford a more or less complete covering to the skin; but it may range between very short, as in the lion, to very long, as in some hyaenas and the striped weasel. However, coat length and quality and sometimes the relative proportions of the constituent elements are affected very con- siderably by season, moult and condition of health; and owing to these factors skins in museum collections are often misleading and sometimes not susceptible to valid comparison. Wrong taxonomic deductions have sometimes been based upon them. Little is at present known of these factors in comiexion with the majority of West African carnivores. In some species, apart from the normal coat there may be a well- developed collar or "mane", or a nuchal or spinal crest ot exceptionally lengthy hairs. These growths are affected by age, sex and possibly season. Pelage patterns. Pelage texture follows a general family plan. In the typical dogs it is frequently long and rather harsh; in the cats mostly much shorter and somewhat softer; the otters have fairly short but extremely dense, rather silky fur. Pelage pattern in this Order varies very widely but often similarly follows some sort of family, or at least subfamily, plan, to which, however, there are sometimes notable exceptions. The larger dogs in West Africa have coats characterized by broad, irregular blotches but the foxes are more self-coloured; a speckled, "pepper and salt" appearance occurs in many mongooses, though, again, some are predominantly unicolorous. It is amongst the cats that the widest diversity is found. Spots are common — the bold markings ot the leopard, cheetah and scrval are familiar; and though the pelage of the adult lion is plain its spotted heritage is clearly displayed m the cubs. A more curious incidence of the loss of spots in the Felidac is shown in the golden cat, which within a single species exhibits in the adult both spotted and unspotted forms, a difference which appears to be related to distribution and is thus racial. The West African Viverridac, that is the civet, genets and their km, also have spotted coats. Though spots may in some individuals coalesce into lines, pure transverse striping as the sole pattern as in the tiger does not occur in West Africa; but through the regular disposition of the light and dark annulations of the hairs ot its dorsal fur one of the mongooses is cross-banded. Bold, broad, longitudinal bands are exhibited by the African polecat and weasel. The ratel is unusual amongst the African carnivores in its coloration. Most mammals have the belly white or at least paler than the back; the ratel exactly reverses this; and so to a lesser extent do the striped polecat and striped weasel. Pattern is produced in three quite different \\a\s. The hairs may be unicolorous throughout their lengths as in the polecat and weasel, where they are in separate bands. 14 THt < ARNIVORF.S OI \^ EST APRU A wholK' black or wholly wiiitc. More ohen, bold spots or lines arc produced, as in the cheetah and other cats, by groups ot hairs that are indistinguishable from the rest of the tur in their pale proximal portions but difter in having black ends. Much more rarely, and in West Africa onlv in the single case ot the banded mons^oose referred to in the previous paragraph, the pattern is brought about by the regularity with which each of the dirterent coloured annulatic>ns on the hairs tall together along the back instead ot Knig hcterogencously mixed as in the ordinar^• "pepper and salt" coat. The backs ot the ears, notablv in the Felidae, otten contrast with the rest ot the coat. In this tamily thev may be wholly grey, whollv black, or have bold black markings with or without an additional white spot. Melanism. Melanism is not uncommon, but albinism appears to be very rare, lied as an outstanding feature ot the pelage crops up in a number ot dirterent families ot the carnivores. It occurs in the dogs, where in the foxes it sometimes constitutes the main colour. It is found also in the cats; in the genets, where it is in some cases at least a component of the spots; and in the mongooses. The striking appearance of a leopard skin when it is seen as a trophy m a room makes it seem that such an animal would be very conspicuous and easily picked out in the field; but the reverse is mostly the case; for w'hen one of these animals is at rest in the undergrowth or amongst the foliage of a tree the pattern merges completely into the dappling ot light and shade produced m the vegetation by sunlight. Only when the animal stands fully exposed in the open or is on the move can it be clearly seen. The alarmed surprise of suddenly discovering oneself in unexpectedly close proximity to a leopard in the bush must possibly first be experienced before the real truth of this can be fully appreciated. At birth and soon after, the coat is mostly short and rather silky, the adult markings either absent or very taint; but lions are born with a pattern that is lost at maturity; and in the cheetah the juvenile coat is very long and greyish-white, c]uite unlike that ot the adult. Sometimes, as in the mongooses, the entire pelage is erectile in anger or alarm; sometimes only the hair along the spine. Skull. A great part of the tissipedes have a rather narrow skull m which the total length IS markedly more than the zygomatic breadth, the bramcase otten not a great deal wider than the posterior end of the rostrum. This last is itself relatively narrow and often long and tapering. In the Felidae, however, the skull is clearly rounder, its breadth across the zygomata not a great deal less than its length, the braincasc broad and subglobular, the rostrum very short. Throughout the suborder there are nearly always pronounced, sometimes ilangc-hke, supraoccipital crests, and often, in mature skulls, especially males, a sharp sagittal crest, sometimes short, but often extending the whole length of the braincase. The nasals are always narrow, often relatively short, their upper front margins always situated well posterior of the forward limit of the premaxillae and incisors, thus leaving a very open, anterior nasal aperture lacking a bony roof The slender coiled bony laminae within the nasal cavities, known as the turbinals, which plav a major part in the sense of smell support a total surface area of receptive membrane incomparably greater than that available to man. The zygomata are alvyays strong, sometimes very strong, and this, in conjunction with the cranial crests, is tn give the nccessar\- anchorage to the extremely powerful GENERAL INTRODUCTION I5 biting muscles. Thcjugal boiic plays an important role since it torms the greater part of the anterior half of the arch, the maxillary zygomatic process being relatively short. With few exceptions as in the otters, the infraorbital canal is of comparatively small size. The orbit is incompletely ringed with bone, there being a longer or shorter gap between the postorbital process from the frontal and, in the majority of genera, an upwardly curving process arising from the jugal. In the ratel and some otters one or both of these orbital processes may be poorly developed or virtually absent. The bullae may occasionally be small but are more often large or very large and betoken a highly developed sense of hearing. They furnish a point of considerable taxonomic interest, providing a distinction between the two superfamilies into which the suborder is separated. In the Canoidca the bulla is simple, consisting of a single cavity. In the Fcloidca, on the other hand, it is partially or almost completely divided into two chambers, of varying relative sizes, by a septum which can usually be clearly seen in a prepared skull through the auditory meatus. The division of the bulla into two parts can generally be detected externally as a shallow fiurrow curving around the body of the structure. It must be added, however, that this classic conception is ques- tioned by Hough (1948) by whom the canid auditory region is regarded as more like the fclid hi its essential structure than has been commonly supposed. This paper furnishes details of this region in the Canoidea in comparison with those of some other present and past Carnivora and draws conclusions regarding their significance in the phylogeny of the Order. The bony palate sometimes terminates about the level of the back of the molar row but very often extends far posterior to this. The anterior palatal foramina are of no great size, round or oval, and lie more or less between the canines; the posterior foramina, often two pairs, arc situated at very different points of the palate according to genus. A highly important feature of: the skull is the form ot the mandibular condyle, that IS the hinge between the lower and the upper jaws. In many mammals, including human beings, this is a fairly flexible affair permitting movement of the lower member in various directions, up and down, from side to side, or back and forth, enabling the occlusal surfaces of the upper and lower teeth to alter their relative positions and to slide across each other, and thus produce a grinding effect. This is possibly best seen in the ungulates in the action known as "chewing the cud". In the carnivores, on the other hand, the hinge consists of a long subcyltndrical condyle on the mandible firmly embedded in a complementary-shaped receptacle, the glenoid fossa, in the upper jaw, completely inhibiting any side to side or back and forth action, ensuring that the jaws close, like a comparable liinged door, in one firmly fixed position. Without this, the cutting action of the carnassial teeth, as described below, would be as ineffectual as a pair of scissors with play at their axis of coupling. Dentition. The dentition is always, even in the smallest carnivores, patently ot a powerful nature (fig. i). All the roots are closed, that is to say that once any tooth has reached its appointed size there is no further growth to replace wear as, for example, in the open-rooted gnawing-teeth of the rodents. The incisors arc invariably f . They arc relativelv small but the outer one, especially above, is often larger than the others. 16 THE (AUMVIMIIS (II U 1,M M 11 1 ( A "^cg-Q-O o ^ ^--crrcs-o _Q^^ Fii;. I. Typical carnivorous teeth, showing the position ot closure: a. right upper |a\v, palatal view above, lateral view below; b. right lower jaw, lateral view above, dorsal view below, rcitiurns zcnia, B.M. No. 25.5.12.22, V, -■ 2 sometimes appreciably .so, even to taking on a subcaniiiitonii appearance. Tlic jaw is nearly always strikingly dominated by the exceptionally tall, strong, curved canines — the "dog teeth" in fact — one above and one below on each side; but the cheekteeth are also ot a remarkable and unique torm. "Cheekteeth" is a convenient term used tor all the teeth which lie posterior to the canitie when it is not desired to ditierentiate between premolars and molars. The symbols /, c, p and in are used to denote incisors, canines, premolars and molars respectively, a tigure being added above or below as index or sutiix to indicate whether the toodi is m the upper or lower jaw and its position in each category reckoning trom trout to back. Thus, p^ is the third pre- molar from the tront in the upper jaw; 1112 tlic second molar in the lower jaw. The total number of cheekteeth ni the Carnivora is very variable according to family or genus, ranging in West Africa between 14 and 26 tor the entire mouth. The premolars may be 3 or 4 above on each side ot the jaw, and 2, 3 or 4 below; the molars i or 2 above and i, 2 or 3 below. Complications exist in that some teeth are occasionally deciduous, being shed with advancing age, the jaw in such cases appearing not to correspond to the accepted dental formula. Moreover, in a few species certain teetli seem to be in tlic process of evolutionary loss, forming components of the dentition in some specimens but not in others, without being actuallv deciduous. 'I'lie dental formula is thus variable. GENFRAr, INTRODUCTION 17 Fig. 2. Camassial tcctli, lateral and surface views: a. right upper; b. right lower. Fclis marfiarita ainiisis, B.M. No. 67.1429, o. - 3 The most characteristic feature of typical carnivorous cheekteeth is the modification of some of them into a highly efficient, sharp, cutting ("sectorial") form, adapted to dealing with masses of flesh and bone. This chiefly concerns the last premolar of the upper jaw (p*) and the first molar of the lower jaw (nn). The crowns of these are either almost wholly compressed from side to side or have at least some of their lobes thus flattened, the occlusal surfaces being in this way reduced to keen, knife-like edges which on closure slide over the face of the opposing tooth in the manner of a pair of scissors. The upper teeth always close over the outside of the lower. They are maintained in the essentia! firm close contact necessarv' to effective shearing by the rigid condyles, as explained earlier. Because of their flesh-cutting function these teeth arc kno\\ n as the "carnassials". In this work, as in most others, the upper carnassial is alwavs reckoned to be the 4th premolar (p*), it being assumed when only 3 premolars actually occur in this toothrow that it is the ancestral ist premolar (p^) that has disappeared and that the anterior premolar as it today exists is therefore p-. The lower carnassial is always mi. The crowns of the cheekteeth lying immediately anterior to the carnassials are tor the most part als(5 laterally compressed and pointed and thus play their part too in a shearing action though in a relatively minor way. Those teeth, if any, posterior to the carnassials are of a less specialized kind suited rather to crushing than cutting, though they are often of much reduced size and then almost functionless. The detailed form of the carnassials varies somewhat from genus to genus ; but, speaking in broad terms, the upper one consists of a laterally compressed outer blade divided into cither two or IS Tin; (AUNivouis oi wcsr atuk a thicc j>oiiiti.'(.l Liisps, toi;(.tlicr uitli .1 Initial, miali lower, cusp whicii plays no role in the sectorial attion (tig. 2). Tlicrc are three roots; two below the blade and the third beneath the inner cusp. The lower carnassial is onl\- two-rooted but has, in general, a broader and more complex crown tliaii the upper one, comprising, basically, six cusps, though most are indistinct and several may be lacking. There is a good deal ot variety ot sliapc and disposition of these cusps; but most commonly it is the two anterior buccal ones wlijcli arc enlarged and laterally compressed to form tlie sectorial blade that exercises a shearing action against the upper. The virtual absence of grinding surfaces in the t\pical carnivore dentition means that cluinks of flesh are cut off and swallowed whole. But not all tissipede teeth conform to this predoniinautly sectorial shape, which is best exemplified in the cats and dr extermination, brought into use to cc^mbat predation upon valuable tarm stock. Canidae are to be tound in verv diverse ecological settings, trom the arctic snows to hot and dry deserts. In West Africa wild species, with whicJr tliis account is alone concerned, do n the individual hairs, which ma\' .iKo in luher ways var\' amongst tliem- selves in eoloiu or .it least tone. Areas ot black or white mav torm a well-marked feature in i ert.nii species; but there is scarceK' ever anything that could be regarded as ,1 formal, fixed and regular pattern ot stripes or spots. Even in the so-called side- slriped laik.d of West Atrii.i, tliL- bl.ick b.nul on the tl.ink, which gives rise to the n.niie. is shiucw ii.it iiu iviisi.nit .nid i>lteii obsi iiii . 'ilie extreme bushiness ot the t.nl C'AN'IDAE 31 (Plate i) gives It a striking appearance and and a characteristic shape that furnishes an immediate point ot recognition. Skull. The canid skull (tig. 4) is long and narrow, tire elongated rostrum reflecting the wcli-kiiown sharp face of all but certain highly specialized domestic tonus of the family. The braincase is rounded ; but, considering the reputed intelligence ot the Canidae, seems disproportionately small and to constitute a relatively minor part of the total skull volume. As tar as West Atrican species are concerned there is always at least some development ot sagittal and supraoccipital crests except in Fciiiicciis (fig. S) in v\hich the former is almost completely lacking, and the latter relatively insignificant, hi Lyciioii (fig. 10) the sagittal crest is tall and knite-like; in Caiiis (fig. 4) it is low; and in Viilpcs (fig. 8) it IS restricted to the extreme posterior part of the cranium. The frontal region is marked bv fairly well-developed, subtriangular postorbital processes, otten torming a slight flange over the orbit itself but lacking any finger-like extension; and since the jugal process is also short there is thus a wide gap in the circumorbital ring. The nasals are long and narrow and always reach back at least as tar as the front of the orbit. The up-curved zygomatic arch is of moderate strength, the jugal bone pla\ing a major role in its constitution. The palate broadens posteriori)-, the cheekteeth, from front to back, curving gradually out and then, more sharply, in again, the dental row from the canine to the posterior molar thus forming a flattened S. The mesopterygoid fossa is mostly broad and deep. The bullae in the West African forms are large or, in Fciiueais, extremely large. In view of the manifest importance of the teeth in this family and the relatively strong construction of the upper jaw, the mandible appears unduly slender and weak, the slightly up-curving rami being, except in Lycaoii, shallow; but it must be remembered that there is very little chewing carried out in this family, the teeth being mostly used for severing chunks ot flesh which are swallowed whole. The coronoid process rises steeply, high above the t\pical carnivore sub- cylindrical condyle. The angular process is small and sharply divided from the main body of the ramus. Dentirion. The dental formula in the Canidae is basically jYjt 4- throughout the family with a few (extralimital) exceptions, of which Otocyoii, the bat-eared fox, occurring from Ethiopia to Cape Province, is the onl\- African example, hi this genus the cheekteeth may be ^ or |. In the West African genera, with which this work is concerned, the incisors, though well-developed, are relatively small, their sharp cutting edges sometimes trilobed. The canines — a name, of general use throughout mammalian dental nomenclature, deriving from their prominence in this family — are always very tall, recurved, strong and tapering to a sharp point, ideally suited, to sinking deep into flesh and maintaining a secure anchorage upon a struggling prew Their build prevents their plaving any further role in mastication. The post-canine gap is at most of verv moderate size, and in Lycaoii non-existent. All the cheekteeth (fig. 6) have cingula, most promiiK-nt!\- developed in the posterior part of the toothrow. The four premolars ot the upper jaw nicrease progressively in THE CARNIVORES OF WEST AFRICA oe a size troin the tirst to tiic last, /)', tlic upper caniassial. beiiii; always the tallest check' tootii. All, when iiinvorii, are sharp and ot triangular profile, though there may b secondary, hir lower, cusp anterior or posterior, or both, to the main one. p^ has a single root, p'~ and /!■' two each longitudinally sited; p^, which is of somewhat more complex construction, has a third root situated transversely to the first, surmounted by a small cusp; and there is a larger secondary narrow cusp in line with and posterior to the main one, forming an important component of the sectorial blade. The lower premolars arc ot the same torin as the upper ones except that /m is simple precisely similar to y)2 and ;)3. The molars are far more complex besides being ot markedly diHerent torms in the upper and lower series. They also ditter somewhat in the two subfamilies. ;»i, by reason ot Its great breadth, is by tar the bulkiest tooth in either jaw. Its cingulum is well- developed and anteriorly torms a small subsidiary cusp. Apart from this, in the Caninae the crown comprises two outer cusps, two much lower inner ones, and an internal heel. Ill- is of similar construction but lesser size. The mandibular molars arc not broad. /!/i, the lower caniassial, consists of a large anterior narrow blade, divided into 2 cusps; posterior to and in line with this is a third much lower external cusp; and there are two similarly small internal cusps, one opposite this last, and one opposite the rear section ot the blade. 1112 is a much smaller tooth, with four cusps; and 1113, the smallest tooth in the mouth, little more than a peg, has two cusps, hi the second subfamily, the Simocyoninae (Lycdon), the molars, both above and below, though of the same general form are r.ither simpler in their cuspidation. Habits. Some of the wild dogs, the foxes, have always been recognized as habitual predators, seeking out and killing their own meals; others, the jackals, have for long been commonly regarded as, next to the hyaenas, the great scavengers, living almost entirely on the remains of the lion's or the cheetah's kill. Recent intensive observation has shown this latter notion to be less true than was thought and that the jackals do, in tact, hunt more on their own account than was supposed. This is dealt withmorefulh' later. However, though all the C.-uiidaeare preponderantly flesh caters they do, never- theless, consume an unexpectedly high proportion ot insects and fallen fruits. Most of them are to a very large extent nocturnal, or at least crepuscular, avoiding, except in necessity, direct exposure to the sun. Daylight shelter tor the purpose ot rest, or more especially for breeding and the earlv protection of the ^■oung, is most commonly found in holes in the groimd, "earths" as they are popularly termed. But sometimes, more particularly in the case of young as yet unmated, and hence solitary, adults, temporary concealment is sought in naturally occurring craiuiics amongst rocks, or even in dense grass opened sufficiently for the purpose by a rotatory movement of the animal before lying down. Earths mav be selt-e.xcavated but arc probably most often basically holes made bv other animals, hares, aardvarks, pangolins or termitc■^. improved and adapted. There arc often two or more exits. The Cauidae all appear to be monogamous, the pairs remaining together tor some tune. They share in feeding and bringing up the family, though in the early stages the male is often kept at some distance bv his mate. Litter size amongst wild dogs seems to range between 2 and as many as ly; and there may be one or two litters a CANIDAE 33 year, die age of the mother possibly being a determining factor in both. No set breeding seasons have been cstabhshed for Africa. The average period of gestation is in the nature of 9 weeks but, though not well-investigated, probably varies a good deal amongst different species and may lie anywhere between 7 and 11 weeks. The young, variously known as "cubs" (foxes) or "pups" (jackals and hunting-dog) are breast fed for 6 to 10 weeks, being gradually weaned to solid food, mostly regurgitated by the parents. An interesting ceremony in this connexion is described later imder the jackals. The gait in the Canidac varies a good deal according to circumstances but follows a common pattern throughout the family. A slow walk is rarely adopted except within a limited range of a few yards extent. Over longer distances the normal means of progression is either, at slow speeds, a four-legged run, sometimes varied by bouncing the hindquarters as a unit; or, at higher speeds, a canter; or, in full piu'suit, a gallop. In this last, the two hindfeet touch the ground in rapid succession, impcllmg the animal onwards and slightly upwards while the forelegs arc stretched forwards until the two pairs of limbs arc fully extended and the whole body is in flight out of any contact with the groiuid. The two forefeet then consecutively touch the ground, and give the animal a second onward thrust while the hindlegs are brought forward until they cross the now backwardly directed forelimbs, which leave the ground, the whole body becoming a second time suspended in flight, but this time with the legs tucked under it, not outstretched as previously. The Canidae have no great powers of climbing; but they can overcome low obstruc- tions in their paths both by leaping and by scrambling over those that offer adequate footholds. The hunting-dog when in full cry intermittently performs leaps to obtain a view of the prc\' which may be hidden by the tall grass. Voices in the African Canidae, so far as they have been recorded, are pretty varied, not only between species but according to circumstances as well. There is, without doubt, an extensive vocabulary of signal notes, for the attraction of a mate, the control of the yomig or the co-ordination of the pack, that has not as yet been investigated or recorded. Nothing truly resembling the familiar bark of the domestic dog seems to be uttered by West African wild species, most of tire sounds being characterised as harsh yaps, reiterated melancholy whoops or long-drawn-out notes. Several factors serve to hold the number of Canidae in check. When yoimg and relatively defenceless they are, unless actively protected by the parents, subject to the same attacks as other small mammals from pythons, eagles, other carnivores or driver ants, hi the adult stage they may be killed by hyaenas, angry lions and the like ; but the risks of destruction they run from these or similar enemies seem to be slight. Their numbers arc kept in control more by diseases, and by loss of efficiency from luider- mincd health or accident which prevent them from maintaining their place in the pack or against stronger, more active and thrusting members of their own kind. The availability of food has a considerable influence on the numerical size of the litter or the proportion of it that can be successfully reared. The relatively low density of antelope population in West Africa, for example, is directly responsible for the general rarity of the hunting-dog there and the small size of the packs in comparison with East Africa. ? I 1 1 1 1 ( \ I! N I \ c ) iM s o 1 w I s r M in I A Tiixoiioni) . XltliiuiiJlli tin C .inid.ic might .ip|H'ar to lie .1 i.lc,u-tut, casiU icnii;- in/.ihlc gnnip. tlKir t.woiioim has, in tact, inorc c(iiiipK\itv tliaii at first sl-chis hkclv. I his Is a matter tor more speeiahzed works and it is pointless to Jo more tlian glance .It the position here. Simpson (i<)4_s) gives a long siimiiiar\ ot the Kinsiderations and opinions involved and tnrnishes reterences to the very extensive hteratnre on the snbject. The tossil record is unusually ricli and the intorination which it others leads to a diversit\ ot possibilities in tlie matter ot plivlogem- and ol consequent views. One ot the chiet questions at issue is the limits which should be placed upon the fanuh' and the closeness ot its assocuilion with, or degree ot separation troin. other group.s, and in particular the Ursidae, the bears. The latter, though with their heavy build and lum- bering plantigrade gait appareiuK' so dissimilar trom the dogs, are, at least in Simpson's opinion, \-er\' closcK' related and, in point ot tact, a tairK- late otisho(it trom them. Bur this IS ot no great concern to this present work. On ,1 narrower issue, the extent ot, or even the propriet\' ot am , subdivision within the ver\ uiiitorm tamil\ Canidae is a ttirther point 111 some dispute. In practical terms, three subtamihes ot living caiiids are in tact tairlv gcneralK' recognized, all occurring 111 Atrica though oiil\- two in the region with which this work deals. The third, the C")toc\ oninae comprising soIeK Liiocyoii, the b.it-eared tox, is contined to southern Atrica and the eastern side ot the continent as tar north as Ethiopia. The question as tar as West Atrica is concernei.1, theretore, reduces to the validit\' or otherwise ot recognizing two subtamihes, and the distinction which may be drawn between them. No one could mistake Lydioii tor anything but a dog, tjiough it is true that it dithers shghtlv 111 minor matters ot general appearance trom the more typical members ot the tamiK', the |ack.ds, toxes. wolves and so torth. 1 lowever, on the score ot its slightK' simplitied molars and its possession ot onl\- 4 (hgits on the toretoot it is retained herein as representing the subtamilv Simocvoninae. though it is admittedly doubtful whether these and other minor distmcticsus should lo^icalK- be accorded .iin greater than generic significance. KEY TO THE SUBFAMILIES OY CANinAI' (Previous ke\' page 2iS) ( 'o.it with varicoloured blotches; forefoot with oiil\ 4 iligits: .idult skull lengtli about 200 mm; /;/' with little or ]io sign / the Carnivorous, Padtydorniatous and Edeulalc Mammalia in the British Museum : 180. Type species C i7i/fc/i.<: >kiill, li.M. No. 21,2.11.28 : . i6 rill, CARNIVORES OI WEST AFRICA The legs, tdic and liiiid, are palely red on their outer aspect; and the forelegs carry a longitudinal black streak to the wrist, sometimes clear, sometimes very much reduced. Skull (tig. 4). In all West African skulls the profde show s a marked descent from the trontals to the nasals as opposed to the relatively flat outline at adiistiis. The zygomatic arch upcurves strongly, forming in some cases an almost semicircular outline. The sagittal crest is low over most of the braincase but becomes sharp and keel-like poster- iorly and joins equally pronounced lambdoidal crests to form an acute pyramidal pronnnence. The rostrum is shorter, less tapering and slender tlian in adtistus; the nasals are shorter. The mandible is more curved and rather more powerfully built than m aJiistiii, the depth of the ramus being greater. The coronoid process, too, is broader, generally with an incurved hind-margin and a slight backward hook at the top, the front margin descending in a broad convex curve from this hook to the base. In iuhisiiis there is no hook, and both margins are almost perfectly straight. The chief feature of the dentition is the relatively greater length of the carnassials. It will be seen from the table of measurements, page 55, that both /)■* and iiii are longer than they are ni luhisliis; and iu addition to this the length from front to back of p* is at least 82 per cent of the length oi ui^ + iifi. and that 0(1111 is well over 130 per cent of /;i2 4- 1113. Habits. Close studies of these animals in the field have been made in recent years by Wyinan (1967) and Goodall (1970), and much of the following accoimt is derived from these sources. Golden jackals arc to be seen on the move b)' both day and night, especially it the latter is brightly moonlit. Like most dogs they can sleep or be active at a moment's notice as occasion demands; and though t\\c\ do most of their feeding by day it is sometimes necessary to follow up kills made at night by lions or hyaenas even though they may probably, apart from a few hastily snatched mouthfuls, have to wait at a safe distance till well after siuirise before feeding can commence in earnest. Often golden jackals are solitary animals; but at mating times and during the raising of a family they are always in pairs; and sometimes these or small family units hang together for extended periods. A. J. Hopson noted that they were frequentl\- to be seen up to tour in a party in the Sahcl woodland and Sulradora pcrsiai (Salt Bush) thickets near Lake Chad (private communication). Their popular reputation is solely that of scavengers, cleaning up, in company of the vultures, when other larger carnivores have killed and eaten their fill. That thc\- do this is true; but it is only part of the story. It is possibly their easiest way of procuring a full meal since it calls tor little more than patience or the skill to see an opening and the agility to dash m, seize a few mouthfuls and nip quickly awav before the heavier and slower-moving feeders can prevent them. But, in fact, their dietary is much wider than this, and they do a good deal of killing n their own accoiuit. They mostly confine themselves to small prey, poiuicing upon hares, rats, ground squirrels, cutting-grass (Tliryoiioinys) and the like; they arc known to take lizards and not infrequently to kill and eat snakes. Ground-haimting birds such as francohns and bustards tall prey to them. They also consume a surprisingly large amount of insects: dung beetles, larvae, termites, or grasshoppers, the last ot which thev may either pounce <->n or catch in flight. The\' also eat a good deal of ctralimita! data in conjimction with Schwarz"s (1915) figures published for cciilnilis — which themselves were derived from a dry skin. From these it would seem that there is not a great deal of dirterence in size except that the ear is possibly shorter in ndiistiis. One animal from liahr-el-Ghazal was said to weigh yy kg, that is slightly more than iiiirciis. Skull (tig. 6). The first obvious ditterence between the skull ot --^-s-^-^ , , 'J CJJ— Q 'J _ ti W S != -^ = 56 I HE CARNIVOlilS Ol WIST AFKICA Gciius FENNECUS Dcsmarcst. TS04 Feiuiccs reiinccus Dcsm.irest, A. G., 1S04, l\'oui'i:im Diciionnaire d' Hisloire Niitiin-llc, 24, Tableau niL-tliodiqiiL- dcs Mamniifercs: iS. Type species FiHiiecus arabicus Desniarest(= Ctinis zcrda Ziniincniiann). The name IS a Latinized form of the Moorish word tor a fox, fainec. Mcgtilolis IlHgcr, 181 1, Proilroniiis systciiwlii Mammalium ct Avium . . . : 131. Type species Caiiii ccrdo Gmehn (= Caiiis zerda Zimmermann). This name was made up from the Greek words tncj^as (mc^al-) large and oiis (otoi) ear. Tills is a monospecific genus distributed over a small area of northern Africa from Morocco to Egypt, as far south only as Air and Sudan, and thence across to parts of Arabia. In other words, it is an animal of dry sandy deserts or subdcscrts. Since there is only one species there is no point in entering into a generic description. FENNECUS ZERDA (Zimmermann) Feniicc I'lilpcs minimus siiannsis Skjoldebrand, 1777, K. svciiska I'etenskAkad. Haudl. 38: 267, pi. 6. Algerian Sahara. This name is regarded as invalid because, as given to a species, not a subspecies, it was tri- nomial. The second name, minimus, is Latin for smallest, given because ot this annnal's dnninutive size for a fox; the third name is a Latinizarion of Sahara. Ciinis zcrda Zimniermami, 1780, Gcographische Gcschichle dcs Akiisclicn und da vicrluszi\;i-n Tltiere 2: 247-24S. Sahara and North Africa behind the Atlas Mountains. The name was said by Zimmermann to be that used in "Barbary". Cdnis ardo Gmelin, 1788, Linnaeus' Syslvnui Natumc, 13th ed. 1 : 75. Sahara. This name is another spelling of zi-rdii. I'ii'ina anrita Meyer, 1793, Systeniatisch-summarische Uebersiclit dcr ncuesteii zoohgischcn Entdeckungcn in Naiholland und AJrika: 91. Bisk.a etc., Algeria. The Latin adjective aurila means having large ears. Fcnncais arabicus Desmarest, A. G., 1804, Nouvcau Diaionnaiic d'Hisioire i\'atiin-Uc, 2i, Tableau metho- dique dcs Mammiferes: 18. Barbary, Nubia, Abyssinia. Mt'^ahiis ccrda Illiger, 1811, Prodromus systcmatis Maiiimaliuni ti Avunn . . . ; 131. The name is a variant oi zcrda. h'cnuccus brucci Desmarest, A. G., 1820, Encyclopedic Mcthcdique, Mammalogie: 235. Libya, Tunis, Algeria, Sena.ar. James Bruce, after whom this was called, was a well-known explorer of northern Africa in the second half of the 1 8th century. GjMij^eimcaK Lesson, 1827, Manuel dc Maninialoi;ic : 168. Vnlpcs denhami Boitard, 1842, Le Jardin dcs Planlcs: 213. Interior of Africa. This was named in honour of Lt. Col. Dixon Denham, a famous traveller m the Siliara and explorer of Lake Chad in the early iQth century. Distribution and general. The range of this essentially Saharan animal has already been given above. In suitable localities it is not uncommon and because oi its small size and rather charming appearance it has tor long been a favourite with writers on natural history, tiguruig far more copiouslv in literature than mam- more widely FENNECUS 57 distributed and rather more important animals. In general appearance it is a mimatiuc fox with a sandy coat, huge ears and a very bushy tail, and it is, indeed, often know^l as the fennec fox, though any really close relationship to the true foxes has been brought into question. It does not occur ever)'where iji the Desert and Subdesert zones bccaure it must have soft sand into which to burrow. Sand dmies are therefore ideal, but not in utterly barren situations since food must, of course, be available in fair quantities. Description. The femiec (Plate i) has a head & body length of about 300 to 370 nun and a tail of 160 to 240 mm. It stands about 150 to 175 mm at the shoulder and weighs about 2 kg. The pelage is long and very soft, both above and below. Broadly speaking, dorsally it is sandy, but there is a certain amount of variation, some specimens beiug rather greyer, some rather redder. The same apphes to a darker, richer coloured, band along the spine, almost absent from some, clear m others, especially on the hinder part of the back. In this area, in some skins, the bristles are dark-tipped with a pure white subterminal band, forming a very prettily-patterned patch. In others the pattern is more diffuse, the majority of specimens having fme bristles with long black tips thinly dispersed over the entire dorsal region. The pelage is so soft to the touch because it consists, apart from these scattered bristles, entirely of dense, very fme, very long, luiderfur. Tliis, iji the North African examples, is pale chocolate-grey in the basal half the extreme base being narrowly white; but in the western specimen from A'ir there is no trace of this basal tijiting; while in the Dongola (Sudan) skin it is relatively pale. The West African examples, too, are much shorter-fiu-red; but they are all youngish animals. The tuidcrparts aiid the insides of the limbs are pure white, the frir being abiuidant and soft, but only half the length of that of the back. The tail, which is roughly half as long as the head & body, is very bushy, the very long hairs with which it is clothed beiiag towards the tips a rather redder brown than the back. There is a deep blackish-browni mark not far from the root of the tail covering a scent gland; and the extreme tip is also blackish-brown. The head, with a broad face and large eyes, comes rather abruptly to a narrow muzzle and is wholly dominated by the enormous, pointed ears which are broad as well as long. On their backs they are sandy-grey, but all around the marginal area on the inside there are dense long white hairs. The crown and front are sandy, but much of the rest of the face, surroimding the eyes, the rhinarimn, the cheeks, and the upper lips, is pmc white with the exception of two dark, reddish-brown lines descending from the inner comers of the eys to the lips. The upper parts of the limbs are, in the northern African specimens, reddish-sandy; but in the Air examples they are nearly white. Skull (fig. 7). The skull tapers fairly sharply from a moderately broad braincase and enormous bullae to a very narrow rostrum. The profde dips appreciably just forward of the orbits to give this narrow and low muzzle. The braincase is rounded; the supraorbital ridges arc well-pronoiuiced but narrow and sharp, hollowed on their upper surfaces, in the maiuier of Vulpes rather than Cants. There is no very marked intcrorbital constriction. The distance across the zygomata is wide, the maxillary process broad, the slender arches sharply up-curved, the circumorbital ring widely 58 THE PAnNIVORES OF WEST AfRICA open. There is little or no sagittal crest and only poorlv developed supraoccipital ones. The bullae are extremely large. The palate is very broad betweeji the carnassials but narrows abruptly and becomes practically parallel-sided anteriorly. Its hind margm is about level with the middle of (/i-. There is nothing particularly remarkable about the lower jaw or the dentition apart from its very sharply cuspidate nature whicii probably facilitates an insectivorous diet (fig. i). Taxonomy. This, despite the multitude of names, actual and in permutation, by which this animal has been knowji to science, is pretty straightforward. It is true that in the early days Butfon (177O, Suppl. 3: 148) referred to it as the anonymous annual, and that Lesson (1S27) wrote ot it (in translation) that perhaps no animal had more engaged naturalists than this; they have made of it by turns a dog, a galago, or the type of the genus Fciiucc. The old coiifusion of naming has now been s\\ ept aside and ti.xed \\ ith apparent permanency as Fciiiicais zcrdd. But, at a very considerably narrower pitch than that indicated in the previous paragraph, some doubt of the animal's precise relationship still remains. The tennec has long and widely been regarded as a kind of fox, and is in fact very frequently referred to as the feiuicc fox; but the cytological researches of Matthey (1954) have shown tliat the chromosomes (diploid number, 2N — 64) indicate that the genus lies closer to the wolves {Caiiis) than to the foxes (Viilpes). So far, no races have been described. The West African material from Air is poor 111 quality and meagre in amount and it is therefore not possible to draw any reasonable conclusions; but superficially the animals from this area seem somewhat paler and shorter haired than those from further north-east; but there is no significant diilerencc of size so far as can be deduced from the mean measurements of three h'om the one area and of five from the other. Habits. Accoimts of the.sc in the wild have been briefl^' given h\ several collectors and observers over a large number of years; but, in all, the information from these sources amoiuits to little beyond the most obvious facts of life in the desert. However, because of the fascination which this miniature "fox"' of charming appearance has long exercised over human-beings, especially as a household pet, the feniiec has been more closely studied in captivity than most carnivores. Good descriptions of its behav- iour and disposition under these circumstances have been given in recent years by a number of writers of whom the following arc the chief: Rcnsch (1950), l^etter (1957), Volf (1957), Hill (1961), Saint Girons (1962), Vogel (1962) and Gauthier-Pilters (1966). Though these accounts concern animals in unnatiu-al conditions thc\- nevertheless cast important light on instinctive behaviours, the more convincing in that many traits are displayed in common b)' animals of different origins at different times and places. It is not possible in a work of the present nature to do more than glance briefly at some of the recorded facts, more especiallv those which most probablv rc\eal the femtec's normal activities. Fermecs are essentially nocturnal or crepuscular in their activit)-; \ct though, like so many desert animals, they avoid the full hirce of the mid-day heat, they are not averse to sunning themselves for brief periods before the dav is far advanced. 15ut FENNECUS 59 Fig. 7. Faiiicais zada: skull. B.M. No. 1939.1746, ?, x i 60 THE CARNIVORES OF WEST AFRICA very little of this insolation suffices for their daily needs and they soon seek shade — though they ma)' emerge for a second spell, a pattern that may be observed in many domestic dogs. The fennec's home is a hole in the groiuid, which it digs for itself, fairly deep probably to avoid overheating during the day. These "earths" are lined with soft material, and they have a number of different escape passages some of which are said occasionall)' to commmiicate with the homes of other fennecs, as these are sociable little animals which in suitable areas live amicably together in some numbers. The unit of these communities is almost certainlv a small family group; and it would seem from indications, though it is not certainly known, that male and female hang closely together for long periods. Favourable localities for femiccs are characterized by soft sand into which burrowing can be carried out with ease. For this reason stable sand-dtuies fixed by the roots of sparse vegetation are ideal, wholly barren desert having no attraction since food must be available in fair quantities. Given a soft soil such as exists in these sites, digging, carried out with the forepaws, is very rapid. All who have kept these animals refer to one habit that makes them difficult to tolerate as domestic pets: the constant and very noisy nocturnal scratching resulting from attempts to dig into unyielding wooden or concrete floors. That the instinct to hide is very strongly developed is illustrated by the fact that even in captivity fennecs always seek to shelter underneath furniture; or, if nothing more resembling an "earth" is available, by pulling over themselves such bedding as they may be provided with. It has already been said that fennecs dislike intense heat, particularly direct insolation; but they very much appreciate warmth. They are, indeed, highly sensitive to cold, and this is another reason for the depth of the burrow so that a wholly equable tem- perature can be maintained. When above ground on the hunt for food, protection against the night cold that develops rapidly after sunset in sandy deserts is to some extent provided by the dense woolly pelage; but, to judge from behaviour in captivity, fennecs are intermittently active for short periods and this might be explained by the necessity, in nature, to return from time to time to the warmth of the home. Another highly important reason for the avoidance of intense insolation and the frequent seeking of shelter in a deep burrow, in which the humidity probably remains at a high level, is the need to conserve water. It is imlikely that the majority of feiuiecs in the wild have easy access to standing water in order to drink, and very likely they obtain most of what they need from moisture in fruits and bulbous roots. Observations on drinking habits in captivity are curiously conflicting. Rensch found that his fennec drank little or nothing for the first two or three years except at the mating season; but Later in life it drank daily. Vogel found much the same thing; but he noticed, in addition, that the animal was averse to drinking out of a bowl but readily lapped up drops spilled on the floor — so much so that it was observed, itself, to scatter water from the drinking vessel with its forepaws or nose. Rensch mentions his fennec as licking drops from a tap. This would seem to indicate that even if these animals had access to oases, or other pools, they would prefer to get what extra supplies they required beyond what is provided in fruit by licking small quantities troin leaves or other surfaces moistened by dew or rare showers of rain. Schmidt-Nielsen (1964), in connexion with loss of bodily water, foimd that urine could attain a high concentra- FENNECUS 6l tion in the fennecs. When over-heated, fennecs can cool off to some extent, hke dogs, by panting; but as this involves loss of essential moisture from the tongue it is not an operation that can commonly be engaged in. Fennecs are seemingly almost omnivorous, and there is little difficulty in feeding them in captivity since they readily eat most things from roast beef to marmalade. Fn their natural surroimdings they live on small rodents, such as gerbils and jerboas, lizards, insects, eggs, small birds and a good deal of vegetable matter, of which fruits and tuberous or bulbous roots form the main part. Vogel observed his fennec, like a domestic dog, to eat grass when opportunity offered. According to Professor Monod of Dakar quoted in Dekeyser (1955), fennecs are very fond of the bright yellow, leafless parasite Cistanche phelypaea Cout. often growing in fleshy clusters at ground level on the roots of, amongst others, the so-called salt bush, Salvaclom persica, commonly occurring in the Sahel and Subdesert zones. Big pieces of food are consumed in a sitting posture; small ones are eaten while standing. In many of their ways these little "desert foxes" reflect the manners of some domestic dogs: as, for example, in their inquisitiveness, evinced in sniffing at or quizzically regarding unfamiliar objects; in hiding away surplus tit-bits of food; in turning round three or four times before settling down. They particularly resemble poodles in their ability to stand and walk upright on their hindlegs, and in the way they stretch their hindlimbs after sleep, or lie flat on their bellies with both fore and hind legs stretched out. In other ways they are rather cat-like, particularly in their manner of cleaning themselves by licking their forepaws and "washing" their heads. The large ears are cleaned on their insides by the hindfeet. Like cats, too, fennecs are very competent jumpers, being able to accomplish a standing more or less vertical spring of 600 mm or more and a horizontal spring of 1200 mm. They also, at times, can purr something like a cat; and when they sit on their haunches they curve their tails sideways and forwards like a cat, sometimes even raising the tip off the ground as cats do. Fennecs, too, are very competent climbers and can ascend vertical obstructions that offer some sort of foothold. They can also squeeze through remarkably narrow crevices. These animals scratch a shallow hole to defaecate and urinate in and they cover their droppings with sand, sometimes shovelling it over with the nose or, more often, flinging the earth backwards with a scraping action of the hindlegs, just as domestic dogs can often be seen to do. This latter ritual is performed automatically even when it is pointless on the hard wooden floor of a cage. Feimecs utter a variety of sounds from a low growl to a higher snarl ; when content and being stroked or fondled the\- make a purring noise similar to that of a cat. They are given to yapping at night. There is an annual moult. Opinions differ as to whether these animals are ever completely tameable. As with almost any wild creature it is probably merely a matter of inbuilt temperament, varying from one individual to another. One fennec, at least, has shown itself to have a very long memory and to be able to recognize its owner with excited pleasure after many months of separation — and tliis at a distance by sight, the possibilit)- of smell being eliminated by a glass front to the cage. These animals have a longevity- of about 12 years. Schmidt-Nielsen records that the young are dug out of their nests, fattened up and marketed for eating. 62 1111 (AUMVOUIS (II W IS 1 AIUKA MatiiisT .md breeding in captivity have been observed and recorded b\ I'etter, Volt. and Saint Girons. After penetration the male turns round so that the two sexes face m opposite directions, and coition lasts i{- hours. There is apparently no attempt at Mibsequem copulation. The period ot gestation is so or si days. Petter found tliat after mating the female continued to act normally but the male grew aggressive towards people whom lie knew well; this became more pronoimccd after parturition and so he was separated from the niotjicr. By then she was so upset that she kept transportnig her newlv born cub from place to place and ended by thus accidentally causing its death. Volf records verv much the same sort of thing. From i to 3 cubs are boni. Their eyes are closed, their ears free though giving no hint of their future huge size. The pelage is fuie and short. Under natural conditions the -soung appear aKi,ays to be born in March or. at the latest, the begiiuiing of April. Gauthier-Pilters (1966) has recorded in sonic detail her observations on play in the tennecs made on 21 animals over a period of 11 years, all born in the wild. Juvenile fennccs play with each inlicr verv like fox cubs or the puppies of domestic dogs, biting each other in the legs and neck and rolling each other over. They also shake small objects as if shaking a rat to death. From the age of about 6 weeks they indulge ill racing plav, chasing each otlier. the pursuer with cars erect and directed forw^ards, the pursued with them laid back, zigzagging or making abrupt changes of direction. Tliis thev will do even, if necessarv, in a very confuted space. Where opportunitv offers the\- may plav a kind of hide-and-seek. Should play become too rough and one of the participants get cornered, or for that matter at an\- time or any age when offensive or defensive postme seems necessar\-, the back becomes arched and the ears are laid back in a threatening attitude. Feiuiecs have no abilitv to raise their hackles like a dog, but instead, the black mark at the root of the tail is erected and displayed. Solitary plav also takes place, using articles ot food or substitute to\-s, tossing these in the air or shoving them about with foot or nose. Chasing tlieir own tails is sometimes indulged in. Fennccs will play even w-ith strange anini.ils which thev have only just met. Gauthier- l^ilters (1966) records games between \'oung teiuiecs, dogs and jackals; but cats appear unwelcome and frightening. Should i-itlier creatures fiil thev pla\' willingK- with human beings. Inclination to pla\- appears to vary with se.isiiii. taljuig to a muiimum at breeding time. This observer found tlie pattern ot female beliaviour with the male to differ before and after parturition, exhibiting in the first place elements of greeting behaviour and in the second of breeding behaviour, that is tlie bringing and offering of food — though the male was, in the event, never allowed to take it. Most pla\' takes pLace at evening or .it night, hideed. witli captive tennecv. to judge from the experiences recorded b\- Ci.aithier-Pilters. this can be somewhat ot a tri.il tor the owners ot these luH turnalK- ver\- active creatures, sijice they persistently play after dark for hours on end. preferabK' over and luider beds, tables and chairs. This author records a T4-day trek with half-tamed tennecs w. hich for safety had to be tied at night to her camp-bed and which, despite their thus restricted freedom, spent their time in romping or, at interv.iK. digging deep holes in the sand. Sueli pla\' activities, espeeialK' 111 the voung, are recogni/abl\ related 10 beluiMour VULPF.S 63 cxliibitcd 111 various situations regularly arising in later life, and for which they arc, in effect, rehearsals, training and strengthening the necessary muscles and inculcating instinctive reaction on occasions demanding offence, defence, capture or kilhng. Table 2: Numerical data for i'enncais zcriia West West Sudan A&ica Africa (Dongola) Northern Northern (Afr) (Air) single Africa Africa means extremes specimen means extremes Vcgetttion Subdescit — Subdesert Subdesert — Number in mean 3 — I 5 — Condylobasal length 83-2 82-8-83 -7 82-3 83-8 78-7-86-7 Basilar length 77.9 77-4-78 -2 76-5 78-2 73-6-80-4 Palatilar length 39-0 38-9-39-I 37-3 38-9 36-4-41-0 Zygomatic breadth 45-4 44-4-47-2 43-8 46-1 43-5-48-6 Upper cheekteeth breadth 24-2 23-3-2S-4 24-0 25-0 23-2-25-9 Nasals, length 27-5 26-2-28 ■ S 26-8 27-1 26-1-28-7 Interorbital breadth 16-5 16-0-17-0 15-8 16-2 15-5-17-0 Postorbital constriction 20-3 19-8-21-; 17-7 20-2 18-7-21 -3 Braincase breadth 3fi-8 36-7-37-0 35-7 36-5 35-5-37-5 Toothrow (c — III-) 3S-I 33-6-36-0 35-3 35-4 32-4-36-4 p* length 7-4 7-2-7-6 7-0 7-6 7-4-7-8 111^ + II fi length lO'I 9-9-10-2 9-4 lO-I 9-5-10-5 Mil length 8-3 8-1-8-4 8-1 8-3 8-0-8-5 1112 ■- 1113 length 8-0 (7-9)8-0 y-i 7-6 7-2-7-8 Head & bodv 355 350-360 360 371 333-395 Tail 170 160-180 190 147 125-187 Hindfoot 93 92-94 92 94 90-9S Ear 90 86-95 90 94 90-97 RATIOS (per cent) Tail/head & bod\- 48 S3 40 Zygom. br./condylob. 1. 55 53 55 Braincase/condylob. 1. 44 43 44 Braincase/zygom. br. 81 81 79 Palatilar l./condylob. 1, 47 45 46 Interorb./postorb. 81 89 80 P*JC— III 21-0 19-8 21-4 p^jm^ — II fi 73-2 74-6 75-2 iiilliii, - (113 104. 114 109 c;cnus VULPES Hemiiig. 1N22 True Foxes I iilpcs Frisch, 1775, Das Naliirsyslcm dcr viir(ussigai Tliicre, 15. This work has been ruled to be unavailable I'y the International ("ommission on Zoological Nomenclature (Opinion No. ^sS of 1954). The name IS the Latin for a tox. 64 THE C.ARMVOHES OF WEST AERICA I'lilpcs Okcii, iSif), Lihibihli da Kalnr^cschiihk\ 3, 2: 1033, 1034. Type species Vtilpcs aviiiiiunis Okcii (= Cam's viilpcs Linnaeus). Okeii's Lilirhiuli is similarly unavailable (Opinion No. 417 of iy56). I ■"//)« Fleming, 1822, The Philosophy of Zoology, 2: 1S4. Type species Catiis rulpcs Linnaeus. (^yimlopcx H. Smith, 1839, Jardinc's Wmiralist's Library, 25 (of the series), 9 (of Mamm.ils): 222. Type species Caiiis corsac Linnaeus. Distribution. This genus is spread tluoughout Europe, much of Asia and of North America, and over the more arid areas of Africa. The common red fo.x of Europe {I'lilpcs viilpes), in one form or another, occurs over a great part of tliis wide range with the exception of the more southerly parts of Asia and Africa south of the Medi- terrajiean region — though its identity with the American red fox, frequently alter- natively designated V. fnlva, is disputed. Rather less than a dozen species in all are recognized, of which three occur in Africa south of the Sahara, two of them being found withui our limits. Foxes are small to moderate-sized carnivores, having a long, soft, dense, often ver\- attractive coat and an outstandingly bushy tail — termed in hunting circles the "brush". This profusely haired tail in conjunction with a sharp face and prominent ears makes it difficult to differentiate foxes precisely from some of the typical (wild) dogs; and, indeed, this genus has often been synonymized with Canis. However, in general, ill Africa, foxes are smaller and have shorter legs — they are much more able than dogs to slink along with their bellies almost in contact with the ground. Nevertheless, despite shortness of limb they can move verv fast over long distances. A more positive character dividing the foxes from the t\'pical dogs is that the tail in the tormcr measures more than half the length of head & body, whereas in the latter the reverse is the case. In foxes the pupil of the eye is a vertical oval, rather in the style of a cat but with not so wide a range of expansion and contraction. Some, if not all, foxes have .1 scent gland on the top side of the tail about 50 to 60 mm beyond the root; this organ exists, according to Anderson (1902), in at least one of our West African species. There are five clawed digits on the forefoot but only four on the hindfoot; the soles of the feet in the African species are abimdautiv hairy between the pads, riieppclli vet more so than piillidd. Skull (fig. n). No general description ot the I'lilpa skull is given here. Its form bears a close overall resemblance to that of Cniiis, differing, apart from a considerable disparity of size, in one minor point. This concerns the supraorbital ridges which in Cauls are dorsally smootlily convex, whereas in Viilpcs there is a slight concavity in the upper surface. It is this lack of any important cranial or dental disagreement that has made taxonomists sometimes doubtful of the validity of generic distinction between 1 'iilpcs and Civiis. Habits. Little enough is known of the lives ot the majority ot species apart from the fact that the palaearctic common red fox has been intensively observed and to some degree studied over a long period of time. This species is popularly regarded as exhibit- ing in its appearance and behaviour the acme of slyness and cunning, and its name has passed proverbially into European languages as the most apt and succinct expression cf thc^e attributes. It has, indisputably, other more admirable qualities: courage, determination, endurance. It is the combination ot all these that has reiideicd this fox VULPES 65 possibly the favourite and most exciting animal of the chase. And it is these charac- teristics, too, that have in large measure enabled these harmful predators to survive in considerable numbers in the face of human expansion and of intensive organized hunting over many centuries, when wolves, bears and other carnivores antagonistic to man's interests have tended towards extermination. In what degree, if at all, this craftiness and skill in self-preservation are shared by African species is unrecorded. But it is perhaps w'orth noting, from an historical point of view, that a Hunt in semi- English style existed some years ago at Zaria (Nigeria), its activities being directed against the sand fox inhabiting the neighbouring fields and hillsides in fair numbers. There is good reason to suppose that proximity to man so far from being distasteful is welcomed by foxes since any occasional danger arising from it is more than com- pensated by the ready availability for food of domestic animals and birds, more con- centrated in amoimt than wild prey and less apt at defending themselves. Most foxes arc in large part nocturnal, but they are also commonly active during daylight hours, especially in the early mormng or late evening. So far as is known, all species of Vidpcs shelter and breed m self-constructed "earths" in the ground or, much more rarely, in the protection of holes formed by rocks. Some live only in small family units ; others are more gregarious. Beyond these generalities it is difficult to go ; for nothing appears to have been recorded for African species regarding the various aspects of breeding, and little about feeding habits. Concerning these latter, it may be safely assumed that African foxes take a wide diet, not only of flesh but of fruits and other vegetable matter as well. Their enemies are the more powerful carnivores, birds of prey, the larger snakes — and, of course, diseases. These last, and their possible impact upon man, have not been investigated for tropical African foxes. Taxonomy. It has already been mentioned that opinion has differed in the past, from Luinaeus onwards, as to whether the foxes constitute a separate genus or should be regarded as wholly one with Caiiis; but the two are now generally looked upon as validly separable. There are no problems with regard to African species, though there has been at times a little confusion of thought — vide Thomas (1918), who cleared up points relating to nicppdli and pallida but fell into the trap of supposing that Gray's I'ldpcs dorsalis from Senegal was indeed a fox, whereas, as pointed out by Ellerman & Morrison-Scott (195 1), it was in fact a jackal. The two species that occur within our limits may be separated thus: KEY TO THE WEST AFRICAN SPECIES OF VULPES (previous key page 3 5) Tail over 300 mm, tip white; dorsal pelage with a bright reddish spinal band flanked with greyish; cars over 80 mm long; breadth across the outside ot the upper cheekteeth over 30 mm; c — • 11 fi over 45 mm rueppelli {pa^c 66) Tail under 300 mm, tip black; dorsal pelage darker medially but not bright red or flanked with grey; ears under 80 mm long; breadth across the upper cheek- teeth under 30 mm; c — • 01- under 45 mm . . . pallida {pa^c 70) 66 I 1 1 I < A 11 M \ ( Ml I s 1 1 1 W I s 1 M in I A VULFES RUEPPELLI (Schm?) Kiipix-ll's Fox (.',iM/s nippclii [sn] .Scliin/. 1^:15, C'liriV; '.v 'I'liiirnitli . . . Am ilciii iV(iH^i).(i.(i//i'/i /rq' ubcncl:! . . . , 4: S08. Ootigola. tdiiard Riippi-ll, attcr whom this was named, explored and collected in various parrs ol norrli-eastcrn Africa tor the Senckenberg Nature-research Societv, Frankfort-on-Main. I'lViif fntitiliatf Cretzschmar, iSzy, in Riippell's Atlns ch i/cr Rcise iiii ihn