THE CARNIVORES
OF
WEST AFRICA
BY
D. R. ROSEVEAR
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BRITISH MUSEUM (NATURAL HISTORY)
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THE CARNIVORES
OF
WEST AFRICA
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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-\iv>. Northern part ot the Kete Kr.uhi
district, Togo, probably near Bimbila. between the Oti and Oak.i Uivcrs. This was called after Us
collector. Professor Mischlich, a German government oHicial.
Lycaon chcrnmicri Matschie, 1915, Shcr. Cos. naturf. I'rcnmic, Boil.: 369-371. Lake Chad area, probably
near Dikwa. This was named in honour ot the German Governor ot the Kamerun, Karl Ebermaier,
who sent the living animal to the Berlin Zoo.
There is a further extensive 20th century synonymy, mostly erected b) M.itschie and mostly applying
to southern .and eastern Africa.
Distribution and general. The luniting dog, somctmus too rcstnctcdk, ,ind
ccrtainlv today inislLMdiiiglv, called the Cape luiiuiiig dog, is eoiitmed to Atrica hut
has a very wide rajige throughout much ot the coiuiiieiit trom the Sahara to South
Africa. In the former of these it is generally regarded as prohably little more than an
occasional rare visitor; hut Heim de Balsac (1936) thinks there is good evidence in
support of a distinct Saharan race. Two specimens \\ere captured north ot In Ouzzal
(Adrar des Itoras) in 1927 (Lhote 1946); and it is said to occur m Tanezroutt. Tihesti
and Enncdi.
South ot the Sahara it is an .uninal ot the more open kinds ot grass-woodland but is
erratic in its occurrence, the more so todav as its numlicrs are, in general, becoming
ever less and less owing to the widening settlement and use ot the land by man and
the consequent reduction ot available food in the torm ot antelopes. A century ago it
might at times be seen to hunt hi packs ot up to a hundred, or possibly even more, and
those of thirty to forty were common. Now, and especially in West Africa, the packs
tend to be smaller, sometimes only six or eigin, and often less; but R. H. Kemp,
of the Nigerian Federal Forest Kescarch Department, saw a pack of at least 30 in
April 1969 in Borgu (extreme western Nigeria), halt ot them pups estim.ited to be
five or six months old (personal communication); and \Mth general g.ime protiction
in this Reserve it seems possible that it is there becoming again more numerous.
There arc records of occurrence from the north ot the lvor\- Coast, the upper Volta
area, and across this belt to Sudan. Eritrea and Somali.i. Thence the species ranges
LYCAON
77
^^
78 "I HE CARNIVORI-S OI WEST A H) H. A
down tlic eastern halt of the continent south to Transvaal and Portugese East Africa
and across to Angohi and Soutli-west Africa. South of tlie Tropic of Capricorn, where
it tornierly d\\ eh in large numbers, it is now absent or only a very occasional wanderer.
In the British Museum there exist trom West Africa only one complete specimen,
that trom Mani (Lower Shari River, Chad), and two partial specimens, the one an
unmeasured skin trom Leri-u-duchi (Zaria, Nigeria), the other a skull from Lake Chad.
The hunting dog prctcrs open coimtr;,- where vision and pursuit of quarry arc not
much obstructed. It is therefore most commonK' to be foiuid in the Sudan and Sahcl
woodlands where the grass is tairly short and relatively sparse; and though it certainly
occurs in the Guinea woodland this is probably for the most part during the dry-season
when, as a result ot the annual grass burn, this type of country becomes very open.
During the rains, on the other hand, the ground cover in most places in this zone is
both too dense and too high — 2 metres or more — to make hunting in the manner
ot these dogs a practical possibility. This applies in a somewhat lesser degree to the
Ooka zone.
Description. Lyciuvi (tig. 9) is easily the bulkiest and most solidly built of West
African canids, standing 61 to 66 cm at the shoulder, with a head & body length ot
about 1 14 cm and a bushy tail of 35-5 to 3S cm. The weight of a tully grown animal
would be some 27 to 32 kg; but the average in a piack, is probably more in the nature
ot 20 kg. The Himting Dog is quite immistakeable with its short coat, blotched with
black, yellow and (often) white, and its very conspicuouslv large and rounded ears.
Though quite obviously a dog, it is, by the teaturcs just enumerated, very readily
distinguisliable from the jackals, ^vhich are much smaller and have pointed cars.
The pelage is harsh, of moderate length, close-lying and not at all dense; and it is,
thus, vastly ditferent from that typical ot the jackals and foxes. It is, indeed, of entirely
different composition since it consists solely of stiff, terete, bristle-hairs and has no
underfur whatsoever. These bristles are, by comparison with the closely packed fur
ot the foxes, relatively widely spaced at their roots and emerge from the hair follicles
mostly in pairs, occasionally in tltrees, rarely singly. Instead of the colour ajinulation
common in the pelage of other canids these bristles are to all intents and purposes
imicolorous trom root to tip, pure white, blackish or pale yellowish-brown. These
tints occur over sharply dcfuied areas of the skin and result in a broad mottling of the
pelage with little ijitergraciing of one patch with the next, the fur thus exhibiting
nothing of the intimate "pepper and salt" mixture common to so many mammalian
coats. The proportion of one colour to the others varies very considerably, as do their
relative positions on the body. Matschic (19T5) tried to establish fixed patterns in
relation to various geograpihical areas, the basis of his proposed specific naming. Not
only were the data on which this sup>position was foiuided mostly exiguous, but also,
where in the British Museum collection more than one specimen exists from a given
locality, constancy of pattern is seen to be non-existent. This is amply confirmed by
photographs of packs taken bv R. H. Kemp in the Borgu Game Reserve, Northern
Nigeria, (Howell. 196SI, and private comiiuinication) ; and also by those illustrating
Kiihme (1965b).
The chest and belK- arc motllcd 111 the m. inner ot the b.ick but the hair is considerably
LYCAON 79
shorter and yet more scanty, scarcely obscuring the skin. In the only two West African
specimens available for study the chin and under the jaw are black; the throat and
upper part of the chest white. The back of the neck and the top of the head to the
forward level of the eyes are pale brown; and a broadening black stripe riuis medially
through this region over the crown and down the centre ot the face to join the all-
black muzzle. The posterior part of this line is sometimes faint or lacking. The muzzle
itself is short, broad and heavy; the nostrils are well-separated and widely open. The
large, rounded, cars, about 115 to 125 mm long, are black on their backs and to some
extent on their anterior faces as well, though here there is also a white tuft situated
over the lower part of the inner edge.
The legs are variegated with the same colours as the back. In this subfamily there
are only four toes on each foot, the short claws being deep and powerful. The feet
themselves are strongly built. The two middle toes are widely separated but joijied by a
narrow naked web (Pocock, 1914b). The proximal margins of the digital pads carry
long fringes of stiff reddish hairs; but the rest of the sole between these and the plantar
pad is scantily clad. The distance between the plantar and carpal pad is luiusually long
and narrow. The hindfect are similar but narrower. The tail is short-haired in its basal
third, and then has a longish-haired black and white bush, the terminal quarter being
conspicuously piure white. The females usually have about 1 2 teats.
Skull (figs 10 and u). The skull is very powerfully built, with strong ridges and
arches and a massive dentition. As canid skulls in general go it is short and broad. The
braincase, as in other members of the famih', seems unexpectedly small ; it is romided
but carries a pronomiced, sharp sagittal crest which joins an equally prominent supra-
occipital crest to form a backwardly-projecting pyramidal "helmet". The interorbital
region is slightly constricted, the pointed, triangular, do\\Tiwardly curving postorbital
processes being wcU-dcveloped, their hind margins continuing and converging post-
eriorly across the temporal region in a diamond outline to meet the forward end of the
sagittal crest. The inuzzle is noticeably short, wide and deep, the anterior nares very
open.
The zygomatic arches are broad and strong, sharply upcurved, and since the pointed
process on the upper margin of the jugal is not very distantly separated from the post-
orbital process the orbit goes some way towards complete encirclement, but not quite
so near as in the hyaenas. The palate is broad tliroughout, being particularly so between
the camassials, and shows little of that marked anterior jiarrowing between the pre-
molars and canines that gives the more typical dogs their sharply pointed muzzles.
Posteriorly the palate comes to an etid about the level of the hind margin o( in~. The
bullae, though prominent, can not, in comparison with the size of the skull and their
relative development in the jackals and foxes, be characterized as large. The mandible
is robustly built, with deep, solid rami and strong coronoid, condvlar and angular
processes.
The dentition is powertul, all the teeth being relatively broad and strongly cusped.
But the posterior upper molar, in-, is much reduced in size and is less in bulk than the
antero-extemal cusp of the caniassial, p*. As in the other West African canids the
cheekteeth are f^. The outer upper incisor (i^) is much enlarged and caniiriform in
So
till ( AIIM \ lUil S ol WIS! M UK A
Fig. 10. Lyiium piain: skull, r\pc.' >>( iliaiim^, li.M. No, 7.7.,s,74,
' ; l,\tcr.il view
shape; the outer lower uicisor (/g) li also enlarged hut not to the same marked extent
and not caiiniitorni but riattisli and bieusped hke the rest.
Habits. The limiting dog lias, over a long period ot vears, attracted a good deal ot
attention which has resulted m an ahiuidant recording ot the chief features of its mode
of life. Some of its alleged habits, traditionally held and passed from writer to writer
or by word of mouth from one naturalist to another, appear to have been directly
contradicted by close and specialized stud\' of the species made in the field in recent
vears. Tlie most prolonged and concentrated investigations have been carried out by
Kiilime (i9''isa and b), Estes & Cloddard (lyfiy) and Goodall (1970), whose published
observations contaiji a great deal of information relating to the daily and iiightK'
habits of packs in the Serengeti plains and Ngorongoro crater (Tanzania), observed at
close quarters over long periods. Much of what has been noted in these areas is doubtless,
niiihitis nitilninlis. also applicable to West Africa.
The interest and repute of hunting dogs rest, as their common English name indicates,
on the \\a\ 111 which tlie\ secure their food; for, although sohtarv individuals arc
from time to time observed, these di^gs generallv live a higlilv communal existence
and hunt, skilfulK' and persistentb', m packs. These vary in number from an extended
famiK' unit of .iboiit half-.i-dozen to a sizeable group ot 30 or so. Ainthiiig larger
tlhiii this IS tiida\', even in East Africa, relatively rare. CertaiiiK' 111 West Africa few
p.ieks of this magnitude are likel\- to be found, though one such has recently been
observed, i'robably a dozen dogs might well be regarded as constituting a more
normal pack for this region. C'oncrete information ot anv kind relating to the area
covered bv this present work is, however, extremely scantw In East Africa packs have
often been touiul to show .i pretlominance, 111 numbers, of males.
LYCAON
8l
Fic. II. Lycai'ii picnis: skull, Tvpc of sliariiiif. B.M. No. 7.7.S.74, v, -< ' ; palatal &' dorsal views
Hunting is normally engaged in regularly twice a day, in the early morning and
late evening; but if the weather is deeply overcast, or for some other less apparent
reason, it may as an exception be carried out during the intervening period. A chase
has been known to be continued till after dark (Estcs & Goddard, 1967) ; and moonlight
hunts have also been recorded; but such nocturnal events in ill-lit conditions are
relatively rare since Lycaon hunts by sight rather than by smell. According to Kiihmc's
observations it would seem that what is acceptable as a meal may vary from day to
day, a change of diet apparently being desirable, readily available prey of a kind that
was one day willingly taken being passed by the ne.xt in the search for other flesh.
Estes & Goddard, however, found that something in the nature ot 69 per cent of the
kills were Thomson's gazelle, and iS per cent juvenile wildebeest. What is pursued is
S; Tin; ('AI!niv(ii(i:s or wkst m-iuca
doubtk'ss dctcnnincd to sonii: cxiait b\' the size ot the pack, the more numerous tliis
may be tlie greater tlie possibihty oi being able to overpower the larger sorts of ante-
lopes. Li West Africa it is unlikely that anvthing as bulky as a waterbuck or roan would
ever be tackled, though attempts would certainly be made on their calves. On two
occasions U. H. Kemp has foiuid the kill in Nigeria to consist of a fully mature harte-
beest in good condition. When a Land Rover stopped near one of these kills some of the
pack made attempts to drag the hartebeest away and two dogs did, in fact, succeed in
drawing it about 27 metres (personal commimication). Probably the main sources of
food in West Africa are gazelles, reedbuck, cob and hartebeest; but bushbuck, oribi
and crowned duiker, though less gregarious than the others, may doubtless also be
taken. A bushbuck killed by hiuiting dogs was in fact observed by David Brown in
the Borgu Game Reserve (Nigeria). Warthog are fronr time to time hunted. Hiuiting
dogs are. mdeed, known to take much smaller fry such as cutting-grass [Thryoiioinys),
giant Gambian rat {Cricctoinyf), hares [Lcpiis), ground squirrels [Eiixcnis) and so forth;
but this they usually do as independenr individuals, not as a pack though others may
be present and all on the move in search of more satisfying prey.
A look must be taken at some of the long-held views regarding the method of
hunting that have now been upset by close and continuous observation. Traditionally
these dogs were believed to pursue their prey tirelessly and relentlessly over vast
distances luitil the quarry faltered and fell from sheer odiaustion, the outcome of pro-
longed terror and ph)sical effort. It \\ as even said that in the case of the larger and
stronger antelopes such a chase might extend over as much as i s or 20 km and that
during this the leading dog would be deliberately relieved from time to time by relays
of others which were less fatigued. What in fact reaiU- happens is, to take the morning
luuit, that soon after dawn the various dogs of the pack emerge from their sleeping
quarters and wander slowly round defiecating and carrying out an extended greeting
ceremony, each with ever\' other member of the pack. This consists of poking the
nose into the corner of the other dog's mouth and of licking the lips and face. If all
are not present a not unmusical bell-like call is uttered to assemble distant members.
EventualK', some half an hour or so after daybreak the pack moves ofi at a trot in
more or less open formation to look tor game. When suitable quarry has been sighted
approach to it is made with some care, at a walk, in a slightly crouching attitude with
head and tail both lowered.
in the past it li.is been commoiiK' held that Lycaoii strikes far more terror tn the heart
of the antelope population tli.iii does the presence of lions; and that not only would
antelopes stampede at the first hint of danger but would actually entirely desert any
district tainted by the mere existence of luuiting dogs (Bere, iys6). Nevertheless, that
.uitelopes do not, in fact, always recognise their danger is shown m E. M. f-ang (1963)
where a herd of gazelles is recorded as actually riuming inquisitively towards an
approaching pack until they were no more than about 30 metres away before realizing
that they were Ln grave peril. Kiihme, in the papers cited above, also found that antelope
herds were cither not visibly wary of near-by hiuiting dogs or took flight only when
an obviously actively hunting pack on the riui towards them was within some two or
three hundred metres, thoui;h Estes & Goddard estimated the alarm distance in these
LYCAON 83
circumstances somewhat higher. It is, however, quite certain that a herd of antelopes
instinctively knows when a pack means basincss, exhibiting no fear even though dogs
may be at very close quarters but stationary, and but little alarm when a pack is merely
walking or trotting. It is this that enables the dogs to approach fairly close before setting
a herd in rapid retreat; or, in the calving season enables them to edge very near to a
mother and her fawn before snatching the latter.
Once the prey has been set in motion the dogs break into a run and follow, still in
more or less open order. If more than a single antelope is being chased there appears
to be no deliberate selection of any particular victim from the start, choice of that
which becomes the ultimate prey being as much a matter of chance as an)thing, resolv-
ing itself nito a question of the slowest member ot the herd. This is possibly a gravid
female or the tardiest to abandon grazing tor flight, most frequently a male. At the
start of a chase every dog follows that beast wliich is most directly in front of it, and
hence nearest, but they tend to switch to any one that is obviously less distant from its
aggressor, until ultimately the hunt nearly always resolves itself into the pursuit of a
single animal. There is no deliberate taking over the task of leading dog in relays, as in
the past supposed. If change takes place it is probably most often due to the fact that
the quarry, failing to escape on a direct course, starts to circle; and whereas the fore-
most dogs follow immediately on its heels others further in the rear can more easily
follow the chord of the arc and so, pursiung a shorter hue, gain gromid over both the
antelope and erstwhile leader. Yet this is not necessarily always the pattern; for Estes &
Goddard (1967) found, in Ngorongoro, that the dominant dog of the pack selected
the victim and led the pursuit. Nor does the hunt always resolve itself into a single kill.
E. M. Lang (1963), for example, describes an incident where a pack divided into two.
The search for prey is carried out at a trot of about 11 km an hour; actual pursuit
is at a steady 48 km an hour, a rate which can be maintained by healthy adult dogs
over long distances. Short spurts can if necessary be made at about 56 km an hour.
Not all the pack can keep up with this, and laggards or young dogs may still be i or
2 km in the rear at the time of the kill, not catching up till the leaders have already had
5 minutes or so at demolishing the victim. So far from the dogs patiently and steadily
rmuiing an antelope to exhaustion before killiiag it, as popularly supposed, Kiihme
observed that the kill was preferably made as swiftly as possible. Prey given a leading
start of 300 metres would be caught and overcome in a further 600 to 800 metres.
This according to other observers is rather short. Estes & Goddard reckon the average
chase to cover i.l to 3 km and to last 3 to 5 minutes; and Goodall found that the hunt,
if not successful, was abandoned after 4 to 5 km; but he did, nevertheless, follow one
that fruitlessly covered somewhat more than this. Since, moreover, this pack had
travelled 8 km m the preliminary search for suitable prey before it actuallv started
to hiuu it must in all have covered some 13 or 14 km.
The idea that these dogs lope untiringly behind a fleeing antelope until it drops
from exhaustion must also be abandoned. The antelope is not always captured; and if
the chase goes on past a certain distance the dogs, as Kiihme observed, can get just as
exhausted as their quarry. This is confirmed by Macintosh (1953) who writes of the
fuiding of an e.xliausted gazelle, pantmg in the grass and just capable of staggering ofi".
S4 Till 1 4, u s I \ (iHi s oi \i 1 s r Ai i(i( A
and some iSO inctrci nr sn awav a hunting ilni; m a similar stati.-. In such cases the
pack rests tor some s or lo minutes and then tonus up for a renewed Iruut. SimilarK'.
it a successful hunt should tarn out to provide an insutlicient meal for all or part ot
the pack, the late-arriving laggards tor example, then a second hunt is almost immed-
iately entered upon, at least by those with unsatisfied appetites.
Hiuiting is, to all intents and purposes. alwa\-s by sight, not smell, and it the quarr\',
during the chase, is obscured by grass the hunting dogs make intermittent leaps to
keep their prcv in view, the white tail tip at these moments becomnig very conspicuous
to an onlooker. Opinions have diftered regarding whether these dogs liimt mute or
not. This is dealt with later. Eventuallv, in the normal hunt the foremost dogs atter a
while fuld themselves almost literalK- at the heels of their prev and tliev then rcpeatedlv
jump t'orward to snap at the hindquarters, flanks or bellv. tearing away ribbons ot
skill or lumps of flesh. At length the animal is brought to a standstill or knocked down
and the dogs leap upon it. There is no attempt to kill it outright. It is either literalK
torn apart bv seizing the limbs and tugging m opposite directions, or eaten alive,
startinr; bv ripping open the bellv and devouring the entrails. It is, of course, impossible
to sav for certain but there is some likelihood that the victim does not suffer so much
111 this as might at first seem, being m a daze and a merciful state ot shock. All but the
tousihest parts — heads, horns, hoofs, and sometimes skin and leg boties — generally
disappear with astonishing rapidity. It has been mentioned above that not all hunts
end m a kill. There are widelv diverse estimates of the success rate. Goodall (1970)
records onlv 39 successful kills in 91 lunits. that is 43 per cent. Estes & Goddard (1967),
on the other hand, analysing a much smaller number of hunts found a success rate ot
over twice this. Ss per cent.
However, success rate apart, hunts do not always result 111 a satistactor)' meal, tor
other, stronger predators have to be taken into accoiuit. Lions are not above robbing
hunting dogs of their legitimate kill, and thev are too powerful tor the dogs to be
.ihle to do anything about it, even as a pack. Spotted hyaenas are another matter.
These too. of course, are bigger and stronger than hunting dogs but they lack proper
pack co-operation; so that the outcome ot attempts at superpredation b\- them depends
ver\- largelv on the relative numbers ot the two contenders tor the meal. There is no
doubt that hvaenas sometimes deliberately sit and watch hunting dogs, or w.mdcr in
their vicinitv eating tlieir droppings, waiting for them, or at times even deliberately
stimulating them, to set out on a himt, in which the\ then follow them. It the chase
ends with onlv one or two dogs m at tlie kill the hyaenas can then leap m, frighten
the dogs off and snatch a Itasty teed before the rest of the pack comes up and, b\- a
reversal of power, establishes ownership. If the hvaenas do pluck up enough courage
to sneak 111 again and seize part of the pre\- rlie\- are chased oft (E. M. Lang, 1963).
.uid thev are reduced to waiting on the outskirts nf the feeding pack until the dogs have
hiiishedand the\- can then safcK' close m and polish ofl the skin and bones or any other
residue that ma\- be left. There is, indeed, an odd sort ot enmity between hiuitnig
dotrs and spotted hvaenas. Sometimes the former tolerate the latter in close proximity;
at other times thev delight in mobbing a single hyaena, biting fiercely at its buttocks
so that ultiniatelv its onlv defence is to squat down and so protect its hindquarters
LYCAON 85
while snapping at dogs that come too near. But there never seems to be any question
of the dogs, even as a pack, killing a hyaena or even doing it much visible harm;
and it eventually slinks oft". Jackals, vultures and other birds of prey which almost
invariably try to share in a kill arc driven off and made to wait, though golden jackals,
in their nimble fashion, often manage to dash in from time to time and secure a mouth-
ful here and there.
Kiihme estimated that the pack he observed occupied a livijig area ot roughly 16
square kilometres and hunted over an area of between 100 and 200 square kilometres.
It will be appreciated that it is thus not normally easy or possible for a casual observer
to follow a hunt at close quarters from its inception to its end, the more so as it is carried
out at a fairlv high speed. Only if one chanced to be actually near at hand at the correct
moment and suitably moimted was tliis possible in the past; and the fmer details of
what actually takes place during pursuit were therefore witnessed by relatively few
people. This accounts for a good deal of supposition, of inaccurate observation or
wrong interpretation of a pack's behaviour. The problem is lessened today by the
existence of motor vehicles that can travel safely and easily across country; and this is
the method of observation employed by Kuhme and other modern field workers.
Himting dogs are little disturbed by the presence of a vehicle at a reasonable distance.
R. H. Kemp (personal communication) twice foimd these animals to pay little attention
whatsoever to his Land Rover, coming towards it and getting out of its way very
slowly or, when it stopped, lying down in front of it. One stood on its hindlegs to
get a better view. Nevertheless, Macintosh (1953) records the attack by a pack on
the tyres and wings of a slowly moving car.
Lycaon, apart from its normally wide hunting range is, except at breedmg times,
by nature a rover and in the course of a year may cover an extremely wide stretch of
countrs'. It is driven to this wandering habit partly by the migrator)' nature of the
antelopes on which it preys since these are pretty constantly on the move to fresh
pastures. On the other hand, some limit may be set to unrestricted nomadism by the
existence of other neighbouring packs, each claiming right over a more or less defmed
area, though there seems to be appreciably less insistance on strict territories in hiuiting
dogs than with many other predators. Anyhow, some degree of boundary observance
has been noted in East Africa where the large antelope population is sufficient to
support numerous Lycaon packs; but in much of West Africa the herds of antelopes
arc comparatively small, the hunting dog being consequently of relatively rare occur-
rence and the territories over which the few packs can roam correspondingly large and
probably, for this reason, less well defmed. During three months of breeding and
family upbringing the pattern of life is, of necessity, different since the interest of the
entire pack is completely centred roimd the earth or earths m. which the pups are
being raised. It therefore, apart from brief daily sorties after food, becomes something
of a sedentary unit. Such a pattern postulates a set breeding season for all females.
Kiihme's pack contained only two adult bitches, each with a recenth- born litter;
it would be interesting to know what happens where there arc several adult bitches
breeding at different times.
It has already been mentioned that hunting dogs, though subsisting mainh' on
S6 Tin; (,ABMVoui:s oi \\\%t amuca
ai;tclopcs, apparently like to alternate tront one species to another; and that when
these are not conveniently available they turn their attention to other kinds of flesh.
Ir is virtually certain that nothing would come amiss to them so long as it was readily
obtainable; and they certainly do not hesitate to eat the dead of their own kind, hi
parts of Africa, and particularly in the past when they were in larger packs and of more
common occurrence, they have shown themselves to be highly destructive of cattle,
sheep and goats; .and they have themselves been relentlessly hunted and shot or killed
with poisoned bait on this accoiuit. It was probably their depredations amongst
domestic flocks that chiefly earned tor them the reputation ot ijidiscriniinately slaughter-
ing tar more than they could possibly eat; but here again Kiihme's observations rmi
coiuiter to this widely held belief since he foiuid that his small pack never killed more
than was absolutely necessary to satisty tlie S adults and 15 whelps. When the Litter
were very yoiuig and ate relatively sparingly the toragers regularly killed and brought
home a single 30 kg antelope morning and evening; but when the pups had grown and
become more dcmandijig they doubled their twice-daily kill. This figure is, of course,
nothing more than a rough estimate; but at the end of the rearing period when all
23 dogs might be assumed to eat practically adult meals this would work out at an
average of something like 5 kg a day. On the other hand, Estes &: Goddard (1967),
working on a pack numbering 2t, estimated that there was an average of 2-7 kg a
day of meat available to each dog; this amoiuit would however vary with the size ot
a pack between 2 and 4 kg, the smaller packs getting the larger anKiimt. It is interesting
to compare these estimates with the figure given tor zoo animals of i.V kg ot meat
daily (Dekker, 1965). Such captive specimens would, of course, get relatively little
exercise compared with those hunting for themselves in the field and their demand
would be correspondingly less. It may here be noted that these zoo specimens were
given an occasional whole chicken or whole pigeon, mdicating that, in the wild,
birds, such as francolins, may be included in the diet of Lycaoii.
There is, however, some evidence in support ot the charge that these ajiimals kill
more than they can eat. R. H. Kemp (personal communication) twice observed in
Nigeria that the kill had scarcely been damaged except for one hindleg and the belly
ripped open. At IS a.m. in the Borgu Reserve he once observed about a dozen animals
which had just made a kill ; but the rallying hoot heard in the distance not only indicated
that the pack was in fact larger but, strangely enough, served also to call away the
dogs by ones and twos trom their newly killed hartebeest, \\'hich still remained un-
touched at dusk. This must be luiusual; but the commonest expLuiation ot only
partially eaten prey is probably that these corpses are the outcome ot second hiuits by
imsatistied members ot a pack, too few in number to be able to consume all ot their
kill. Hiuiting dogs, unlike many other carnivores, never feed a second time from the
same kill.
Kiihme never observed his pack to drink, and it is to be presumed that much ot the
necessary water can be picked up trom blood or from licking their coats when wet
with dew, or from like sources; tor it is known that these dogs can go tor long periods
without drinking. But they certainly do drink on occasion. (loodall says that they will
do so daily it possible, and quite clearly deliberateK' seek out water-holes to this end.
LYCAON 87
Apart from tlieir understandable fierceness in the pursuit and capture of tlieir
necessary prey, these animals seem by nature to be of an essentially friendly and sociable
disposition amongst themselves. They do not seem to engage in the fierce protection
of boiuidaries practised by the spotted hyaenas; they arc not even aggressive towards
other packs unless these approach too near to the breeding burrows (Goodall, 1970).
There appear to be no records of any attack b)- hunting dogs on man ; and in fact,
they seem in nature to regard human beings witli some indifference. They can, however,
be successfully tamed but have an imwelcome drawback as a domestic pet in their
strong odour. This is probably due to glandular exudation; but they have also attaching
to them another smell that seems to permeate their very bones and persists for many
years after their death. A cupboard containing thoroughly cleaned skulls of Lycaon
has a particularly nauseating odour, shared, it is true, with lu'aenas and some other
flesh-eaters.
Amongst their own kijid these wild dogs are playful and rarely quarrel, and then
only briefly. They share the same shelters, even at breeding time, and peacefully look
after each other's young. They have, according to Kiilime (1965 a & b) developed a
classless social system that involves the ready sharing of both food and labour. Such a
view is diametrically opposed by Goodall (1970) who, while agreeing about the sharing
of food and labour, fmds that there is a recognized social order not only throughout
the pack as a whole but m each of the sexes separately as well. This conclusion was
reached as the result of long and close observation of a single pack, and was derived
trom the interpretation of submissive body postures. The pack was normally led in
the hmit by an old male; but even when on one occasion this dog fell into fourth place
he was still foimd to influence the direction taken by the leaders. Estes & Goddard
(1967) found that there was a pack leader, cither male or female, but no other rank
order.
It has been noted earlier in this present work (on page 48) that Wyman (1967)
observed a food-begging ritual amongst the jackals. An exactly similar ritual had
previously been discovered in the hunting dogs by Kiihme. That is to say that on the
arrival home of the toraging party those dogs that have remained on guard, and those
young which are ot an age to require a full meat diet, excitedly thrust their noses at
the lips of the returned hunters luitil these latter, in response, regurgitate some of the
meat with which they have recently, and partly with this end in view, been filling
their stomachs. This performance is repeated until those that have remained at home
have received enough food to satisfy their hunger. The yomig, in their imrestrained
impatience, often go beyond the mere ritual of nose-thrusting and become aggressive,
actually biting the adults in their lips or legs. It is this voluntary sharing of food that
enables labour also to be parcelled out between those that hunt and those that undertake
the vitally important duty ot guarding the helpless young against marauders. It is
extremely interestijig to note that the males take a tremendous part in the raising of the
young. On their return trom himting they make a point ot regurgitating food for
them in preterence to the females ; and, it was observed in one pack, that when, through
death, no adult females were left the feeding and bringing up of the orphan pups was
willingly and successfully undertaken by the males alone (Estes & Goddard, 1967).
SS illl. (:A1(M\(MMS (11 W 1 s I AlKliA
N(i niotlicr, thcrctorc, is essential attcr the first slmrt pennd oi milk feeding. Care
of tlie voting, botli in tlie steps taken to guard them and in the matter of seeing that
tliey are properly ted. seems to plav a dominant role m pack behaviour; tor later in
lite wlien the yoiuig dogs reach the point of accompanying their elders on the hunt
thev arc given absolute priority over the kill, all the adults retiring until the pups have
first tully satisfied themselves. This, according to Kiihme, is the onl\- kuul of pre-
cedence recognized within the pack.
This begging ritual at the homecoming has been extended to a similar, but un-
rewarded, greeting ceremonv carried out between adult members of the pack m the
earlv morning and late afternoon before setting out on a hunt, and especially between
those about to remain behind and those who sallv forth after food. Any ot the voung
who attempt to take part in this ritual m which the adults tonnallv register with each
other their fellowship ot the community are snarled or snapped awav; and it is not
until they are accepited without protest at this twice-daily ceremom of mutual recog-
nition that they can be reckoned as tully adult members of the societ\'.
In the course ot their hunting, feeding and the ripping ot their still living prey to pieces
these dogs become well splashed and smeared with blood; but thev take good care
later to lick themselves quite clean. This is. of course, easier with their short, sparse
bristle-fur than it would be had thev the dense tangle ot long tuie undertur possessed
bv the foxes, hi this type ot pelage, too, there is less shelter tor fleas and other ecto-
parasites; but, ncvertlicless, a good deal ot time is spent scratching at their tiu'.
Like other dogs, these utter a variet\' of sounds according to mood or circumstances.
Their most famous note is the ott-repeated bell-Iike "liooo-liooo" used as an assemblv
call, either to gather the resting pack together tor hunting or, when c]uartering tm'
game, a find has been made and there is immediate need to concentrate all forces on
the line of assault. It is also used should a dog tuid itself separated from the others. The
cogenc\' of this call is indicated bv the observation made b\' R. H. Kemp, noted above,
that it was sutticientlv compelling to induce a dozen dogs to abandon a newly-made
kill without feeding. Although it is soft and musical it carries a ver\' long distance,
2 or 3 km.
It seems that most ot a chase is earned out in Mlence, but the dogs break into guttural
or raucous velps when closing in tor a kill and emotion rises to a pitch. This becomes
an almost bird-like chirruping at the kill. Tins last sound is uttered alsii at other mom-
ents ot high iuiticipatorv excitement such as when about to set forth on a hunt, or
when mobbing hyaenas. The normal cr\' ot alarm at possible danger is a growl breaking
into a short, deep, harsh bark which, however, is not really much like that of a domestic
dog. A softer bark is used as a greeting; and there is a good deal of whining as an
aeccnnpaniment to begging for food.
ComiiKiiiK', hunting dogs, leading a nomadic lite, he about during the da\ or
sleep at night simply curled upi on the surface in whatever slight cover the vegetation
affords; but when it is a question of breeding, the whole life ot the pack becomes
centred round a burrow or burrows made in the abandoned holes ot aardvarks, giant
pangolins or other excavators. These luiderground shelters have not been described
111 aii\- detail but appear to be lined with grass. Though their main purpose is for
LYCAON 89
parturition and tlic protection of the young they may also be used by any aduk as an
occasional refuge against the n^iddax- heat or inclement weather or tor sleeping at
Jiight. Not infrcquejitly two or more bitches litter in a single earth (Brand & CuUen,
1967) or in closely proximate liolcs, cither without quarrelling or with at most minor
disagreement. Indeed, as part of the commiuial life and communal sharing out of food
and duties they look atter, and even suckle, one another's yoiuig.
The abscjice of precedence within the pack appears to extend to the matter of
mating, the males not contending for the females as they commonly do in otjier groups.
Coition takes place several times over a matter of a couple of days. The period of
gestation is from 69 to 73 days; and there may be any number between 2 and 19 pups
in a litter (Goodall), probably 7 or 8 being an average size. The pups, which weigh
350 to 380 grammes at birth, arc suckled for from 10 to 12 weeks, their eyes opening
between the loth and 14th days. This is a tricky period and should anything happen
to disturb the mother she may, as has been found many times in captivity, eat her
yomig. If she is upset or thinks them in danger she carries them in her mouth from
place to place. The presence of the male seems to be necessary at the period immediately
following birth (Cade, 1967; Crandall, 1964). Crandall foimd, indeed, that the mother
did not clean the new-born puppies, this being done by the male, whose active duties
in comiexion with the litter were then over. But though the close presence of the father
becomes for a period tuiwelcome he must remain within easy distance.
From the time of the openijig of the eyes onwards the pups, though very much
centred on the nest, quit it more and more frequently and for increasingly longer
periods. Much of the suckling, if not all after the first couple of weeks, is carried out
above groiuid, actually in or not far from the entrance to the earth. Diuring this act,
which may take place either at the direct vocal invitation of the mother or in response
to whining requests from the yoiuig, she may either lie down or stand. In the latter
case the pups stretch up on their hindlegs steadying themselves with their forefeet
against the mother's belly. Suckling sessions last only from 2i to 3 minutes (Goodall),
in sharp contrast to the lengthy bouts indulged in by young spotted hyaenas.
Though breast feeding is continued for nearh' 3 months the pups are in fact given
meat after about a fortnight. This is specially prepared for them by the female chewing
to small size some of the meat she has begged from the returned himtcrs and regurgitat-
ing it for the second tune for her offspring. The male parciu, too, may take some part
in this, as he does, now, in the general care of the litter. Gradually in this wa)-
the young are weaned luitil they take large pieces of flesh for themselves, regurgitated
by the hunters ; for from this stage the eiu'nc pack evinces a lively interest in the welfare
of their next generation ; and, as has already been said, should, through accident, no
adult female remain the males competently see to the upbringing of the litter. The
whole process of raising a family to the point where the young dogs can take care of
themselves and join in the himting occupies about three months. Throughout this
period the breeding-groiuid is never, or only exceptionally, left unprotected, a varying
Jiiunbcr of dogs and bitches always remaining on guard duty while those in search of
food are absent. Dining the period of upbringing the site of the home burrow is
changed, over a short distance, from time to time, especialK- it any danger has tlircatencd.
90 THE CARNIVORES OF WEST MUKA
To do this, in tlic early stages, tlic pups arc carried in tin- mouth to tlic new site, other
teiiiales besides the actual motlier sometimes assisting.
It is interesting to note that in order to escape very pressing danger a jiunting dog
lias been kno\\ii seeiningl v to teign death with complete success though it is not clear
whether this was, in tact, merely very convincing acting or an actual coma brought
about by an attack of very powerful domestic dogs that had been so closely pressed
home as to cause real death, by worrying, of a companion. The hunting dog in
question maintained its apparent demise though dragged over the ground tor some
distance, and for some further time atter its aggressors had gone away ard left it lying.
Nothing is definitely known ot the expectancy ot life of these animals in their natural
enviromnent but one has been known to live in captivity to an age of about lo years.
Taxonomy. It has alreadv been mentioned, in connexion with the general des-
cription of the Lyme;; pelage, that Matschie (1915) used the differing proportions of
colours and their varving situations on the body to create a large number ot indepen-
dent species. The material c-in which this study was based was extremely limited for
each of the separate species proposed; and the characters on which these species were
based, moreover, were such as might be suspected as intrinsically iticonstant. Further,
apparent diflerences of size cannot be ascribed to entire populations on the strength of
single skulls possibly ot varying ages and of unknown background. Holz (1965),
as the result ot statistical study, which embraced skull measurements, came to the con-
clusion that there was absolutely no foundation tor Matschie's proposals at specific
level, and that valid separation even at racial level was doubtful as ajiy distinctions
appeared to be clinal. It is, of course, possible that, at some future date, given a statisti-
cally significant number of specimens from each of a variety ot localities, some valid
racial separation may become evident. But any present attempt to maintain such dis-
tinction on the basis ot the tew, mostly single, specimens that now exist seems to
pretend to an luidcrstanding ot the situation that dots not. in fact, exist, and is without
real meaning or value. Subspccitic names are therefore ignored in this present work.
The accompanying table shows the measurements of the onlv two British Museum
West African specimens, together with those two of Matschie's proposed species for
which measurement data arc given in his paper.
91
Table 4 : Numerical data for Lyiacii piclus
Mani
Djannaga
Kete Krachi
(Shari River)
Lake Chad
(Togo)
(Togo)
Type of
B.M. No.
Type of
Type of
sharicus
1937.7.10.1
mangucnsh
mischichli
Vegetation
Sahel
Sahel
Sudan
Guinea
Number in mean
I
I
I
I
Condylobasal length
189
210
—
—
Basilar length
173
192
C.I 90
—
Palatilar length
8S
98
—
—
Zygomatic breadtli
122
135
125
128
Upper cheekteeth breadth
68
77
7^
74
Nasals, length
67
71
65
76
Interorbital breadth
37
46
49
49
Postorbital constriction
37
42
—
4a
Braincase breadth
69
72
69
69
Toothrow (c — in-)
n
89
82
87
p* length
19-4
22-2
—
ml + Ml- length
21-5
25-6
—
—
nil length
22-5
27-8
—
■ —
(112 + H13 length
14-9
i6-2
—
—
Head & bod\-
900=1
1 150''
1130"
Tail
325a
—
375
410
Hindfoot
220*
—
200
215
Ear
C.90
—
120
125
RATIOS (per cent)
Tail/head & body
36
—
34
36
Zygom. br./condylob. 1.
64
64
—
—
Braincasc/condylob. 1.
36
34
—
—
Braincase/zygom. br.
57
53
55
54
Palatilar l./condylob. 1.
46
47
—
—
Interorb./postorb.
100
109
—
116
p^jc-iifi
25-2
24-8
—
—
p*jm^ + nfl
90-0
87-0
—
—
"'l/'"2 + '"3
151
172
—
—
*. As given on the labels; Thomas's type diagnosis figures differ.
''. These appear to have been taken from the prepared skin.
9i THE CAnNIVORES OF WEST AEKICA
Family MUSTELIDAE Swainson, i(S35
Polecats, Weasels, Otters, etc.
General. The Mustelidac constitute aai important family of very wide distribution
throughout the world except in Australasia, Madagascar and the polar regions, being
most common in the northern temperate belt. The family contains animals with such
\\ell-known English names as polecat, stoat, ferret, mink, marten, badger, ratcl,
skimk, weasel, otter and others. These are carnivores ot from very small to mediinn
size, often with strikingly patterned coats and mostly with a greater or lesser degree of
objectionable odour, the secretion of special glands, in some cases invohmtary and
constant, in others deliberate as a defensive act.
The mustelids may be terrestrial, arboreal, rivcrme or lacustrine; and there is an
cxtralimital genus tjiat lives in tlie sea. In general diey have long relatively slender
bodies and markedly short legs for their size; there are five digits on each foot, webbed
more or less deeply and armed with claws that are non- retractile. Progression is, in
African mustelids, mostly digitigrade. hi the Mustelid.ie as a whole development ot
an aural bursa is variable; but as far as all the West African representatives ot the
family are concerned it is always absent. 13y nature these animals are fierce hiuiters,
being responsible for the destruction of large quantities of small mammals and birds,
and in some cases a certain amoiuit of insects. To the great majority of Mustelidac
any kind of flesh is welcome, preferably freshly killed by themselves but they will
consume that killed by others if it is readily available, and even carrion it they are
hard pressed. The main prey is rodents, hares, slnrews and birds — in Africa up to the
size of a francolin or the young of larger breeds. Eggs, too, are welcome, and, less
commonly, reptiles and amphibians. More exceptional diets are tound in the river-
haimting otters, which live on tish, and in the ratcl. which often robs wild bees' nests
for honey and grubs.
Skull. The rostrum is short and blunt; there is no alisphenoid canal. There may
be 32, 34 or 36 teeth, each of these totals occurring in West Africa. This is due to the
different arrangement of the cheekteeth in the subfamilies: 3-j or 'j-^ or jy,. There
arc always 3 incisors and i canine, top and bottom on each side.
Taxonomy. Five subfimilies are now generally recognised (Simpson, 1945). The
position, liowever, is by no means as straightforward and clear-cut as this might
imply. Palaeontological evidence of atlinities and present structural morphology are
both complex and in the past the tamily has been grouped and regrouped in a number
of diff^erent ways. Pocock (1921c) gave a useful summary of the variously held opinions
between Gray in 1869 and the time of his own paper, hi this last he assessed the differing
views, gave an accoimt of what he regarded as relevant external characters and evolved
yet another, virtually independent, classification in which he recognised no less than
fifteen subfamilies in respect of living forms alone. Simpson (1945) once more reviewed
the position, in the light of both living and extinct forms; and, regarding Pocock s
treatment as "excessively inflated", reduced the subfamilies of ext.int mustelids to the
five uo\\' widck accepted.
MUSTELINAF. 93
Only three of these subfamihes occur in West Africa, the two that do not exist
there being the Melinae (true badgers) and the Mcphitiuae (skunks). The tlircc sub-
famihes relevant to this present work may be distinguished by the following key.
KEY TO THE SUBFAMILIES OF MUSTELIDAE
(Previous key page 28)
1. Dorsal pelage very long and with a well-defmed longitudinal pattern of black
and white bands; total length of skull luidcr 70 mm; cheekteeth 3-5.
Mustelinae {pai^c 93)
Not like this 2
2. Fur coarse, the upper side typically whitish or pale grey from the crown ot
the head to the root of the tail (but this area often reduced or, not infre-
quently, entirely black like the rest or the pelage) ; claws of the forefeet
very long and powerful; ears with no free external shell; skull with
very little iiuerorbital or postorbital constriction; bullae large and
subglobular; cheekteeth 1^ .... Mellivorinae {p(,oc no)
Fur soft and sheeny; postorbital constriction fairly marked; bullae rather
flat and subtriangular; cheekteeth ^ (but ;)i often missing).
Lutrinae {page 128)
Subfamily MUSTELINAE Gill, 1S72
Polecats, Weasels, etc.
General. This subfamily contains most of the smaller mustelids, fulfilling the
important function of holding the rodent population in check. They, ot course, kill
other prey too, mostly birds, insects and some reptiles. Their bodies are often long,
slender and lithe, the whole effect being added to by an unusually lengthy neck; but in
the African species these features are not so pronounced as amongst the European
and American forms. There is no bursa on the ear piruia. The tail is short to moderate,
that is to say at most never quite as long as the body; and it is sometimes showy.
The pelage is often striking ; and in the case of the mink {Miistcia, subgenus Liitrcola)
is so much sought after as the most luxurious of commercial furs that it has become the
subject of a flourishing farming industry and of selective breeding experiments directed
to the production of distinctive colourings. Some members of the subfamily inhabiting
the colder northern regions undergo a pelage change in the winter, the brown coat
becoming white; in the case of the stoat {Mustcla enninea) the tip of the tail remains
black and the fur is much sought after on this accoinit, most notably as a decorative
trimming to official robes. Another well-known member of the subfamily is the
ferret, a domesticated, and often albino, form of the Asiatic polecat, bred, and to
some extent trained, as a hunting animal to flash rabbits or other subtcrrestrial species
from their burrows.
The t;eiuis MiistcLi penetrates into the Mediterrane.in region of northern Africa;
94 II" (AUNUDiti.s oi \\i:sr aiuila
but ill so tar as the true Etliiopian region is concerned tliere are three Atrican nnistehne
genera, laoiiyx. Piwilictis and AuTi/iu'j/e, ot which only the first two are found witluii
the area deah with in this present account. PoccUcfJiilc is essentially a soutliern African
genus but extends as tar north as Tanzania and parts of the Congo and Uganda; and
there seems no obvious reason why this animal, which closely resembles htoiiyx in
external appearance except tor a tar more slender bodv, should not have spread turther
westwards across tlic continent. Tlie two genera witji \\hich, however, we are here
concerned iiiav be distinguished as tollows:
KEY TO THE GENERA OF MUSTELINAE
(Previous ke\' page 93)
White marking on the tace continuous across the trout; tips ot tlie ears only
very slightly white; total length ot skull under so mm Poccilictis (/)ii^'c 104)
White facial marking broken above the eyes; tips ot the ears conspicuously white;
skull ss mm or more ....... Ictotiyx (jhit^c g^)
C.ciuis ICTONYX Kaup, iN.^s
Atrican Polecats
Zi'iillii Okcii, 1S16, Li'liiiiuch dcr Natin\^fscliklilf, 3, Zocil. pt. 2; m. Okeii's work has been ruled to lie
imavailablc by the International Commission on Zoological Nomenclature, Opinion No. 417 ot 1956.
This name is the Spanish zoiilln, a diminutive ot zona a fox.
ZoriUa I. GeofFroy, 1826, Dii'lioiiiiaiif iliusitjHLit'Histoin- Kalnirllc, 10: 215. This name has been suppressed
by Opinion SiS of the International C'omimssion on Zoological Nomenclature and placed on the
Olticial Index ot Rejected and Invalid Names in Zoology with the Number 1912. {Bull. :ool. Noiiiciicl.
M- 1.S3-154)-
hlpiiyx Kaup, iS^s. Ddi 'fliiincicli . . .. i: 152. liy Opimou MS ot the International Comnnssioii this
name has been placed on the OHicial List ot Generic Names with the Name Number 1759. Type
species. b\' inon. A Inpsm calami tor liioiiyx.
This is a moiKispecitic genus the description ot which is, thus, adequately tunii.shed
by the accoiuu ot the species which immediatelv tollows. There has been very con-
siderable, and heated, argument m recent years over the tjuestion ot whether the
name laony.x shoulci replace the commonly used ZorilLi. The matter, now resolved by
Opinion XiS. w .IS .1 higliK complex one the various aspects ot which can be toujid bv
those interested ^et out 111 1 11111.1 (ii)(ti. mjCis and n/iCi) ; Van Cn Idcr {ti)<>(t) ; and I Icrsli-
kovit/ f K/i^ and igOd).
ICTONYX 95
ICTONYX STRIATUS (I'crry) African Striped. Polecat or Zorilla
Bradypiis sirialiis I'cny, 1810, Aruviit or ihe Miisvuiii oj !\'alnral History, part 1 1, pi. (41) and text. Cape ot
Good Hope. A note on the validity of this name is given by Hollister (1915); and the name siriatus
Perry has now been placed on the Official List o( Specific Names in Zoology with the Number 2202.
The name siriatus is the Latin adjective meaning marked with channels or bands, given in reference to
the dorsal pelage. The generic name is fronr the Greek hraJys slow, and pons foot, meaning slow-
footed.
Mephitis capviisis A. Smith, 1S26, A Dcscripiiir Cataloi^iic of tin- Soiiili Ajricaii MiistKiii, pt. i. Mammalia:
20. No locality named. This generic name is the Greek word tor a disgusting smell, and refers to the
body odour.
Miistcia zorilla p. Si-ncgalciisis ]. B. Fischer, iS2y, Synopsis Aiaiiinialiuiii: 219. Senegal. The generic name
is the Latin for a weasel.
Mustcia zorilla Smuts, 1832, Disscrtalio Zoolo^ica, eiiuiiicrationviii Maiiinialiuiii Capcusinm (omincns, 12.
Not Viverra zorilla Schrebcr ( = Spilogalc).
Mephitis africana Lichtenstein, 1S3S, Ahli. preiiss. Akad. IViss. for 1.S36: 284. Cape of Good Hope, Scne-
gambia, Abyssinia, Barbary.
Rhabdo^ale iiinsteliiia Wagner, 1841, in Schreber's Saufttihierc, Supplement, 2: 219. Cape of Good Hope.
The specific name is Latin for weasel-like.
Zorilla striata var. seiiegalaisis Gray, 1865. Senegal. Type in the British Museum, No. 50.7.8.38, }; skin
good except for the terminal part of the tail probablv missing, skull good.
Distribution and general. The African striped polecat is very ohen, both in
literature and speech, termed the zorilla or zorille, sometimes zoril, a Spanish name
having no real application to Ichviyx. The steps by which this came about are involved;
but reduced to their simplest terms the position stems from confrision arising in the
early 19th century between the African polecats and the central American spotted
skunk {Spilogale), of rather similar appearance and of which zorilla was the local
Spanish-American name.
This is one of the most striking ot small Atrican mammals, with its conspicuously
banded black and white coat; but it must not be confused with the closely similar,
though very much smaller, striped weasel which may occur in the same locality.
When the animal is at rest in grass or other cover this bold pattern doubtless serves to
disrupt its form and conceal its true nature; but such coloration is just as likely, in the
open into which the animal must often enter, to act as a warning to rapacious birds
and other beasts ot prey, cautioning them to avoid attacking a victim capable of
emitting a nauseating fluid. Couspicuousness in these circumstances is added to by
the absence from this entirely black-bellied animal of the usual protective coimter-
shading.
Ictoiiyx striiitiis ranges over the drier zones of Africa from the Cape northwards to
Sudan and Ethopia on the cast, and thence across to Senegal in the west. It also occurs
in South-west Africa and parts of Angola. This is an animal chietlv ot the drier open
woodlands, mostly (as tar as West Africa is concerned) the Sudan and Sahel, but
penetrating also into the Doka zone. It would seem to be pretty common in the southern
and eastern part of its range; Shortridge (1934) characterises Ictciiyx as one of the most
ubiquitous mammals in Southern Africa, in all kinds of vegetation, down to the sea
coast and even in town gardens, [t also occurs up to 2000 metres on mmuitains. It seems
yd Till- CAUMVOHliS (.U WLST AlUICA
to W nukli less ccmniuiii in tlic area dealt witli in diis present account — tliougli this
apparent relative rarit\' in West Atnea ina\' be due merelv to lack ot collecting. Onl\'
S skins and s skulls exist in the British Miiseuin Iroiii tins area: from Senegal (2),
larniso. Nigeria {2) and Zaria (Nigeria).
Description. Tlie African striped polecat (tig. 12) varies a good deal in size, and
such dirterences have been used, at least in part, diagnostically in tlic erection of local
races. The males are generally held to be appreciably larger tlian the temales; but this
distinction is not evident in tlie scant data available tor West Atrica — one male skull
compared w ith tour females. Tlie pattern of the pelage is broadly the same thr(Highout
the animal's wide range but the proptortions of its coniponcnt elements, that is to say
black and white, var\ , seeminglv trom place to place and so commonly constitute the
m.ijor basis ot proposed subspcciation. The lengtli and quaht\' ot the fur also var\.
In West Atnca this polecat attains a ]icad cV body length ot between 320 and 350 mm,
the narrowly bushv tail being some 250 to 2N0 mm. The body is tairlv, but not
markedlv, slender, and the legs short, tlie animal standing abi-nit lis to 130 mm at the
shoulder. Tjie weiglit ot an aduh may be about j kg en- sometimes a little more,
Tlie dorsal pelage is tairly long and dense but loose in texture. It generally has a
sheen. The underparts are considerably shorter and often sparser, sometimes almost
naked over the bells and chest. On the back it may be regarded, purely tor convenience
ot description ot West Africa specimens, as basically white marked with three highly
conspicuous longitudinal black bands. (Tlie converse of this prob.iblv more accurateU-
expresses the true position). The centre one ot these black b.ands, running along the
spine, starts trom the crown ot the head; the two lateral ones originating a little short
ot this, at the tore part ot the neck. These three lines are here narrow (roughly s mm
or less broad) and they riui approximately parallel to not quite the middle of the
back. There the two outer ones broaden somewhat (to roughly 12 to is mm) and
diverge, curvinc; outwards to the top ot the flanks and then in again to the root of the
tail, where they meet or nearly meet, thus enclosing a regular elongated ellipse. The
medial black line continues its path to tlie tail through this \yliitc ellipse but at about
tJie midpoint broadens into a black blob, roughly ot diamond shape and 30 mm wide
at its broadest point, then narrows again posteriorly, meets the two outer bands and
continues on to the basal part of the tail. The sides of the neck, the shoulders and the
entire tmderparts, together with the legs and teet are wholK black. The long-haired
but nevertheless rather narrow tail is an intimate mixture ot black and \s'hite, the
latter colour dominating in West Atrica (except in the basal 70 mm) though not alw.iys
elsewhere.
The pelage is composed ot long tine undertur .uul (.oiisulerabK longer and stouter
bristles. These elements are selt-eoloiired throughout, either all-black or all-white;
and. in general, the two colours occup\' clearly defined areas ot skin without inter-
mixing. 13ut there are exceptions. I lere and there one or more isolated pure white
bristles occur amongst black patches; and sometimes there are small islands of white
m the black .neas; and, more rareb', in some (extr.ilimital) specimens or forms, bristles
(dlouri'd .It the b.ise ami arising 111 the bl.ick arcis. beconic. .l^.llnsl the prewiiling
character ot pi^meiit.ition, white in their termin.il li.ill. Sneli aberrations, ,\\](.\ especialK'
ICTONYX
97
,xlw
Fk;. 12. African Striped Weasel (Pocciliclis liby7-j6S. Tvpc species IWciliciii liliyni
(Heinprich & Elireiiberg) [-- Mmlchi Ubyca HcmpricK & Ehreiibcrg) from Libya. This name is trom
the (Ireek jh^ihi!o< v.iri-cttloured or pied and Jt7/V weasel, in reterence to the black and white pelage.
Distribution and generaL The striped weascK are, at least in West Africa, ver\-
closely similar in external appearance to the striped polecats though ot much smaller
size. For a long time the two were, indeed, reg.arded as congeneric; but there are
certain small external and cranial differences which led Thomas & Hmton to erect
for the weasels a separate genus. Seven species have at one time or another been named
but these are all now regarded as either synonyms or merely forms of a single species
lihyui. The range of this, and hence of the genus, is restricted purely to the northern
portion ot Africa, from about 12"'' Nortli, that is within the Sudan woodland zone, to a
little short of the north coast from Morocco to Libya and lower Egypt. Poccilictis is
thus essentiallv a dry or very dr\' country animal, about half-a-dozen local races being
recognised. The general generic diagnosis and the points which distinguish Poccilictis
froin Ictonyx can be sufficiently gathered from the description of Uhyca which follows.
Taxonomy. Whether Poccilictis merits separate recognition from Ictonyx as a genus
IS open t'ives rise to ditfereiit "tonus". Soinetniu's 111 these dorsal variations
MELLIVORA 113
tlic divisoii bct\\ccn black and wkite is more or less clear-cut; in other cases tlie black
is '"frosted" with white, verv lightly over the hindquarters, more densel\- over the
shoulders and neck.
The pelage, though contorninig throughout the species to an overall basic pattern,
nevertheless exhibits some variety in the fuicr details of its composition. Two elements
are present: abundant long, strong, straight, flat bristle-hairs, which dominate the
pelage; and much finer, curly, terete underfur. Both are most commonly cither all-
black or all-white. The white bristle-hairs arc approximately 0-15 mm in width, their
length varying between 2Z and 30 mm; the black arc a trifle less broad, about 0-13 mm,
but their length is generally somewhat greater and may, indeed, reach as much as 38
to 40 mm, especially on the lower back. There is, in fact, some tendency for the fur
to lengthen from neck to rump. The underfur is conspicuously fmer though, at 004 mm
diameter, still pretty coarse by comparison with softer-furred carnivores. In the majority
of cases bristle- hairs and underfur arc of one colour throughout their lengths — all-black
or all-white; but si^iiata is exceptional in that some of the white bristle-hairs have black
tips and some ot the black bristlc-hairs a white subterminal band, not sharply detmcd
but about 5 mm broad.
The white areas of pelage ma^• consist purely of white bristle-hairs and ■\\'hite imder-
fur; but t]uite commonly the latter is black though, sijice it is short and well overlain
by the white bristle-hairs it has little or no eifect on the overall colour. However,
when such an area is invaded bv black bristle-hairs as well, the total effect is grey. The
black areas are generally pure black; but sometimes there are scattered white bristle-
hairs amongst it. The underfur is normally about 18 to 20 mm long and is generally
relatively sparse and so plavs little part in the overall tcxtiurc or appearance of the iur;
but in some cases it is distinctly more abundant, more undulate and much longer,
reaching 24 to 27 mm. In the form known as si<^iiata the basal portion of the bristle-
hairs, as well as the underfur, tends to curlincss.
The body of the ratel is fairly long — some 60 to 70 cm — but distincth- thick-set
and broad across the back. The skin appears to be remarkably loose; and many field
observers have maintained that it almost seems as though it were independent of the
body and that the latter could turn and twist freely inside it. The head, for this bulky
body, is unqcstionably small, rather flat, and with a short muzzle, the whole tront of
the face being black. The eyes are not very large; and the ears, though shaped and
folded rather in the manner of human ears, have no independence of the head but
appear as little more than ridges of skin. They have no bursa. The legs are short and
sturdy, the animal standing some 23 to 28 cm at the shoulder. The feet have 5 toes each,
armed with very strong claws, those of the hindfeet short, but those ot the front
remarkably long, the central three of much the same length (25 to 30 mm) and forming
a unit connected by short webs, the ist and 5th digits lying posterior to these. The soles
are thickly padded and naked to the wrist, the posture being semi-plantigrade. The tail
is short, naked below at the base but clad with long hairs for the rest, though narrow
in form since these are fairly close-lying. The circumanal region is hairless; there are
two very large scent glands situated within the anus and these according to Pocock
11-1
TlIF, CARNIVOUI-.S or W'fST M I! 1 ( A
Fin. i6. .l/i7/ii'iira uipciisii: skull, li.M. No. 26.S.7.
I ; lateral \'icw
([920) "discliargc cnpiousK a surtot.itmg Huid cxactK- as in tlie Skunks . . .'" There
are two pairs ot abdominal mammae m tlie temales.
Skull (tigs. 16 and 17). The specimens in the Bntisli Museum sliow. irrespective ot
age, a wide range of size and some variety ot detail. All, however, are very solidK'
built; the sutures fuse at an early age, and in tully adult skulls quite literally leave
absolutely jio trace of independent bone structure. The braincasc is relatively rather
broader than m the dogs. There are slight, pointed postorbital processes but little iji
the way ot interorbiral constriction and usually not much postorbital narrowing, the
trontal aspect riuming almost parallel-sided from the temporal region to the rostrum.
The last is ver\- short and broad, the nares ^videly open. The same applies to the posterior
narcs, the intervening nasal structure being Imely complex ijidicating a highly developed
sense ot smell. The zygomatic arch is strong, the glenoid fossa deep. In fully mature
skulls there is a slight sagittal crest which joins a very marked, though shallow, supra-
occipital crest. The whole posterior aspect ot the skull is one ot firm solidity of build,
including the foramen magnum and its adjacent condyles. The anterior palate is short
and broad: posteriorly it continues, parallel-sided, well back of the toothrows. The
bullae are large though not highly domed, and like the rest ot the skull extremely
strongly built, being fused posteriorly to large paroccipital processes. The mastoids
arc well-developed and prominent.
MnLMVt)liA
"5
Fig. 17. Mcllivora capcnsis: skull, B.M. No. 26. X. 7.1, j, x \ ; palatal & d
orsal views
The dental formula is jyyy = 32- The teeth seem often of rather irregular develop-
ment, one or more being exceptionally small, set at an unusual angle, or lacking.
The type o( si(^iinta has a second lower molar on the lett side but not the right. The
carnassials appear to be slow to erupt; and in several skulls, even with wcll-ossified
sutures, two sets of teeth exist, what is presumably a very adult-looking milk dentition
being replaced by the fuial set. The cheekteeth are in the young skull very sharply
cuspid; but, for an animal reputed to live to a great extent on relatively soft food, all
the teeth seem to wear very badh'. The incisors are often pretty irregular; but when
properly developed form, top and bottom, a very compact, strong cutting unit, the
M(' Till ( AIINUIIIU s Oi WrST A I R 1 r A
upper OIK'S being ei>n>Klcralil\' Luger tli.m the lower, the outer pau" iji eacli case beijig
somewliat bigger than tlic others. Though it is not always detectable, in the lower jaw ii
is bitid, i-2 tritid, and 13 generallv unnotched, though it occasionally shows faint traces
ot trifidv. The canines arc stout but. tor carnivores, relatively short.
Habits. One ot the foremost characteristics of the ratel is its extreme bravery and
general toughness. It appears to be quite without fear, and when flight seems of no
avail will turn savagely to attack man or anv other creature. It can take any amoiuit of
punishment and is so tireless in combat that it has been known to e>draust and overcome
tar larger .animals. Indeed, one is said to have killed a buttalo in the Kruger National
i'ark. One ot the things that makes it so ditiicult tor dogs or anything else to deal with
is the e.xtraordinan.' tougluicss and looseness ot its skin. The tirst makes it well nigh
impossible tor teeth to penetrate; the second enables the ratel. in spite of being held,
to twist and tuni so readily that it can inflict a fierce bite on any aggressor that has
seized it in hand or mouth. Sikes (1963) tound that the only sate grip was by the skin
on the back ot the head; anvwhere else, including the scrufl of the neck, was highly
dangerous. How impenetrable the skin is is illustrated bv an accomit given by T.
Rawson-Shaw (in Fleetwood, 195N) in which he relates t];at blows from a matchet
■"which would have cut any other animal of that size in half merely bounced ofl",
leaving a shallow gash on his hide, and it took about ten of these and tour .22 bullets
to kill it".
Others have tound much the same; and it is commonly held that a direct shot m
the head trom a tairly powertul rifle is the onlv certain way ot killing a honey badger.
In the Bauchi area of Northern Nigeria it was well knowii that arrows and spears were
almost useless and that the best, indeed reputedly onK', way ot certainly killing one ot
these animals was to club it over the back ot the head. As opposed to these views the
plain tact remains that though a tew are damaged a very high proportion of the skulls
m the British Museum have perfect craniums. In tussles with dogs it is usually the ratel
that succeeds in sinking its teeth into its opponent, hanging on tirelessly, with jaws
clenched like a vice, despite being banged on the ground or against trees or rocks, and
tmally, ^\hen the dog is completely exhausted, makhig ott apparently none the worse
tor the experience. Sikes (1964a) tound that, m pla\', a captive ratel being swiuig
round hanging on to a sack actually appeared to enjov being bumped roughK up and
down on the ground.
Caught young, m tact, these animals take very kindh' to captivitN and become not
only docile but activcK- triendK' (Sikes, 1963 and 1964a; Hoesch, 1964). They are
attectionate and very playtul, even inventing games (Sikes, 1963). They recognise
tnends and voices, distinguishing various tones in sumntons, command, warning or
reprimand. Hoesch's pet used to nibble in a restrained tashion at hands or clothes m
triendship. But in captivit)', as 111 the wild, ratels can with some abruptness work
themselves up into an ecstatic tury. During such a bout the hair stands on end, the
.mimal toanis at the mouth and becomes almost literally blind to any external calming
influence. Such moods, however, quickly vanish. Sikes (1964a) supposed that these
rages, which ot coiu'se are an invaluable reaction to aggression in the wild, w^erc
brought on or auirmented b\- unusualK' larije releases ot adrenalin into the blood
MELLIVORA II7
Stream. Hoesch (1964) found liis captive animal to be clean in its habits. Sikcs (1964a)
observed hers to urinate or defaccatc into a suitable hole, which if not of approved
size or shape would be industriously altered. The droppings were rarely covered. The
scent emitted by ratels affects different people in different ways, some finding it
extremely repulsive, others (Sikes) after a time not unpleasant. All agree that it differs
from the ordinary musky smell of other mustelids.
Ratels are mostly solitary animals but may at mating time hunt in couples, or a
mother may be seen with her two young ones. When not active in the search for food
they shelter in holes, either in the ground, in tree trunks or logs, or amongst boulders or,
in suitable country, in a cave. Earthen burrows may be dug by themselves, this being
one of the functions of their powerfully clawed forefeet; but, like so many other
subterranean dwellers, they make full use of aardvark holes, warthog dens or termite
mounds. No accoiuit has beeai given of the extent or nature of these underground
homes, whether they have side passages or any kind of lining. No one appears ever
to have attempted to dig out a ratel; and, mdced, it can be gathered from the previous
paragraphs that this might well prove a hazardous undertaking.
The ratel is generally recorded as being purely nocturnal, a reputation doubtless
deduced from many a midnight raid on fowl houses and the fact that it seems to be
relatively rarely encountered by day. However, not unusually it is active at dusk or at
da\vn ; and in certain conditions it is undoubtedly on the move in full daylight. Probably
these animals suit their habits to prevailing circumstances. In areas where they are
liable to be much harassed by man they make constant use of the cover afforded by
darkness; but in remote districts of low human population they probably become
bolder. Certainly, captive in zoos, they fmd no difficulty in adopting a tully diurnal
habit; and their known behaviour in coimexion with honey seeking, detailed later,
would seem to postulate pretty regular daylight activity.
When on the hunt ratels move at a steady jog-trot with the fore toes characteristically
turned in. Tliis at best is not so fast as a young man can rmi, and it has been often said
that they can keep tliis pace up tirelessly; but this is notliing more than a guess hazarded
on the leisurely nature of the gait and of the known general stamina of the animal.
No one has, in fact, ever followed a ratel very far; as soon as it becomes aware of
being chased, as it soon must on open groimd where it can itself be seen, it makes for
cover in undergrowth in which it is quite impossible to keep track of its direction.
Nor have any experiments been carried out with marked animals to determine,
without continuous observation, how far they travel. During the course of its hunting
or other excursions a ratel defmes or redefines its territory by means of its scent glands,
squatting at intervals to press its anal region on the groimd or against strategic trees
or rocks. As no specific field investigation of the ratel has as yet been carried out the
extent of the hunting range is unknowii.
Despite the name honey badger these animals are by nature essentially carnivores,
and the pursuit of food therefore concerns itself a good deal with flesh of various kinds.
Almost certainly a hungry ratel would take any sort that came conveniently to hand,
including carrion; but its normal range of food embraces small rodents, birds, eggs,
insects, lizards, tortoises and frogs. Some, at least, of the rodents it may dig out from
IlS THE CAUNIV01U;S Of WhST AflUCA
their burrows — gcrbils, groinid-squirruls aiul otlicrs; the birds arc sucli as (.iwell and
nest upon the ground or pay it trequent visits: trancolins, bustards, plovers, quail,
doves and so forth. Frogs, too, arc sometimes dug up: Angus Buchanan (1926: 251),
writing ot the race ot Mclliuora from Air named after him refers to "a regular warren
ot holes scooped out in the night by a ratel 111 search ot dormant trogs buried in the
sand a toot or two beneath the surface"".
Ratels have sharp, backwardly-pointing papillae on their tongues which enable them
to deal etlectively with tough foods. Tortoises present no difficulty to this animal
which can readily crush the shells with its very powerful jaws. Snakes also arc taken,
including highly poisonous ones; and that they are not always of small size is shown
by an extraordinary record in Aliican IViU Life, 1964, 18 : 37 which tells ot a ratcl
tighting, killing and feeding on a python some lo or 11 feet long. The begiiuiing of
this exceedingly noisy and energetic combat was not seen, but the battle must have
continued tor some half an hour, at the end of which the snake "was so mutilated that
it looked as it it had been run over by a train". The ratel has, at the Asiatic end of
its range, earned a reputation tor excavating yet another kind of food: inadcquatelv
buried human corpses. Though doubt has often been cast upon this, Pocock (1941 : 465)
says that the truth ot the belief has been proved beyond doubt. Like other small carni-
vores ratels welcome vegetable food such as berries and other fruits, roots and bulbs
as a regular part of its diet. They inquisitively lift stones or tear flaking bark from
trees.
The ratel's partiality for birds very often leads it into becoming a pest in farmyards
or private compounds where domestic fowls are kept. Such is its strength and per-
sistence that it is difficult to keep out. It has been recorded (Fleetwood, 1958) as ripping
thick planks from a strongly built hen-house, or burrowing beneath stone foiuidations.
A ratel is known to have climbed the mud wall of a hen-house to a small window
covered with wire netting fastened down with staples, ripping this obstruction aside
to gain entr)'. Another, unable to get in in any other way, tiumelled under the wall
and up through the floor, which was paved with large stones set in mud mortar. The
sturdy, muscular legs and long, strong claws of the forefeet enable tiuuielling to be
effected speedily even in hard groimd. Once entrance has been effected to a fowl-
house it often happens that everything inside is slaughtered and eaten, nothing beyond
a few scattered feathers remaining. In one episode recorded by Fleetwood (1958)
17 Muscovy ducks were lost, and in another 36 halt-grown pullets; and the same sort
ot thing has happened to many others. Rawson-Shaw, to whom these figures are due,
reached the conclusion that a ratel may have an ancillary following of other carnivores,
such as civets and jackals. For just as the lion's kill may soon be made use ot by a waiting
band of scavengers, so other less adventurous and able carnivores may well be at hand
to take advantage of the courage, strength and determination ot this pioneer burglar.
It should perhaps be made clear that in many such cases it has been quite defniitely
proved, by shooting, trapping or other means, that it was indeed a ratel responsible
for the entry and at least initial damage. It must be added that visits may be repeated on
successive nights either until there is no further attraction or the raider has been killed.
Sikes (1963) found that in captivity a young ratel would eat a whole dove, but that
MELLIVOKA
119
i::o rm c ai!Ni\ ours (ir wr.si .\i kka
an adult requires a hill-growii towl. Skiu, liau% tcatlicrs and bones ot a vietini arc all
eaten as well as the flesh. The food is held down by the forepaws.
But the tood and the robberv for which Mcllimtra is most widciv hinicd concern
lu>ney and wild bees' nests. Since about the middle ot the i6tli century, when a mission-
ary priest wrote an account of Portuguese East Africa, ojie of the travellers' talcs that
must have circulated in Europe was ot a bird that for its own ultimate benefit deliber-
ately guided men, by song and a clapping of the wings, to stores of wild honey. It was
not luitil some 200 years later, however, that this story was extended to include the
ratel as possibly the bird's chief medium tor breaking open the nests and laying bare
mutually beneticial supplies of honey, wa.x and grubs. The story of this strange co-
operation, it not symbiosis, between animals ot othcrv,'ise quite different ways ot
lite, and wliich set the natural history world wondering tor tuo centuries, was due to
the writings ot a Swedish naturalist, Sparrman (17S6, 2 : iHo-i8i) who accompanied
Captain Cook on his second voyage to the south seas and explored the interior ot
South Africa on the way back. His historic, though certainly not reliable, account
make both amusing reading and a good starting point tor the examination ot this
reputed partnership:
"The itilcl, a sort ot weasel or badger, by nature destined to be the adversary of the
bees, and the unwelcome visitor ot their habitations, is likewise endued with a particular
faculty tor discovering and attacking tliem within their entrenchments. His long
claws, besides assisting him in digging the dark subterraneous passages which serve
him for an asylum, are likewise ot use to him in the occupation he is frequently
employed in of undermining whole colonies ot bees. Now, as a man placed at the mast-
head can easiest descry a sail or land at a great distance about sun-set, so probably
this time of the dav is the most convenient for the ratel to look out for his supper;
for he is likewise said to be particularly attentive to his business about sim-set, when he
will sit and hold one of his paws before his eyes, in order to modify the ra\-s ot the
sun, so as to render them inoffensive to his organs of sight, and at the same time to
have a distinct view of the object of his pursuit: and when, ui consequence of peering
in this manner cin each side ot his paw opposite to the sun, he sees any bees fly, he kjiows
that at this time they are going strait forward to their own habitation, and consequently
takes care to keep in the same direction as that in which the\' fly, in order to tnid
them. He has besuk^, as well as the Hottentots, the Caffres, and the peasants ot the
Cape, the sagacitv to follow a little bird, which flics on by degrees with the alluring
note o( clu'ir, clicir, clwn, and guides its followers to the bees" nest . . .
"Those bees' nests which are built up in trees, are m no danger whatever from the
ratel. In the fust transports of his rage at having sought after these bees in vain, he uses
to bite or gnaw the trunk of these trees; and these bites arc sure marks for the Hotten-
tots, that a bees nest is to be found up that tree. I should myself have entertained many
doubts eonceniing all these properties attributed to the ratel, had I not obtained various
accoiuits of this curious animal, entirely corresponding with each other, from many
experienced farmers and Hottentots living in different parts ot the coiuitry".
Tlie bird to which Sparrman made reference is known as the greater honey guide
(Jihlicdtor iiulicdtor Sparrman), a representative ot a family, the liidicatoridae, com-
MHLLIVOKA
prising four genera, largely African but extending their range to Malaya and the East
Indies. Nearly all of the eleven species are known to eat wax as part of their diet;
and all, so far as is known, arc parasitic, laying their eggs cuckoo-fashion in the nests
of other birds. Some two or three species without question seem to guide, or to
attempt to guide, men to bees' nests; but whether similar behaviour could in truth be
related to ratcls as well, as Sparrman had asserted, was not so clear. It so often happens
in natiu-al history that early held beliefs or superstitions, especially when they concern
the unusual, are copied from work to work, year after year, without closer inquiry.
They make good reading. Like others, the story of the ratel and the honey guide was
incessantly repeated, both verbally and in written accounts; yet critical enquiry showed
it to be nothing more than hearsay, no wholly reliable and competent observer ever
having openly laid claim to actually witnessing the alleged behaviour. The story,
in fact, might have been on a par with the pelican pecking its breast or the porcupine
shooting its quills.
At length, however, Friedmaiui (1955), Curator ot Birds in the United States
National Museum, spent many years making an extremely interesting and detailed study
of the Indicatoridae and their habits. He came to the conclusion that while Sparrman's
account was in many respects inaccurate, the reputed association of the ratel with the
honey guide was true. He arrived at this opinion only as the result of a great deal of
sifting of old records and circumstantial evidence. No one has, in fact, ever witnessed a
complete sequence of events from the initial attraction by the bird of a ratel's attention,
through the stages of guiding, to the discovery and breaking open ol a nest. It is merely
that McUii'ora and Iiulicatorhavc been seen together in a number of isolated circumstances
that are capable of being joined together into a coherent series of events. Whether the
behaviour, either in its observed parts or presumed whole, has been correctK' inter-
preted is altogether another matter.
Briefly, what takes place, with a ratel as with humans, is that the bird, perched not
very high up on a tree, utters a great deal of unceasing chatter, climiii, chiirra, cinirra,
to attract attention; and having achieved this end flies, still chattering, a few yards
ahead and settles on another perch. The ratel responds to this chatter with occasional
grunts or growls. This is continued imtil within soimd of a bees' nest the bird becomes
quite silent. Such a course may cover a few yards or half a mile or more; and, corres-
pondingl)-, a tew seconds to half an hour. The route taken, except when extremely
short, is never direct but circuitous, meandering or even criss-cross. Having arrived
in tlie vicinity of a bees' nest the honey guide will remain silently perched quite
patiently for a very long time luitil the store of honey has been laid bare and pieces of
comb are left lying about to feed on.
This is no place to discuss in any detail the alleged guiding habits of Iiidicalor except
in so far as may be necessary to draw attention to the fact that the apparent co-operation
between bird and manunal is not to be lightly accepted at its face value. The uncritical
animal lover may all too easily follow others into a trap. When this apparent partner-
ship was first reported it seemed to be nothing other than a case of an intelligent and
hungry bird discovering a store of honey, seeking out a better equipped ally, and
deliberately c.xcituig this essential participant to curiosity and leading him or it to the
THE CARNIVORES OF WEST AFRICA
mutually desirable spoil. All of this is very nnich open to question. It has been amply
shown that hiuiger is not tlic operative excitant snice birds exercise the guiding ritual
even with tull stomachs. It is very possible that there is, indeed, no deliberate seeking
out ot a helper but merely that the chance sight of certain animals leads to an agitated
reaction in the birds. The complex path taken to reach an apparent goal suggests that
no specific goal was actually in sight at the commencement but that by criss-crossing
the bush over a sufficient distance a nest is eventually hit upon. It is a common fact
that in much ot the African woodlands there arc plenty of wild bees of one knid or
another and tliat by quartering the bush it is not excessively difficult even for a human
being to discover some sign ot a nest in a tree or a bank. There may, therefore, be no
true guiding, the expedition being, in fact, little more than a voyage of mutual dis-
covery.
If the bird stimulates the ratel to action with its chatter, tlie ratel in its turn keeps
the bird in a state of excitement by responding from time to time with its deep growling
grunt. African hunters who know this utter the same sound with a view to keeping
the bird interested. Guiding does not necessarily take place to a nest well stocked with
honey; it may be to an almost new one; and it is knov^ai that a long-deserted nest
containing good stores of honey holds no attraction for the honey guide. It is the
sight or soiuid ot numerous bees that brings the bird to a halt and causes its chatter to
cease. This is in the vicinity of, rather than actually at, the nest; but the assertion that
the bird will, if necessary to an unintelligent helper slow in detecting the desired honey,
go further and point to it with its bill is, as might be expected, nothing more than a
picturesque embellishment.
These arc some ot the many aspects ot so-called guiding brought out by Friedmann's
classic study, which make it clear that it is not the simple purposetul act that early
naturalists assumed it to be. A tew other points may be glanced at in conclusion.
Firstly, it is interesting to speculate on how this parmership between bird and beast
evolved; what can have been the original stimulus, and the route by which it came to
its present state. The co-operation serves no vitally essential purpose as it does in
true symbiosis; honey badger and honey guide can each flourish without the assistance
of the other. There are, indeed, signs that the habit is, troni force ot changing circum-
stances in modern Africa, on the decline. We are, in this present work, strictly concerned
only with the habits o{ Indicator iudiccitor in relation to the ratel; but it may be briefly
added that it is possible that the bird occasionally leads baboons to honey, and may
make, apparently wholly unsuccesstul, attempts to interest other animals. A caution
should perhaps be given that the course taken by the bird may not always lead the
follower profitably to a bees' nest; for more than one man has foimd himself suddenly
face to face with a dangerous animal. Such an occurrence, once regarded as deliberate
on the part of the bird, is most probably a purely accidental outcome ot quartering
the bush in its search for a concentration of bees. Finally, for those who may tuid
themselves in proximity to a noisily excited bird in the grass-woodlands, the greater
honey guide is of moderate size, some 120 to 150 mm long with an additional tail of
about 60 to 80 mm. Its coloration is variable with age, sex and other factors but in
general is dark drabby or olive brown above, the wing coverts and some of the tail
MELLIVORA 123
feathers white-edged, and with a small chrome-yellow bar on the shoulder. In the
young the breast is bright yellow.
Sparrman asserted that the ratcl was quite unable to climb and that bees' nests in
trees were therefore unattainable. This is not strictly true. A ratel may not be able to
climb up a siurfacc that others no foothold, or very far up a vertical smooth-barked
trunk; but given a reasonably rough surface, and particularly one at a slight slope,
such as so many open- woodland trees have, it is a pretty competent climber. A mud wall
or a coarse-barked tree can be scaled with some ease ; and any African bee farmer who
fails to set liis hive sufficiently high and along a sufficiently narrow branch is liable to
fuid it robbed. Since a fair proportion of wild bees' nests are made in hollow trees this
ability to climb is useful if not important. Ratels can climb wire-netting or expanded
metal without difficulty, bite through wire, scratch ajid gnaw to pieces strong posts,
and burrow through hard earthen or wooden floors so that a cage to retain a captive
animal must be of very solid construction.
Little is known of mating or breeding despite the many specimens which have been
kept in zoos. The gestation period, not very accurately determined, is said to be about
six months. There are commonly two yoimg at a birth, bom blind. They are moved
from place to place if necessary in the mother's mouth. They utter a plaintive whme.
Details of development are unknown ; but there are two or three records of ratels
having lived in captivity for approximately 24 years. Hoesch (1964) found his tame
animal to drink only rarely; but it used its drinking bowl for keeping cool.
Taxonomy. Since its fu-st alleged variant was described in 1792 M. capciisis has
been something of a favorite for the naming of subspecies. G. M. Allen (1939) lists
II currently recognised for Africa, and Ellerman & Morrison-Scott (195 1) 5 more
for Asia. No less than 5 races are, indeed, said to occur in West Africa. The points
v^'hich have been taken into consideration in the diagnosis of races are the extent of the
pale dorsal colour; its degree of whiteness of greyness; the annulation of or absence
of it from, the hairs; general size. Differences based on these are often clear enough in
single specimens; but whether a variation has any constancy throughout a population
and hence constitutes a valid local subspecific distinction can be established only by the
existence of adequate material. Of the 12 West Africaji specimens available in London
from which to confu-m the validity of 5 reputed races, 2 have no skins and are therefore
racially indeterminable; and of the 10 remaining, only in three cases (Air, Tarkwa and
Abenasi) is there more than a single specimen from a given locality. Of the 8 specimens
which have skulls, only 4 can be regarded as mature; and of these, 2 are badly mutilated
and incomplete. From such material it is quite impossible to reach any imequivocal
conclusion.
In the matter of pelage pattern, it is dirticult to believe that Thomas & Wroughton's
concisa, with merely a small area of the white lacking from the lower back, is anything
but a minor individual variation, the more so as signs of a comparable reduction is
to be seen amongst skins from other regions. The possibility, though unlikelihood,
of this occurred to the authors; but they held, if such did prove to be the case, that its
smaller size was alone sufficient justification for their proposed race. As the type of
omisa is a young male w ith quite unworn teeth, the sutures not fully ossified, and no
124 THE CARNIVORES OE WEST AFRICA
development ot crests, size comparison is scarcely a reasonable basis for argument.
Size plays the major role, too, in Thomas's huchanani; but the type of this is quite
juvenile, with unfused sutiurcs, teeth not tully erupted with some of the milk dentition
still in place. It is true that Thomas cited a paratype which, though described as a
fully adult female with worn teeth, has skull measurements (as given by Thomas
since this skull is no longer available for assessment) that differ only minutely from
those of the juvenile type, as shown in the table on page 127. It is, ot course, possible
that the ratels of the inhospitable Subdesert. feeding largely on frogs and lizards,
are truly of a smaller size; but argument based on present study material is not altogether
convincing.
It is tempting, and not entirely unreasonable, to suppose that differences in the grey
dorsal patch have little to do with locality and are nothing more than the sort of
individual variation that one might expect in a pelage of this kind. But, looking
outside West Africa to districts from which a much wider range of study material
exists, there docs often seem to be a strong overall uniformitv in pelage coloration
from a single area — as, for example, in the dozen skins from Somaliland. The argument,
however, is weakened by the frequent ability to match patterns from widely separated
regions, a striking example of this being a skin from Suakin on the Red Sea (No.
4.8.2.27) having a verv distinctive pure white flank stripe precisely similar to one from
Namaqualand in South-west Africa (No. 4.2.3.54). The hiicliaiiaiii skins vary somewhat
amongst themselves; and when the reputedly mature female is put with the Somaliland
specimens it is seen to differ very little from them except for size.
The situation might he explicable by the postulation of ecological rather than geo-
graphical races. This brings us to the consideration of cottoiii, the all-black form. It has
been suggested, and is very possible, that this is nothing odier than a melano which,
in the nature of these things, might turn up in any population. Yet the fact that the
onK- 5 knowii Mcllivom skins from Ghana are all of this type miglit suggest that the
black coloration has some kind of racial connotation rather than that ot an occasional
mutant. The form was originally described from the Ituri forest, over 3000 kilometres
away on the other side of the continent; and it is also known from Cameroun. The
factor which these three regions have in common is high forest; and it would seem that
cottoiii might, indeed, be an ecological form typical of wet, closed-forest conditions.
Yet this, too, would appear to be negatived by the fact that of Bates's 4 Cameroim
specimens, 3 are all-black and 2 have very white backs presumably typical oi Icnaviota.
It is true that onlv one of these four specimens, a white one, is associated with a definite
localit\', Bitve; but practically all of Bates's mammalogical collecting was carried out
in a restricted area of high forest. It is recorded also (J. A. Allen, 1924) that partly white
forms (which, incidentally, appear from photographs to be excellent matches for the
West Afncait signata) occur in the Congo in the same region as the black form.
Age, too, mav play its part in the question of blackness. The three black Ghana
skins accompanied by skulls are all very old animals with extremely worn teeth;
and Pitman, quoted in Shortridge (1934), expressed the opinion that "As the Ratel
gets older the white patch on the back becomes gradually darker, until in some very
old males it is onlv just distinguishable ". On the other hand, if this ^\cre so one \\ oiild
M K L L I V O H A
125
have supposed that attention would have been drawn to the fact in coiuicxion with
the several specimens known to have lived in zoos to a ripe age.
The situation, therefore, is confused. The present writer is of the opniion that many
of the variations of pelage are purely individual and may turn up in widely separated
places though there is possibly a tendency for some to be commoner in certain ecological
conditions than others. They are, thus, rather "varieties" than regional or truly eco-
logical races. They are retained herein as such and described below for what they are
worth.
Fig. 19. Mcllivom capaisis: dorsal coat patterns, showing the extent of white in tlie v.irioiis
tonns: a. laiconota, h. biicliaimiti, c. coticisa, d. sif;iialii, c. coiioiii
A table showing such measurements as are available for West Atrican specimens
will be foimd on page 127; but since for the reasons given above these figures necessarily
embrace a mixture of ages from juvenile to very old they are not reliably comparative.
The diagrams in fig. 19 will aid in the identification of the ditfereiu forms; but
there seems no reason why all manner ot intermediate patterns should not tinn up.
126 THE CARNIVORES OI WEST AFRICA
Mellivora capensis leuconota P. L. Sclater White-backed Ratcl
Tliis was originally said to conic troni "West Africa" without closer definition;
but it has since been held (e.g. G. M. Allen, 1939) to range from southern Morocco
to the tormer French Congo. No specimen definitely identified as Iciicoiiotn occurs in
the British Museum; but it is to be assumed that two of Bates's Cameroun specimens,
one from Bityc, arc this form. Though onh- the apparently smaller of the two has
external measurements (given on page 127) it appears to rank among the largest of all
ratcls.
In this torm the entire upper side troni the hice to halt way along the tail is pure
creamy white with a minimal and almost undetectable admixture of black hairs. The
white hairs may have narrow black tips, but these, agaiii, are almost invisible.
Mellivora capensis cottoni Lydckker Black Ratel
This form occurs in the liigh forest, or at its edge, in various parts from at least
Ghana to the north-eastern Congo. The known West African specimens all come from
Ghana: Tarkwa, Abenasi and Oda.
As its name implies this form is topically entirely black though some skins liave an
extremely small number of scattered white bristle-hairs only detectable on close in-
spection. But there is one exceptional partial skin from Abenasi in which there is an
even scattering of white or white-tipped bristle-hairs throughout the whole of the
deep brown, rather than black, dorsum. The coat is thin and harsh, but the whitish
hairs become rather denser over the top of the head.
Mellivora capensis concisa Thomas (^' Wroughton Lake Chad llatcl
Only one specimen of this is known, from Lake Cliad, but it is almost certainly only
a minor variant of the ratel that occurs eastwards throughout the Sahel and Sudan
zones, as tar as Somaliland and there called hrockmani Wroughton & Chccsman.
The dorsal coat consists of a preponderance of very long pure white bristle-hairs
amongst long fine black undcrfur and a minority of black bristle-hairs. The feature
which is held to distinguish concisa is the fact that, for all practical purposes, the white
bristle-hairs are entirelv lacking from an area starting on the spine at about the small
of the back and broadening a little to the tail, which is also all black.
Mellivora capensis signata I'ocock Speckled lUtel
The t\pe, from an unspecified locality in Sierra Leone, via the London Zoo, is the
onlv specimen known in the British Museum; but according to Monard (1940) tlie
race occurs at Catio, Portuguese Guinea. The characteristics of the type are that although
the pelage is the normal dense white over the crown, the pale colour starts to thin out
over the neck and shoulders, and continues thence to the rump only as scattered white
"ticking " amongst essentially black fur; and also that the white in this part of the coat
is not due to wholly white bristle-hairs, as in other forms of ratel, but to subterminal
white anniilation of otherwise black bristle-hairs. The "ticking", already scanty.
MELLIVORA
127
tadcs out entirely just torw arc! of the rump, the whole hindquarters and tail being
completely black. The skull has an extra lower molar (nio) on the left side.
Mellivora capensis buchanani Thomas Air Ratel
The question of the small size of this form, known ojily by three specimens from
Elmcki River and Tarrarc River in Air (Subdescrt), has been discussed above. Apart
from this matter of size it is doubtful whether this form differs essentially from the
Sahcl and Sudan zone ratcls referred to above under coiicisd. It is a little whiter over the
head but this may be due to the young age of the specimens now to hand. For the
rest, it is essentially a grey-backed form, the pelage being an intimate mixture of long
pure white bristle-hairs amongst fine long black undcrfur and a fair proportion of
black bristle-hairs. The white mixtmc spreads to the basal part of the tail, one of the
specimens having a small white tip as well.
Tabic 7: Numerical data for Mcllirora capciisis
hucliaimiii
atiicisa
siiiimia
(Type &
kiicoiiota
colloni
(Type:
(Type:
paratvpe :
(Bitye)
(Ghana)
Lake Chad)
Sierra Leone)
Air)
Vegetation
Forest
Forest
Sahel
?Forest
Subdesert
Number in mean
I
3
I
I
2
Condylobasal length
—
142-3
134-8
125-8
toS-5
Basilar length
—
129-8
120-7
120-1
97-2
Palatiiar length
(67)
59-1
57-5
52-2
46-3
Zygomatic breadth
S7-0
8i-i
71-0
66-7
62-3
Upper cheekteeth breadth
49-1
49-2
42-y
44-0
38-6
Nasals, length
25-2
—
::3-4
23-7
21-6
Interorbita breadth
36-7
35-6
30-5
29-6
29-2
Postorbital constriction
30-3
30-5
32-1
28-5
30-2
Braincase breadth
66-3
58-:
58-5
52-8
Toothrow [c — nO-)
41-8
40-7
38-8
33-5
32-5
p^ length
14-4
13-5
12-6
13-0
II-O
"ji length
4-3
4-1
4-3
4--
4'i
"H length
15-4
(i3-y)
14-4
13-0
I2-I
Head & body
S40
736
590
4S9
Tail
240
240
156
—
140
Hindtoot
135
120
95
—
85
Ear
40
3.S
ty
—
29
RATIOS (per cent)
-
Tail/head & body
29
33
26
—
29
Zygom. br./condyiob. 1.
—
57
53
53
57
Braincase/condylob. 1.
—
47
43
47
49
Braincase/zygoni. br.
—
82
82
88
85
Palatiiar l./condylob. 1.
—
42
42
4^
43
Intcrorb./postorb.
121
116
95
103
97
p^jc—m^
34-4
33-1
3-'5
38-8
33-8
p^iiA
335
3^y
-94
310
262
128 TUF. (.AKN'IVUKI.S Ol WEST AI-RICA
This is one ot the tew specimens with any field notes: accorduig to Buchanan it
liiirro\v>; tor deeply buried frogs, the stomach being found to contain remanis of
these and ot li/ard'-.
Siibtainilv LUTRINAE Baud. iSsy
Otters
Distribution and general. The third and last West African subfannly of the
Mustelidae has a wide distribution not only throughout much ot Atrica but also in
North and South America, and in Asia to southern China, japan and some of the
larger East hidia islands. Five or six genera, covering a score ol species, are recognised,
all ot them at least semi-aquatic, mostly riverine or lacustrine, but one, liiiliyilia
Fleming, marine in the northern Pacific Ocean. Two of these genera occur in Afric.i.
including the region dealt with in this work.
Vegetation is of little moment to the otters. They occur m all kinds, their mam
criterion for existence being the availability of ample stretches ot water harbouring
adequate supplies offish and other food materials, and preferably fringed with enough
plant growth to atford secure cover. This last, however, is not absolutely essential,
tor they are sometimes tound along relativelv open stream-banks. Otters, thus, mainly
trequeiit rivers, lakes, swamps and. since they do not object to brackish water, estuaries
and tidal creeks. This does not mean that they are to be foiuid everywhere in such
situations; for they are for the most part shy, generally confining themselves to quiet
localities little frequented bv humans. For this reason there are large stretches ot poten-
tially suitable water quite devoid of these animals, hi West Africa otters are known to
range trom Senegal to Cameroiui and from Lake Chad to the mangrove swamps.
They appear to be commonest in the rain-torest, but there arc not a great many study
.specimens trom which to draw more than very general conclusions regarding either
abundance or distribution. More than one species may exist in a single locality.
Yet though they are largely associated with and mainly dependent upon water it
must not be thought that they are in any sense wholly aquatic. Otters, indeed, spend
a good deal ot their lives on dry land, coming ashore to eat, to sleep, to detaecate or
urinate, to breed or merely t(i indulge in play. More surprisingly, they occasionally,
and tor no very obvious reason, travel long distances overland and may be unexpectedly
come across many miles from water, sometimes at night picked up by the headlamps
ot a car.
Description. Otters vary a good deal in size, hi West Atrica they may range trom
.1 weight of about 30 kg to one of five or six times as much. Similarly the head & body
length may range trom some 60 cm to nearly 100 cm with, in addition, a tail of roughly
halt these lengths. But even the largest ot these is small in comparison with some extra-
liinital species, one trom the Amazon basin being said to attain a weight ot nearh'
3> kg. The very short-legged body is long and beautitulK' streamlined in shape,
increasing gradually in diameter trom head to hips. It is intensely muscular and solidk
built but extremely lithe, hideed, the extent to which an otter can, and regularly does,
lU'x its bod\' 111 all directions must be seen to he belie\ed.
LUTRINAn 129
The head is broad, sliort-inuzzlcd and markedly flat; the neck ot approximately
equivalent diameter. The facial vibrissac are an extremely pronoiuiced tcature, long,
strong and abundant. There is a tuft below the chin and particularly large tufts on the
lower checks. The rhinarium differs in size and shape in the various species, being quite
diagnostic (Pocock, 192 rb) ; the nostrils arc usually widely flared on land but arc capable
ot closure on submersion. The roundish eyes are of moderate or even small size;
yet, by their wide-open brilliance and large roiuid pupils, tliey contrive to be a very
conspicuous feature, even at night when they pick up and reflect ligiit from the lapetiiin
liicidiiiii — in varying colours, ruby-red, cmcrald-green or amber according to species
(Harris, 1968). The oval ears are small, set well down and back on the side of the
head, and are devoid of any bursa. They, too, can be sealed by the closure of two lips
diu^ing diving. The tail, in hunting circles usually termed the "pole" or "rudder",
is a very notable structure. It is roughly half the length ot head & body, often rather
more or sometimes slightly less, highK' muscular, thick and compressed dorso-ventrally
at its base but tapering thence continuously to a narrow tip, thus completing the
overall streamlining. The short legs arc powerful, the paws terminating in five digits,
both front and rear. These are webbed to a greater or less extent, in some species to
the extreme tips ot the digits, in others to only about half-way or less. In the most
typical otters the toes are armed with fairly powerful, sharp, non-retractile claws;
but in a few, including two of the West African species, the so-called clawlcss otters,
there is nothing beyond small vestigial nails, and even these arc absent from the majority
of the digits. The palmar and plantar pads arc various in shape, sometimes rather poorly
develofied (Pocock, 1921b). The hind teet arc always larger than the toreteet.
On the uiaderside of the tail, near the root, there arc paired scent glands which
abimdantly exude a quantity of milky fluid with a powerful musky odour but not so
repulsive as that of some other mustelids such as the polecat. Their purpose seems to
be rather for recognition and the demarcation of territory than tor dctence. Females
may have, at most, three pairs ot mammae, abdominal and latero-pectoral, but some
ot these are often lacking or obscure.
Otters arc noted for the character of their pelage. The tact that this is dense and
handsome has given it some commercial value, both locally and in the world fur
trade, with consequent adverse effects upon the survival of various species. The colour
differs, ot course, in the difterent species but in general is, dorsally, lighter or darker
scpia-browii, though it is in some cases distinctly pallid, and pure white areas occur in
others. Albinos are known. The dorsal fur is sometimes slightly "trosted"; and it
always, even when quite dry, has a sparkling sheen. This is due to the composition ot
the pelage, which is remarkably constant throughout the genus. It is short, of miusually
uniform length over the entire body, as much below as above; very soft to the touch
and exceedingly dense. The hairs are of two kinds. The short, crinkled underfur,
about 8 mm in length, is extremely closely packed and is of outstanding slenderness,
being, at o-oi mm diameter, amongst the finest ot all hair. Despite this, its surface,
seen luider high magnification, is rough with long and relatively coarse scales. In the
majority of cases it is white for most ot its length but becomes lighter or darker brown
near the tips, so that if an otter's coat is rubbed up the wrong way it is seen to be
130 Till ( AliNlV01)l:S Ol «EST AfKlCA
wliitc-bascd- latlKT surprisingly smcc tlic extreme densit\ ot the tur iiornialK' com-
pletely obscures what lies below the closelv-packcd tips.
Mixed amongst this crowded underfur stand much longer hairs, by comparison
widely spaced yet in sum forming a continuous coat that entirely overlays and conceals
what lies beneath. These bristle-hairs, which arc i6 to iS mm long, are very slender and
whitish in the basal halt so that they are there indistinguishable by eye from the under-
hir; but beyond the reach ot this latter they expand into a thick, usually densely pig-
mented blade. This is not concavelv guttered as in some mammals but has a slightly
convcxiy oblong section: and though by comparison with the rest ot the tur relatively
broad, it is. in fact, only about o- 1 mm wide with a length ot some 6 or 7 mm, tapering
terminally to a sharp tip. It is these blades that glisten and give the coat its characteristic
sheen. Because of their long slender stalks the hairs have nothing harshly bristly about
them, hi the newly-born otter, at least in so tar as the West Atrican Liitia maculicolli^
is concerned, the pelage is quite ditterent. There is no sign at all ot dense undertur, the
coat being almost entirely composed ot long, straight, rather wiry hairs. A somewhat
older specimen ot Aoiiyx aipciisis shows the development of the dense coat well under
way but the ultimate outer cover of glossv bristles onh' beginning to be visible amongst
the. as yet longer, wiry baby hair. It is possible that the reluctance ot young otters to
take to the water may be due to the inadequacy ot their protection against cold.
The composition of the belly tur is precisely similar to that ot the back; and, what
is more unusual, so is that ot the tail too. It has been said that this pelage is waterproof;
but though this is not absolutely true, it is a fact that the rightly packed underfur renders
Its penetration by water extremely difficult, and the hard, glossy bristle-hairs, though
they bimch together when wet, arc very rapidly dried out. An otter gets rid ot the
water in its coat not so much bv vigorous shaking, as a dog does, but more by rolling
or nibbijig against a dry surface. This is possibly because its legs are too short to give
itselt a really effective shake. It may be added that the skin ot an otter is tough, rather
like that ot a ratel; and similarly it is also prettv loose so that the animal can twist and
turn dangerously if held by it.
Skull (tigs. 20, 21, 23 and 24). There is some ditierence ot style as well as ot size
between the three otters ^ylth which this book is primarily concerned. There are also
variations due to age and sex, chietly to do \\ ith development of the crests. Detailed
description is reserved tor the accounts ot the different animals given later, the tollowing
being some ot the more general features of the lutrine skull.
The typical otter skull has a broad, flat bramcase which contrasts strongly with the
narrow, often parallel-sided mterorbital-intcrtcmporal region. The muzzle region,
^ltuated anterior to the orbits, is extremely short, being even more abruptly truncated
than 111 Xkllivofii, the anterior nares \yidely open. In the most typical otter skulls there
are prominent sharp postorbital processes, the posterior ridges ot whichjoin the sagittal
crest, helping to outline in this interorbital area a tairly well defined pentagonal plate.
In tully developed males of most species there is a well-developed, sharp sagittal crest
extending from the postorbital region to the equally pronounced, upcurved supra-
occipital crest, which continues as sharp ridges round the sides of the braincase forward
to the very large mastoids. In females, though the supraoccipital crests may be present.
LUTRINAn 131
the sagittal line generally carries nothing more than a very low ridge which in really
old skulls may rise to a brief crest posteriorly. The zygomatic arches arc broad and
strong; the maxillary zygomatic process divides clearly into two branches which enclose
an exceptionally open intraorbital canal, obviously transmitting muscle as well as
nerve, nothing at all similar occurring in other West African Canoidea. The palate is
continued, parallel-sided, well beyond the back of the toothrows; the bullae are
unusually flat and elongated, relatively insignificant. The bone structure of old animals
is dense and solid, fusion of the sutures leaving little or no trace of original independence.
The dentition is often very irregular through the loss, or whole or partial failure to
develop, of a nimiber of cheekteeth. These, when all arc present, are jr^, giving a
total for the whole mouth of 36 teeth; but this not uncommon deviation from normal
should be kept 111 mind since it may give rise to some doubt or difficulty in the recog-
nition of lutrinc skulls. The most frequent tooth to be lacking is the very small anterior
upper premolar (pi), which, when not missing is generally sited in close proximity
to the inside face of the canine, p- may also be in contact with the canine, as though,
through the great shortening of the muzzle, there is becoming less and less room to
accommodate these teeth and they are in the course of being eliminated. The exterior
upper incisor [P) is much larger than the two interior ones (i^ and /'-) ; in the lower jaw
the distinction is not so marked and the incisors as a whole are often irregular and not
in line.
Habits. Otters in Europe, since they are the subjects of regular organized hunting
for sport, have been under close observation for many centuries ; and, the world over,
they have commonly been kept as domestic, and often very intimate, pets. Many
books and articles have been written on this latter aspect, especially in recent years,
and a good deal, therefore, is known of the behaviour in captivity of several different
kinds of otter. Yet there are often unwelcome gaps in field knowledge of the more
hidden aspects of daily or yearly life; and this is particularly true of the three African
species dealt with in this work, one of which, indeed, is scarcely knowni except for
skulls. But even in the less secret matters of everyday biology the views of specialists
arc sometimes strangely at odds about such things as senses of sight, hearing and smell,
body odour, the use of glands, or readiness of the yoiuig to swim. Nevertheless, there
remains wide agreement regarding broad general behaviour; and doubtless individual
otters differ from one another in their physical and mental attributes, as other animals
do, and care must be taken m assuming the general from the particular. So much,
indeed, has been written of specific individuals in relation to sometimes rather unreal
circumstances that one difficulty is to know what, in a brief account, can be taken as
representative of the class as a whole in its purely natural environment. The most
comprehensive account of otters in general, with descriptions of all the species, together
with their habits in so far as they are known, is to be found in Harris (1968). It must
be emphasized here that though in all likelihood most of the account of habits that
follows is applicable to African species these last have, in fact, been relatively little
studied in detail in the field and therefore it may be discovered that behaviour in the
three species with which this book is primarily concerned differs in certain aspects
\}1 TUF (MINIVoniS (ir WIST AIUK A
trom tin- i^ciicr.il pitturc givcji. This may be especially sd as regards breeding. Doubtless
the three species dirtcr amongst tliemsclves in this as in (ither respects; and it is, indeed,
vaguely recognised that tlie only two of the three that have been observed in lite do
live somewhat difterently, though this is more a matter ot general impression rather
than ot tirniK' established tact.
Otters are active by day or by night depending to some extent upon local circum-
stances as well as prctcrcncc. It there is much danger ot human interterence they become
rather carctui of exposing themselves to view and are more inclined to torage nocturn-
ally, particularly on moonlit nights. In any case their hunting tends to be more especialK-
.\n early morning or late atternoon occupation. However, both species and individuals
ot them ditfer somewhat in their degree ot sh\-ucss, and otters have been known to
come into towns at night or to raise families in drains near or under houses; and once,
even, beneath the floor-boards ot a living room.
For the most part otters dwell near water, most commoiiK' tresh, that is to say
rivers, lakes or inland marshes; but they are by no means averse to brackish or even
salt water and so not infrequently are to be found in creeks or estuaries or along the
coast. They sometimes travel overland to a considerable distance trom water, but there
IS no evidence that they spend long m such situations. Though they like to live near it
thev do not pass all, or even the major part, ot their lives in water; they mostly take
to it primarily tor the business ot seeking tood, but this vital business, as will be seen
later, is tempered with a good deal ot sportive activity. Sleeping, breeding, detaecation,
urination, and nearly always tceding are essentially dry-land occupations, as well as
exteirsive periods ot out-ot-water play, and it will thus be seen that, though some species
are more acjuatic than others, otters m general spend at least as much time ashore as
they do in the water. It is customary to reter to the males as "dogs" and the females as
"bitches"; but in spite ot this analogy with dogs the young are generally spoken ot
as "cubs", not "pups ', though the term "whelps ' is sometimes used.
On shore, an otter centres itselt on a set shelter tor the purpose ot sleep or breeding.
This is nearly always subterranean but may occasionally be in dense vegetation. These
dens, or "holts" as they are technically termed, may be self-constructed or taken over
from other animals or adapted trom naturally-occurring holes suclr as those amongst
riverine tree roots, amongst boulders, or even in a cave. Generally they have an under-
water entrance — but extremely little is known of the exact nature ot these homes in
Africa. It at least seems highly likely that the African clawless otters would find some
difficulty in digging except in the softest of swampy mud. The shelters are situated
well above flood or seepage level and are frequently lined to some extent with vege-
table matter, grass, leaves, reeds or moss; tor, although they spend a good deal ot time
in water, otters like very much to be warm and dry. 13rying ofl ot the outer coat,
by rolling or rubbing, is in tact one ot an otters tirst urges on leaving water. Custom
in nest-lining, however, varies; pcssibly when the soil itselt is of a warm and dry natm-e
no nest lining is called for. Such conditions would seem to exist over much of tropical
Africa, except possibly in the forest belt at the height of the rains.
Holts are occupied sometimes by a lone otter, sometimes by a tamily party. Such
parties usually consist ot a bitch and her offspring up to the subadult stage; the dog otter
LUTRINAE 133
is, as tar as the holt is concerned, generally kept at a distance, and occupies a separate,
though neighbouring, shelter.
Some observers have maintained that otters combine and co-operate in fishing
parties; but it is doubtful whether such collaboration, if it exists at all, ever extends
beyond the immediate family. Most otters are by no means gregarious creatures and
holts are generally fairly widely separated; and it would seem probable that each unit
has its own recognised stretch of water, especially when one remembers the powerful
scent-glands which otters possess and their probable use for demarcating territory.
Yet a fixed territory implies some degree of permanence, and otters have a considerable
reputation as wanderers, never dwelling long, perhaps only a night or two, in one
place except at actual breeding time. Such nomadism or semi-nomadism may be for
safety; or it may be that after a while a given stretch of water begins to become ex-
hausted of food, or the fish, by constant hunting, accentuatedly wary. Whatever its
motive, this continual moving renders the field study of otters a matter of some
difficulty except at breeding times since the observer often does not know where to
fuid his animals from one day to the next.
It is commonly thought that otters feed solely upon fish but this is far from true.
Fish may form a large or even the major part of the diet but a good many other things
living in or around the water are eaten, such as crabs, crayfish, prawns, mussels, snails,
large water-beetles, and, on land, groimd-birds, water-fowl, eggs, rodents and possibly
other small mammals, lizards, including geckos, snakes, frogs, toads, grasshoppers,
crickets, mole-crickets and other large insects. These are generalities, for each species
has its preferences and each locality a differing availability of food materials. For
example, it would seem that the two commonest West African otters differ in their
feeding demands, Liitra maailicollis being predominantly a fish eater, Aonyx capensis
consuming more crabs. But no set work has been done on this or on the kinds offish
preferred by each species in West Africa, though it is known that the latter species,
at least, consumes, in Lake Victoria, a fair number of Lungfish {Pivtoptcms) and probably
clariid catfish as well. Eels, which form a considerable part of the food of otters in
Europe, do not occur in the rivers of West Africa except possibly in cstuarinc waters.
These considerations arc of some importance since it has been held in Europe and
elsewhere that otters cause a great deal of economic harm both in fisli-liatcheries and
amongst the more valuable "game" fish. Farmers join with tishermen in their con-
demnation, saddling otters with poultry thefts of which, often, they are not guilty.
Such charges have, on the other hand been strongly denied; and there is a growing
appreciation that otters may, in sum, effect far more good than harm in many ways,
not least by getting rid of much fish that is diseased or otherwise tmdesirable.
Small articles of food taken in the water, such as snails, mussels and other mollusca,
are then and there swallowed with a bite; small fish may be eaten in an upright position
while the otter is treading water with head and neck above the surface, or while the
otter is floating on its back; but anything large is brought ashore to be consumed.
It may, if really large, be held down by the paws and eaten thus on the ground; but
moderate-sized fish are held up to the mouth between the hands and eaten gradually
from end to end. It has often been asserted that a start is always made at the tail end.
134 II" <'AHN1\()UES or WKST AlKICA
tlic licad tuially bcuig discarded; but tlirs is by no means always so. Food is invariably
eaten fresh at once or discarded, never stored, and, in tact, otters are usually believed
not to eat ;uiytliing but the tood they have just caught; but Stephens (1957) records an
opinion that, at any rate in Europe, the\' will, and regularly do, consume a large amount
of carrion.
Ability to catch tish implies a superior ability to swim. The adult otter is certainly a
past master in this art. On the surface, with the head partly above water, an otter
progresses in a rather leisurely fishion, using its teet alone in a sort of dog-paddle;
but once it has dived in earnest the whole demeanour immediately chajiges and the
animal becomes a lively and remarkably dexterous hunter. Propulsion is mainly by
powerful thrusts of the hindlegs, used as a pair, aided sometimes by a sinuous and
rhythmic up-and-down movement ot the body. The forelegs arc occasionally but not
always held back against the belly. Tlie tail acts as a rudder, hi this way a moderate
speed is achieved over a short distance; but it is rather the otter's facility, with its supple
bodv, to equal or outdo its prey in swiftness oi turn that makes escape for the fish a
matter of some hazard. None the less, it seems that otters as a rule prefer to go after
the slower moving kinds ot fish; and certainly the two categories mentioned earlier
as being commonly eaten in Africa are bcuh sluggish.
The eyes are kept open dm^ing submergence and probably tunction more etticiently
luider water than on land. The ears and nostrils are scaled; but no one seems to have
determined how, when an otter opens its mouth to seize a tish, it avoids its liuigs
being swamped with water. Otters swim on their backs as well as on their bellies, and
they sometimes float thus idly as though half asleep. How faultlessly streamlined tlie
body is is well demonstrated not only by the easy progress ot the submerged animal
in rapid pursuit but, as well, by the manner in which an otter habitually slides into the
water with scarcely any disturbance ot the surface and consequent alarm tor the lish.
To all intents and purposes the only sign ot an otter's diving lies in the trail ot rising
bubbles, probably from air trapped in the fur.
Otters, being mammals, must, of coiurse, come up from time to tune to breathe
otherwise they would drown. Normally they remain submerged tor a matter ot a
few seconds; Mortimer (i9''i3), observing the speckle-throated otter in Zambia, found
the commonest time to be from 10 to 1 s seconds, and he never wimciscd a dive lasting
for more than 45 seconds, and that only on one occasion, but when they so wish or
it is necessary they can remain imder water tor some four or five minutes; and Ma.xwell
(1961) timed ari Asian otter for just on six. This, in human estimation, is a very long
time though it is brief in comparison with some other aquatic animals — a quarter ot
an hour for a manatee or a seal, and an hour or even two for some of the whales.
Though no investigation appears to have been carried out on submersion actually m
the otters it would seem likely that the mechanism by which prolonged abstention
from breathing can be achieved is much that found amongst other groups. This, in
brief, is a marked reduction of oxygen consumption brought about firstly by a very
considerable slowing down of the heart-beat, and secondly b\ a constriction ot the
blood vessels in less important areas ot the body while maintaining a full flow to the
brain. There are contributory mechanisms such as, during preliminar\' breathing.
LUTRINAE 13s
a nearly complete change of air in the luiigs, a greater ability in the blood to hold oxygen
and less sensitivity of the respiratory nerve centre to increased concentration of carbon
dioxide. Ability to stay under water has nothing whatsoever to do with extra lung
capacit)' which, indeed, by increasing buoyancy would be more of a hindrance to
diving than a help. Harris (1968) suggests that because of this adaptation to prolonged
submersion otters are diiiicult to anaesthetize; but Stephens (1957) refers to two cases
within her experience without hint of trouble. In one, morphia and chloroform were
used; in the other, nembutal.
Whatever the precise mechanism an otter can, in five minutes, swim a long distance
luidcr water; and though fishing and other underwater foraging is most commonly
carried out at shallow depths it has been shown, by the capture of otters in traps, that
at times dives of at least 18 metres are luidertaken. Ability to remain below water is
to some extent dependent upon age, young otters being incapable of as long periods
of submersion as adults. Despite outstanding mastery of water it is, indeed, generally
held that yomig otters arc most luiwilhng to enter it and, if they do, prove themselves
to be nervous and highly inept swimmers. There seems little doubt, from numerous
cye-\s itness accounts, that in most cases the mother has to teach her yoiuig both how to
swim and how to catch fish. To judge from Maxwell (1961) it would seem as if this
might well apply also to the clawless otter which occurs in West Africa; but doubtless
idiosyncrasy plays its part in this as in other matters, for Mockford (1967) fomid two
young captive otters of this species to take to water quite naturally and easily at an
age estimated to be no more than six or seven weeks. This is the more remarkable in
that Aony.x is incieasingly regarded as markedly less aquatic than other species.
It seems likely that the exceptional development of the facial vibrissae may assist
in the avoidance of rocks and other obstacles in cloudy water; and, by their sensitivity
to the currents produced by even slight disturbances, aid in the detection of moving
tish. Though both ears and nostrils are closed under water, hearing and smell neverthe-
less probably play their part too; certainly on dry land both these senses are tairly
acute, smell, to judge from the complex internal nasal structure in comparison with the
poorly developed auditory bullae, much more so than hearing. Cdours, indeed, must
play a constant role in the otter's life, as m those of most mustelids. Signals are con-
stantly being left on rocks or on deliberately gathered bunches of vegetation; and the
odour lett by otters in normal passage over the ground is strong and very persistent,
it being well known to those who hiuit these animals in Europe that hounds may
often be deceived into picking up a trail already a day or two old. The faeces, doubtless,
have their own particular odour and convey their message to others. Otter droppings,
which are most commonl)- black, rather liquid and slim)-, are technically termed
"spraints"; and it is well known in Europe that set spraintmg places arc used, some-
times over a period of very many^ years. Whether this applies to the same extent in
Africa is not recorded; but Mockford (1967) fomid young Aoiiyx in captivity always
to use dcfmite spots and, therefore, to be scrupulously clean. E)Te (1963), on the other
hand, foiuid an otter of this species awkward to keep about the house since it persisted
in registering o\\ nership in this way. E.xpericnce of those who have kept otters as pets,
as well as the abundance of spraints in the wild, indicates that dcfaecation, always
136 THE CARNIVORES OF WEST AFIUCA
sinuiltancously accompanied by urination, is frequent; and tlierc is good evidence,
troni observation and troni examination oi the nitestines, that digestion and passage
ot food through the gut are rapid. 1 lowever, tlie spraints always contain some indica-
tion ot the food taken, in the form ot undigested iish bones and scales, or chitiuous
fragments from Crustacea. Excretion is performed witii straddled hindlegs and tail
erect, and is accompanied by a constant dancing movement.
Though otters so obviously, in nature, thus constantly leave traces of their presence
or passage it is a remarkable fact that no one ot the very many who have kept them
as pets has ever expressed objection to their smell, despite the animals' sometimes
habitual use ot carpets, chairs and otten beds. In fact, otters have been recorded as
having no smell at all imlcss kept out of water tor several days ; and some have described
the faint odour ot the dense fur as very pleasant.
Besides the evidence of their spraints otters trequently leave indications ot then-
existence in an area in the torm ot rejected or discarded portions ot food. Feeding,
except of small, readily consumed objects, always takes place on shore and very often
at detmitc sites such as a secluded strip of river bank or a conveniently flat rock. Such
favorite feeding spots arc often littered with the heads or tails of fish, or even whole
bodies with a single bite out of them, for otters are sometimes wanton killers of more
than they need. One ot the most frequent signs in Africa is the remains ot crab shells
or claws, since ^4();))'.v appears to have a preference for these arthropods above fish.
Another sign of the recent presence of an otter is the footprints in soft sand or mud.
These, amongst those who specialize in himting otters, are knowni as "seals", a technical
term that has been in use for at least three centuries. These tracks differ amongst the
West African otters, since some feet are clawed, others are without them ; some are
webbed between the digits, others not.
Although they are fiuidamcntally land animals otters are not there so completely
adept as they are in water. Their walk, though brisk enough, is a kind of slight waddle
from one side to the other with head low and back humped. Yet some are known to
cover considerable distances, up to 20 or 25 kilometres, at this pace, and apparently
without tiring. When put to it they can gallop moderately fast, "humping" along fore-
teet and hmdteet alternately. However, apart from their not very expert gait they still
remain highly flexible, swift and versatile in movement, rolling, twisting, turning,
scrambling over rocks, logs and other obstacles. They can climb to a certain extent,
and, in fact, have been recorded (Stephens, 1957) as sometimes sleeping up in low
trees. They often stand upright on their hindfeet, using their tails as a third support.
Otters have several different sorts of call or other vocal soimds according to circum-
stances. Animal noises are always difficult to express in writing; moreover, they differ
quite a lot in the various kinds of otter and are not very well recorded for the species
occurrhig in West Africa, if at all. Not much, therefore, can be said on this subject.
The shrill nocturnal whistle of the European otter is, apparently, never uttered by
African species. Harris (1968) describes the soiuid made by .4('/))'.v when suddenly
alarmed as "a strongly aspirated and explosive 'HaliV ". The same author says that
this species uses "a querulous moaning wail" to express anguish, apprehension or,
sometimes, greeting. Shortridgc (1934) says he has frequently heard African otters
LUTRINAE 137
"barking"; and, quoting Stevenson Hamilton, gives the sound made by Aoiiy.x at
bay as a tliroaty "Kwa-a-a, kwa-a-a". Yoiuig otters, as those of so many other kinds
of mammals, continually utter bird-like twitterings and plaintive squeaks.
Though otters may occasionally idle in water they do not often do so, wakingly,
on land. Unless they arc actually asleep in their shelters they are generally busily active
and rarely, like so many other animals, lie merely basking in the sim. When they are
not engaged in eating or excreting they inquisitively examine all the details of their
surroundings ; or when this eager inspection palls they indulge in play, either alone or
with others of their family. Otters have earned a remarkably wide reputation for
play; and they arc, without doubt, amongst the most dcdicatedly playful of all animals,
their activities in this field having every mark of real play and utter enjoyment without
hint of anything more piu-poseful behind it. The stories of those who have had the
good fortune to watch young otters m nature or who have kept them as pets are
endless and are pervaded by a bewitching charm that is, at best, no more than palely
reflected in even the most persuasive accounts of other animals of any kind. This is no
place to record these, and only brief generalities of behaviour can be given.
Play takes place equally in water and on dry land, and as much by a lone otter as
between two or three. It not infrequently involves the deliberate selection and use of a
toy, and the otter's capacity for deriving continuing amusement from the simplest of
sources is a matter for some amazement. In water, play may consist of delighted and
quite aimless gambolling in the form of dives, twists and turns, or of endless rapid
revolution about the longitudinal axis. It may consist of dropping a stone and diving
to catch it in its descent; or it ma)- be a walk on the bottom, shuffling a shellfish along
with its nose. One otter is recorded (Maxwell, 1961) as taking a ping-pong ball down
to the water to enjoy, and expertly appreciate, for lengthy periods its lively antics
when released at a depth. A ball has its fascination also on land, being struck along with
the nose or dribbled football fashion with the feet or flung from the mouth over the
shoulder. In nature stones are used for this exercise in place of a ball; and an otter may
occupy itself with a smooth pebble for a long time. The young may sometimes play
a species of "tig" with one another. One of the pastimes for which otters arc most
widely famous, either alone or in company, is sliding. This is easily done in winter in
cold climates on snow or ice, set slides being deliberately made and repeatedly used;
but when or where these elements do not exist mud slides down steep river banks are
made. These are by no means uncommon in Europe, but whether such slides exist any-
where in Africa seems never to have been definitely recorded. Water-chutes are similarly
used for play where rivers fall over flat rocks in rapids.
The amusing and interesting things that otters can do in the artificial surroiuidings
of a human habitation caruiot be gone into very deeply here, the more so as circmn-
stances must differ in every case. But it may be recorded that the otter's forcpaws,
which are so fashioned that they are expert at holding things, arc rarely idle, exploring
the possibilities and uses of anything within reach. This is particularly so with the West
African Aoiiyx in which the fingers are imhampcrcd by claws and the structure is not
unlike, in appearance and manageability, a monkey's paw. So dexterous are these
hands that Aoiiy.x can open tins or bottles (Eyre, 1963) or peel hard boiled eggs, and
138 Tiir, cARNivonns of west atrica
amongst other tilings has shown itself to be an expert pickpocket. When indulging in
such an operation, or on the rare occasions when they may attempt slyly to rob a
companion of some tit-bit, these otters have the engaging habit of distracting attention
from their act by turning their heads away and gazing abstractedly into the opposite
direction.
It may be added here that otters have proved themselves to have most excellent
memories both for persons and situations, recognising tnends after prolonged absence
(Pitt, 193S). They are long-lived, one being recorded as existing tor 23 years in captivity.
There is no doubt that otters, caught yoimg. make highly intelligent, enchanting,
and indeed lovable, pets, being not only extremely entertaining but affectionate and
loyal as well. Nevertheless, a warning must be given. Thcv are ccrtainlv not for every-
one or for those who like to preserve a well-equipped and tidy house. Their insatiable
inquisitiveness, their sense of play resulting in mischiet or destruction, their deter-
mination to get their own way, and their constant liability to soak clothes, furniture
and beds calls for an even temperament and a forbearance not possessed by all. Add to
this, occasional lapses into bad temper resulting in painful and sometimes serious bites,
and it will be seen that the care of a babv otter is not a task to be entered upon lightly,
cspeciallv if it is accompanied bv admission to the house.
Something must now be said of breeding, though remarkably little that is detuiite
is know-ii of this even as regards the long-observed European otters. Copulation takes
place in the water, lasts an hour or more and is repeated several times over the course
of a day or two. The period of gestation is very much in dispute. Probably it is normally
in the nature of 9 weeks or a day or two less, but this period may be very much length-
ened by delayed implantation of the fertilized ova in the womb. There may be any-
thing between i and 5 cubs at a birth; but both these extreme figures arc unusual and
the normal expectancy is 2 or 3. The newly-born cubs are covered with fme, short
fur but the eyes arc closed and appear to remain so for a matter of 5 weeks, though
opinions differ largely as to this. Probably all the figures here given for these matters
will be toiuid to be only approximately correct for species in West Africa. The young
are, if necessary for safety, carried from place to place by the bitch using the loose
skin at the back of the neck. They are even said, still as blind juveniles, to be transported
thus actually under water from one river bank to another; which would appear to
indicate that not only do the nostrils and ears close reilexly but that, also, the oxygen-
conserving submersion mechanism described on page 134 conies automaticallv into
operation at a very early age.
It has been said (Stephens, T9S7) that the yoiuig are born toothless; but two juvenile
skulls of West African Lulra and .■lc)//)'.v in the British Museum are both well supplied
with teeth. Otter bitches make very good mothers, caring for their litters assiduously
and protecting them fiercely. They alone bring up the family, at least in the initial
stages, the dog otter not being allowed to come near the nest though lie remains in the
vicinity, occupying a holt of his own. Tlie cubs seem to develop fairly slowly and do
not leave the nest for several weeks. TJiereafter they have still to be taught swimming
and diving and to acquire proper proficiency in hiuiting. At this stage the father may
return and take part in the training. The cubs probably remain with the mother untij
LUTRINAE 139
they are sexually mature, which is after about two years. There seems to be generally
at most one litter a year, possibly less. Even in the diversely seasonal climate of Europe
litters may occur in any month of the year, winter as well as summer ; in the tropics it
would seem that season, for water-haunting animals, must matter even less.
Otters have been found to harboiu: a number of different kinds of internal parasites,
both in the gut and the bloodstream, of all the usual groups of wonns, flukes and
protozoa; but nothing is known of this in connexion with African species. They have
also been recorded as carr)'ing ticks, and it has long been believed that otters visit the
sea from time to time in order that the salt water should rid their coats of these and
other ectoparasites. In the normal course of events otters spend some time cleaning
their fiu: by scratching, rolling, rubbing, biting and possibly licking. The last does not
seem to be a common habit; but as otter fur is at times a constituent of the droppings
(Stephens, 1957) they must at least occasionally clean themselves, or their cubs, in this way.
Nevertheless, external parasites seem to be less common than on some animals, possibly,
sea-cleansing apart, because the extreme densit)- of the underfur makes life amongst
it and passage through it difficult. The hazard of drowning, too, must always exist.
Apart from disease otters have two main enemies, man being the chief. There is,
indeed, reason to believe that the otter population of the world, and especially of
newly expanding areas such as Africa, is markedly decreasing, partly by direct persecution
and partly because their once quiet haunts arc more and more being uivadcd and diver-
ted into economic use. The other enemy, in tropical waters, is crocodiles. Otters have
often been observed in places infested with these reptiles and though they show no
visible awareness or signs of fear it is obvious that they must continually be on the
alert, ready to make a swift avoiding tiurn or to flee to the bank. They may also be
aware that a crocodile's periods of entry to the water have something of a set daily
rhythm dictated by the demands of their cold-bloodedness and virtual lack of bodily
heat control except by alternate smming and immersion. It is possible, also, in the
tropics that pythons fmd their \\ ay into otters' holts; and though an adult dog or bitch
would give a good accoimt of itself in a fight with such a snake — though it might well
be hampered in a restricted space — juveniles would easily be taken. And there is always
the danger, on land, of being surprised and sprung upon by a leopard or other of the
larger felines.
Taxonomy. The position of the Lutrinae as a distinct and valid subfamily of the
Mustelidac has never been brought into question since Gray (1865) fu'st suggested this
classification. Within the subfamily itself the genera are fairly clear except that Paraonyx
is often regarded as no more than a subgenus o( Aoiiyx since it was so treated by Eller-
man, Morrison-Scott &: Hayman (1953). This course is adopted here. Some of the
characters picked on by Hinton in the type as diagnostic of Pi!M()/!]'.v are seen not to
be valid in other and better skins; and while there is, indeed, a remarkable difference
in the teeth it is one of size rather than of anything phylogenetically more fundamental;
and the rest of the characters, both cranial and external, denote an undeniably close
affinity with Acnyx.
The second West African genus is the almost world-wide Lutra, which ranges over
much of Europe, Asia, Africa and America. In consequence there have almost inevitably
140 THE CARNIVORES OF WEST AFRICA
been attempts to subdivide it eitlier into independent genera or at least subgenera. Tlie
sole African species of this genus, macuUcollis, was first gcncrically separated from the
others by Gray as HytJrogalc. This name was later found to be preoccupied and the
proposed genus was eventually restyled Hyihiais by Pocock who, by reason of a
number ot small dirterences, both cranial and external, supported Gray's interpretation
ot the position. Once again, the distinctions drawn appear too minor to warrant full
generic separation and Hydrictis has commonly been reduced to the status ot a sub-
genus. It is so dealt with herein; but see page 141 for further comments on this.
The fullest study of the taxonomy of the Lutrinae is a long monograph by Pohlc
(1920) dealing with all the genera, species and races. Pocock (1921b) described some
aspects ot external characters.
Tlie two West Atrican genera may be told apart thus:
KEY TO THE GENERA OF LUTRINAE
(previous key page 93)
Cheeks, lips, throat, sides of the neck and the entire chest wholly white or cream;
forefeet without long claws and with only slight webs; skull length more
than 115 mm; mastoid projecting very prominently behind the aural oritice
Aonyx {pai^c 148)
Wliite or cream area contuicd to the mid-throat and upper chest, and then to a
greater or less degree irregularly blotched with the normal dark pelage colour;
a similar spotted patch often on the after part of the belly; toreteet with well-
developed claws and webs; skull less than 115 nnn; mastoid not remarkably
prominent ......... Lutra [\h^gc 140)
Genus LUTRA Brisson, 1762
Typical Otters
Lutra Brisson, 1762, Ri'^tuiin Aniiiialc in classes IX distrihnlnm . . .. 2nd edit.: ij|. Type species Miistcla
lutra Limiaeiis, Sweden. Lulra was the Latin name for an otter. (It is doubtful whether Brisson's Ri\;iuiiii
Atiiiiiale is properly available under the hitemational Code; but certain ot his names, including Lutra,
were proposed to the Commission tor validation, see Ellcrman & Morrison-Scott, 1951 : 3, though no
action has been taken on this and the matter has been dropped. Even in the case of rejection the name
still stands as of Briinnich.)
Lutra Briinnich, 1771, Zoologiae Jundamcnta . . . Grumie i Dyrctocrai. Type species Mustila lutra Luni,aeus.
Hydrof^alc Gray, 1865, Proc. zool. Soc. Loud.: 131. Type species Lutra utaculUollis Lichtenstein, South
Africa. This name was preoccupied having already been used twice, by Kaup, 1829, and by Pomel.
1X48, in each case tor a shrew. It is derived from the Greek liydor, liydr-, water, and ijfi/c, a wcisel.
Hydrictis Pocock, 1921, Proc. zool. Soc. Loiid.: 543. Type species Lutra macuUcollis Lichtenstein. Tlie
name was compounded from the Greek words hydor, hydr-, water and ictis, a weasel. Valid as a sub-
genus.
There is further synonymy ot little concern in African litcr.uure.
General. As already stated above, the typical otters are widely spread over a good
deal of four continents. Many of the chief characters and habits of the genus have been
LUTRA 141
given in the general introduction to the subfamily and there is no need to elaborate
further what has been said there; the more important distinctions between it and Aonyx
are set forth in the key just given and will become more apparent ni the detailed
descriptions which follow. It remains only to consider briefly in what characters the
African section Hydrictis differs from the most typical, Palaearctic, members of the
genus, which, beyond this, are of no other taxonomic concern to this work.
Hydrictis was, following Gray, proposed by Pocock as generically sepiarablc from
Lutra on the groimds of a reduction in the size of the rhinarium and a simplification
of the external ear; larger, more fully webbed and hairy-solcd feet, the palmar and
plantar pads being less well developed. The skull was distinguished by "many cranial
characters, especially the shortness of the muzzle, length of the orbital floor, and the
generally immatiu'e aspect of the skull owing to the feeble development of constrictions,
crests and prominences". This last is a very apt description, and even the oldest Hydrictis
skulls are in no way so robustly built as those oi Lutra Intra. The most noticeable feature
is the narrowness of the interorbital region and complete absence of postorbital pro-
cesses, as shown in figure 21, and the consequent lack of the usual lutrine interorbital
pentagonal plateau, observable in figure 24 o£ Aonyx. The sagittal and occipital crests
are relatively insignificant; the mandible and its condylar structiure weaker. There is,
indeed, quite a strong case for regarding Hydrictis as valid at full generic level.
Subgenus HYDRICTIS Pocock, 1921
African Long-clawed Otters
Since Hydrictis consists of a single species, maculicollis, no description ot the subgenus
is called for beyond the differential characters just given above and the account of the
species which follows.
LUTRA MACULICOLLIS Lichtenstein Speckle-throated Otter
Lutra maculicollis Lichtenstein, 1835, Arch. Naturgescli. I: 89-92, pi. 2, f. i. South Africa (Bamboos Moun-
tains). The specific name was derived from the Latin words macula, a spot, and collum, the neck, in
reference to the markings on the throat.
Lulra grayii Gerrard, 1863, Catalogue of the Bones of Mammalia in the . . . British Museum: loi. Port Natal
(= Durban), South Africa. A nonten imdum. This was named after Dr. J. E. Gray of the British Museum.
Lutra malschiei Cabrera, 1903, Boln R. Soc. csp. Hist, nat., 3: 182. Rio Muni. Named after Professor
Matschie, German zoologist. Possibly subspecifically applicable in West Africa.
Lutra maculicollis nilolica Thomas, igii, Ann. Mag. nat. Hist. (8) 8: 726. Malek, upper Nile, Sudan. Possibly
applicable in West Africa.
Distribution and general. The name speckle-throated otter is that most generally
accepted for this animal; but it is a little misleading in that the picture conjured up by
"speckling" is of a much fmer order than that occurring in this animal For that reason
it has sometimes been termed the spotted-necked otter. This, though in one respect
more accurately descriptive is in another less so since the back and sides of the neck
have no markings. The most correct name would be spotted-throated; but this has
14- Tin; (AKN'IVOIMA OI WEST MIUC.A
never Ihx'h used — possibly Ixwuiso it is less euplionious than the term most eonimonly
.ipplicd.
The speckle-throated otter is widely spread throughout Africa trom near the Cape
northwards to Ethopia in the east and Liberia in the west. Specimens have been recorded
from most West African territories but in the British Museum exist only from Sierra
Leone (Waterloo, and near Kenema); Nigeria (Oban, Maiduguri); and Camcrouji
(?Ndop): six skins only, two of them juvenile, and two skulls, one alone of which is
adult. Nevertheless, maculicoUis is thought to be not luicommon in suitable localities
tliough, being secretive and wary, it is not often come across by collectors. A.J. Hopsoii
observed it to be frequent on tlie shores of Lake Cliad. It is also plentiful aroinid Lake
Victoria (Proctor, 1963); and it is possible that the species is more at home in lakes
and similar wide expanses of water than in rivers. Skins are not commonly seen on sale
in local markets. Indeed, T. S. Jones (personal communication) thinks tliat, at any
rate as far as Sierra Leone is concerned, tliis is a much scarcer species than Aouyx.
Amongst about 30 otter skins lie examined during his years in that coimtry only one
was Lntid — that cited above from Kenema. Kuhn (1965) gives this species as occurring
in the following places in Liberia: Farmington River, Gbanga, Kpeaple, Biadatou,
Deaple. Harbcl, Kahuple, Siron, Tappita, Towaitown and Zwcdru.
About 10 different races have at one time or another been named, 6 of them recog-
nised in G. M. Allen's Checklist (1939). None of these is actually from the area dealt
with in this book, but it is possible that matschici Cabrera described from lower Camer-
oun applies to otters of this species from the forest belt of Nigeria; vN'hile iiiloiica
Thomas may cover specimens trom the Sahel zone. The position, however, is by no
means clear. Harris, indeed, cites Liberia, Nigeria and Cameroun as comitries of
occurrence of the nominate race though these areas are, of coiu-se, geographicallv, and
probably ecologically, most distantly separated from the type locality.
Description. Liiiii'i inaciilicollis (Plate 2) is the smallest of our three otters, \\ itli a
head &; body length of about 65 cm and a tail which is rather more than halt this. The
full-grown weight is about 4-5 kg for a dog otter or 3-5 kg for a bitch (Mortimer,
1963). These figures arc the outcome of actual weighings of living animals; the often
quoted "not more than 20 lb (9- 1 kg)" sems to be nothing more than a visual estimate.
The pelage of the speckle-throated ottt r is almost entirely a deep rich red brown —
intense sepia — both above and below. The base of the fur is, as in all otters. \\'hite.
The only exception to tins deep brown lies on the throat and sometimes the chin, the
fore part ot the chest and the after part of the belly which are white, spotted or blotched
with the normal dark ground colour. The size of these mottled areas varies a great
deal from individual to individual, being sometimes verv much reduced or, particularly
as regards the belK' patch, lacking Tlie upper lips arc also narrowly white, but this is
not very noticeable It is the extent of these white markings which best serves as a means
of field recognition; for although the other two West African otters are bulkier and
have a rather paler pelage, age mav affect size, and wetness the apparent colour of the
tur. In the species now under discussion the white is verv strictly confined to the lower
surface ot the bodv, whereas in both Aoiiyx and P show tliat tlic specklc-
tliroated otter corresponds in liabits and beliaviour tairly closely to other Liiira species
in other parts of the world.
rishing may take |ilace at any hour of the day, or sometimes night, although on the
relativelv undisturbcxl sliores of Lake Victoria the species a]ipears to be chiefly diurnal;
it is commonest soon after sunrise; but the otters may actually alrc-ady have been swim-
ming about for some time before this in the scmi-darkness. From Procter's account
it would seem that maculkollis is markedly more social than most. He records that
parties of from 1 to 6 are fairlv common, larger numbers than this getting progressively
Fig. 20. Lutra maaiUiollis: skull. B.M. No. 23.1.22.44, se.\
I ; lateral view
rarer; but he once observed a schoiil totalling about a score. Small parties are mostly
bitches; the larger ones dog-otters, often voung. The two se.xes arc mostlv difficult
to distinguish apart in the water, though full-grown males are appreciably more
heavily built than females. On shore, identification is easier, particularly at times of
excretion since in the dogs the urine is projected forwards, in the bitches towards the
rear. Defaecation is invariably accompanied bv urination, but not I'l'a' versa; and,
as in other otters, there are recognised sprainting places. The spraints are mostly of the
common, blackish slimy, otter type; but drier faeces of artliropod shells and fish scales
are also dropped. In excretion the body crouches low, almost touching the ground;
a bitch holds the tail out stiff horizontally, the dog vertically. Passage of food through
the gut may be extremely rapid. Hoase-cleanluicss is evident in captive specimens.
Like other otters, macuhcollis indulges m play both in and out of water. Amongst
other things it is not infirequently given to teasing its prey, flinging fish back and forth
I.UTIIA 145
liom l.iiul 10 w.iirr .uiil |nirMiini:, it, sonu'lniu's giviiii; U a sliglu nip In ilisaMc it
p.irli.illy ,ukI slow down us iiiovciiK-nts. (!r.ibs .nul smh like ,ur Inittcil alioiit with the
nose, i'roeier, tluniii; loni; observation oftiie speikle-tliroated otter arouiul tiie shores
oil ake VieliMi.i never saw nukl-slKles like tiiose reeorileil as made elsewliere hy other
Fig. 21. I.iilrii iiiiUiiliiollir. skull, 11. M. No. ij. 1.22. 44, sex ?, X I ; p.ilatal t\
ili»rs.u N'lcws
species; Init Mortimer notieeii what might well have been one ol these, the more so as
there were otter tracks around it. Incidentallv, the tracks (".seals") lett hy this otter
can be readily distingiiislied troin those ot the two otlier West Atricaii species by tjie
clear prescjice otclaw marks and, on very soft mud or sand, the taint imprint of webs
reaching to tlie ends ot the digits.
146 rill-. CAUNIV ORES or WKST AlUICA
Wlu'ii diviiii; to tisli. the liod\' is arLlu'd .ilinost clear ot rlic \\ate\ and tlic subscqiioiit
dive is luarlv vertical. This otter, as other species, exhibits astonisjiing grace and
agilit\-, especially 111 the rapidity ot its turns: but there is not much actual speed m
swimming. Mortnicr gives measured figures tor short distances which indicate a rate
ot" progress through the water ot only about 4 km an hour. Both tore and hindlcgs
are used in swimming, the main thrust coming from a vigorous use ot the latter.
Apart trom swimming, diving is also graceful and skilled, being earned out trom
heights of up to 6 feet, leaving scarcely a ripple on tJie water. One ot these otters,
however, is not above an ungainlv scramble into the water with a large splash wlien
suddenly alarmed.
On land the gait is rather awkward but progress somewhat taster, f-or ordinary
purposes a rather clumsy walk is used in which the feet are moved alternately or the
hindlegs as a pair — this latter a gait sometimes used by domestic dogs as an alternative
to the normal trot. A taster speed can be achieved by a run, the legs moving alternately.
Mortimer's measured speed tor this is about 5.2 km an hour; and tmally there is a sort
of gallop, the animal ■"humping" along, moving fore and hindteet in pairs alternately.
This was found by Mortimer to be at the rate of about -j-z km an hour, though it is,
said that over a short distance one ot these otters can keep up with a running man,
which would be at least twice as tast as this. Mortimer's experience with a young
tame otter made him doubttul whether this species could travel long distances overland
as this one, at least, appeared to become very soon overheated. Clambering up and
over rocks is effected without diiiicultv; and Procter observed tliat a jump ot at least
one metre could be made across a gap.
Although various toods, including crabs and moilu cs, .nv taken tish seem to con-
stitute the chief diet in this species. Procter tound Hiiplochivniis oi from 10 to 20 cm to
be a favourite on Lake Victoria, a genus that occurs, but much more rarely, in West
Africa. TiLipia is readil v acceptable but often proves too tast to catch. Mortimer observed
that a numiier offish would be landed betore the meal was actually commenced; but
this IS certainly not always so. His tully-grown bitch ate trom 4-1 to 4-7 kg a week;
and, as otten popularly held tor other species, invariably started at the tail end. Smaller
tish or molluscs are eaten in the water, either upright while treading water or while
tlie otter is floating on its back. Vegetable tood seems not only acceptable but desirable;
tor Procter observed one of these otters to eat a small quantity ot sedge, and Mortimer's
tame one deliberateK robbed the garden ot carrots, beans, potatoes and peas.
When not in the water these otters spend a good deal ot time grooming their tur.
On emergence from the water the head is shaken but not the body, the drying ot wliich
IS aided bv the usual otter practice ot rolling or rubbing. Vision on land is obvious!)'
only good over very short distances; hearing and smell are better developed. Mortimer
observed the latter to make immediate response to seemingly quite insignitic.uit quan-
tities of water in flower vases or other small vessels. A pungent musky smell is emitted
as a reaction to fright, in yoimg animals at least. The soiuids uttered are various. Procter
describes four. The commonest call is "a shrill chikkering", used when playing or
scolding. The second is "a prolonged mewing ya-a-ii-ii', probably a challenge. Another
LUTRA 147
is "a squealing whistle", made when excited, as in play-figliting; and he once heard
an "ic-yaiig" call but never anytliing resembling a bark. Mortimer described the
commonest sound uttered by his iiuiadicollis as a high-pitched squeak which, in certain
circumstances, changed to a high-pitched trill.
Very little has been found out about breeding in the speckle-tlnroated otter. It
would seem that a litter of 3 is a common number. Procter thouglit the period of
gestation might be a little over 2 months. Mortimer, who observed and recorded growth
over a period of nearly two years, found the weight of his female to be 1-36 kg at
li months; 2-5 kg at 5 months; 3-3 kg at 6i months; and 3-5 kg at 7A months, at which
time it was apparently full grown, for the weight thereafter remained stationary at
this figure, at least up to 20 months when the weigliing ceased. The body length, too,
remained luialtercd. Teeth were still being cut at 6 months.
Taxonomy. Nine forms have been named from different parts ot Africa, six ot
them currently recognised in G. M. Allen's Checklist (1939). These have been diag-
nosed, often as independent species rather than races, on the characteis ot size or colour
or markings, and mostly from single specimens or otherwise quite inadequate material.
Colour is notoriously uiu-eliable and in iiiMulicclIu is known to change with age (Procter,
1963). The same author indicates that the spotted white underside markings are even
more luidependable since even among the limited population on Lake Victoria "there
is a very great individual variation in the amount of white on the throat. Every gradation
is seen firom no white spotting at all to the whole throat, chest, belly, front and sides
of the fore-limbs and part of the hind-limbs being almost completely white". It has
been suggested that both mat0 THE CARNIVORES OF WEST AFRICA
Distribution and general. This is possibly the most widespread, it not the com-
monest, otter in Africa. Most otter accounts, both ot wild and domesticated animals.
seem to conccrr. th;5 species rather than Ltara. Described originallv irom the Cape of
Good Hope. :? somewhat misleadingly todav referred to as the Cape clawless
otter, it has .^^.^c j^cn collected or reported from a large number of places in most
countries ixom South Africa northwards to Ethiopia in the east and Senegal m the
west. It is as much a forest animal as one of the more open and arid woodlands ; and
it is knov^Ti to occur on mountains at over 2000 metres. G S. Child (private communica-
tion) records it from Kainji Lake and the Borgu Game Reserve in the Doka bush of
e."ctreme ■western Nigeria. It is said to inhabit, sometimes, the same localities as Liitra;
and its unmbtakeable tracks have certainly been observed by A. J. Hopson in the
sand some 350 metres from the shores of Lake Chad, on which ituiculici^llu is kno\\ii
to be tairly common. Moreover, skins of .-iLi/iy.v are frequentlv exposed for sale in
Malamiatori market, not far ir -ke. In Sierra Leone Aonyx is. according to
T. S. Jones (personal commun:,:^:.-.. , plentiful in all areas, being frequent in the
mangrove swamps. Skins are, there also, commonly offered for public sale. It seems
possible that Aonyx sensu stricto is entirelv replaced in certain areas bv the subgenus
Paruonyx.
Nine 'West African specimens exist in the British Museum from: an unspecified
locahtv- in Fernando Poo; Calabar (forest belt), Zaria (Doka woodland) and Maiduguri
(Sudan w-oodland), all Nigeria; Ashanti and Asikum near Oda (both forest) in Ghana;
Sierra Leone, Bonthe and an unspecified localirv- but almost certainly forest; Bolama
Island. Porruguese Guinea (mangrove?) ; and Gambia, locality unkno\\ni. Two are
complete juveniles; of the remainder onlv two have skulk, and there are no external
measurements whatsoever.
Description. The Cape clawless otter (Plate 2) is a bulky animal. At 16 to 20 kg
it is tar larger than L. inacuUcoUis; and even bigger specimens than this have been
recorded in South Africa, up to nearly 30 kg. The head & body length is from about
750 to 925 mm and the tail about 110 to 5S0 mm, but precise measurements for West
Africa do not exist. Indeed, of all the Aonyx skins in the British Museum only three
have measurements, one from Kenya and two from Ethopia. The coat is variable in
colour from a deep red-brown, almost as dark as L. indLiilicollis, to a distinctly paler
mid-brown, the spinal zone being a little darker than the sides. In some cases the
hairs have pale tips, panicularly over the shoulders and neck, but these are not so white
as in Puraoiiyx and the overall effect not nearly so "frosLcd". The feature which, apart
from size, visually distinguishes this otter from maadkollis is the wholly white or
cream, quite unspotted, chin, throat, upper chest, side of the neck and of the face to
the level of eye and ear. The margins ot the ver\- small ears are white.
It is the feet, however, that constitute the most important ditFerence berw,-een Aonyx
and LutTd. They are quite devoid of the usual carnivore long, sharp claws though in
some cases furnished v\.-ith rudiments. These last are to be found only on the hindfoot,
usually as tiny flattish nails on the 3rd and 4tli digits. The digits themselves are finger-
like in appearance; and, indeed, the whole paw is very similar to a small hand, the
more so as the interdigital webs are much reduced on the forefoot. The structure is.
AONYX 151
in fact, used for grasping much in the maimer of a hand and is far more deft than the
usual mammalian structure apart from the primates. The palmar and plantar pads
are better developed into a miited central pad than in Liitra. The rhinariimi is broad
and rather dog-like.
Skull. The mature Aoiiy.x skull is an incomparably larger, stronger and more rugged
structure than that o( Ltitra maculicollis. The rather flat braincase is not only broader
but has also, in the males, a sharp, deep sagittal crest and pronoiuiced, upturned supra-
occipital crest which continues round the sides of the braincase to exaggeratedly large
and prominent niastoid processes. There are also large, slender paroccipital crests.
In the females this cresting is not so higlily developed ; while in the young of both
sexes the cranium is roiuided and quite smooth.
The long intertemporal and short interorbital legions are abruptly narrow and more
or less parallel-sided ; there are sharp postorbital processes and slight ridges joining
them to the sagittal crest, thus marking out the pentagonal plate usual in this subfamily.
The arches are strong with well-developed jugal hooks reaching up towards the post-
orbital processes. The rostrum is extremely short and falls away almost vertically;
the anterior nares very large. The palate is rather narrow and continues, parallel-
sided, well to the rear of the toothrows. The bullae are small and flat. The condylar
hinge is deep and strong; the mandible ver\' powerfully built, with a tall coronoid
process, much excavated at its base in the ramus to accommodate a large muscle.
The dentition is powerful. The incisors of the upper jaw form a nearly straight,
serried transverse row, those of the mandible being often a little more irregular. The
canines are exceptionally long and sharp, suited to penetrating deeply and holding
securely a peculiarly slippery and active prey; the fust tliree premolars, above and
below, are relatively small, p'^, when not lacking, and often p- as well, being practically
in contact with the inner face of the canine. The result of the smalhiess of these teeth
is that, though there is not strictly what is technically known as a postcanine gap (see
page 18), there is enough space between the two toothrows anteriorly to accom-
modate a fair amoimt of flesh and thus increase the firmness of the canines' hold on a
fish. The cheekteeth as a whole form a short compact series; the upper caniassial has
one long pointed cusp, the lower three, set in equilateral fashion, but soon wearing
do\\ii and together with the very broad posterior molars forming more of a crushing
than sectorial imit, well suited to dealing with the soft flesh of fish and molluscs. In
spite of the general impression of strength the teeth as a whole seem ver\- liable to
both wear and damage. It is the great breadth of the posterior cheekteeth wliich
chiefly serves to diflerentiate Aoiiyx seiisti stricto from the subgenus Paraonyx (fig. 22).
The maximum crowii width o( m^ in the former is of the order 13 to 15 mm as con-
trasted with 95 to 10-5 mm in the latter.
Habits. Aoiiyx has often been kept in capti\'ity, and if procured at a sufficiently
early age makes an agreeable, interesting and amusing pet, friendK" to both human
beings and other animals. Its behaviour in domestication has therefore been well
observed and corresponds closely to that of other otteis in similar radier anificial
circumstances. But its way of life in nature is a different matter. Although the species
has often been wxitten about most of these accounts consist, in fact, of a good deal of
152 THE c:aknivoiu-s or \vi;st afiuca
surmise aiid generalities and remarkably little that is actually positive. Statements,
indeed, seem sometimes to be diametrically opposed; as when {fide Shoitridge, 1934)
Lancaster says that Aoityx capcnsis is solitary as a rule, while Moseley asserts that it
hunts in small companies. Both, of course, may be true, the latter applying, as in any
other otter species, to a bitcli and her yoiuig.
The African clawless otter may, in remote streams, lakes or swamps, sometimes be
come across fishing or sporting in the water at high noon; but where it has reason to
tear man it confines its activities to the early and late parts of the day, and possibly
becomes largely nocturnal. A great drawback ot having to adopt a nocturnal existence
must lie in the increased difficulty of detecting and capturing tish in the dark. Possibly
crabs are easier. Aony.x is always held to be much less ot a fish eater and more of a
crab eater than Liitra iiuuulicollis. It is a fact that the spraints contain a good deal ot
crab shell, and the feeding sites, constantly revisited, are littered with claws and other
remains of these crustaceans. The species is said to take, also, the usual list ot other
foods — molluscs, lizards, small rodents, birds and so forth. Eyre's tame ^4ci;/j'.v greatly
appreciated freshly-killed moles. It has in southern Atrica a great reputation as a
fowl and egg thief; but no clear evidence of this seems to be forthcoming and any
such robbery might as well prove to be the work of a ratel. Certainly any interest in
eggs does not appear to lie in their attraction as food; tor Eyre (1963) records an
occasion when a domesticated otter, having ot itselt discovered two dozen eggs in a
cupboard, instead of treating them as a desirable meal, hurled them, presumably one
by one for amusement in the usual otter fashion of dealing with a pebble, all over the
floor. Zammarano (1930), as the result of first-hand experience in the field, decided
that the clawless otters were endowed with very acute senses, and he characterized
them as cautious and cunning in the highest degree and the hunting ot them, con-
sequently, as anything but easy. The species has otten been observed on the sea coast;
it is, or was some years ago, to be seen in the large lagoons, not far from the sea, between
Tiko and Douala (Cameroun). T. S. Jones obtained a specimen in a comparable situation
of mangrove and sea at Bonthe (Sierra Leone).
Clawless otters make good pets if taken young and brought up on the bottle. In
captivity these animals, though demanding in time and patience, e.xert considerable
charm. They display quite unexpected intelligence, solving problems which they
would not come up against 111 nature such as opening tins, bottles and cupboards with
amazingly dexterous fuigers. The paws are used almost as hands, not only holding
things but also to throw small articles (Eyre, 1963). This author describes a variety of
sounds made; a high-pitchcd squeak, demanding attention; a purring growl with a
warble in it when pleased; a whine of frustration; a hiss and a growl when trightened;
and a "most luiearthly scream, most trightening, when he is in a temper". The explosive
"Hah I" uttered bv .4i'/i)'.Y in the wild has already been referred to in the general obser-
vations.
Nothing defuiite appears to have been recorded of breeding. The normal litter
seems to be 2 or 3, but as many as 5 has been estimated trom a tield observation of a
bitch and cubs. T. S. Jones (personal communication) had numbers of young brought
to him in Sierra Leone and from these the litter size appeared to be invariably 2. The
AONYX 153
period of gestation is unkiiowji but has been guessed at as about 2 months. There is
probably no set breeding season in West Africa but T. S. Jones reckons that it is nor-
mally between September and October, the whelps being born in a hole in the bank.
On the 5th October he obtained a very young animal which had its eyes open but which
he thought was still being suckled. The skull of this specimen is, in fact, very thin and
there is no sign of the permanent dentition. The skin has the correct adult coloration
though the fur, as noted earlier, has not its fmal composition. On the other hand, two
newly-born specimens exist, both wholly white, above and below. Eyre (1963) says
that in a young animal the round eyes are a deep navy blue. As with other otters, the
young have to be taught to swim by their parents — a fact that must be borne in mind
with captive whelps. They are often somewhat reluctant.
Taxonomy, hi spite of the very v/ide range of this otter there is little doubt of there
being but a single species of ^4o»)'.v in its restricted, subgeneric, sense. The question of
races is more obscure. Pohle (1919) recognised four, With, poensis Waterhouse in addition
as a separate species; G. M. Allen (1939) six, none of them from West Africa. There
have, however, been four attempts to relate special forms to this region, three of the
proposed types resting in the British Museum. In 1838 Waterhouse named pocnsis
from a skin which, from its size and the nature of the pelage, appears to be that of a
young animal. The diagnostic characters given, largely colour, are without much value ;
the skin has no feet and it is therefore not absolutely certain that it is Aoiiyx at all.
In fact both Thomas (1889) and Lonnberg (1910) considered it to belong to Lutra
maculiaillis. However, the throat, neck and sides of the face follow the pale, unspotted,
pattern of Aonyx, but instead of being white or creamy they arc, as Waterhouse
described them, "of a rich deep golden yellow with a faint brownish hue" — though in
the past 130 years this has become obscured by London grime. Pohle (1919), disagreeing
in respect of synonymy with Gray, Trouessart, Thomas and Lonnberg, accepted this
unusual colouring as having full specific validity. He further entirely rejected Thomas's
and Lonnbcrg's opinions regarding the genus of this specimen on the grounds that the
throat was luispotted, a character that never occurred in the niaculicollis group. That
Pohle himself seems to have been wrong in this is demonstrated by the iiiaciilicollis
skin, B.M. No. 34.9. 16. i, with indisputable claws on a forefoot but without spots,
and not very dissimilar from the specimen now under discussion. The possibility
therefore yet remains that the poeiisis type is, in fact, a Lutra; and nothing further can
be decided regarding its generic, specific or subspecific standing until more and better
material emerges from Fernando Poo.
The next proposed form, calaharicm Murray from Calabar in south-eastern Nigeria,
was foimdcd on a skull very poorly diagnosed by Murray, not only as a species but as
the type of a distinct new genus as well because it had one fewer premolar in the
upper jaw than Lutra vulgaris of Europe. This absence oi p^ is now known to be a not
unusual dental variant, and J. A. Allen (1924) regarded the species as "practically indeter-
minable" from Murray's diagnosis. The skull, now in the British Museum, appears to
dirter in no material respect, except possibly its slighter size, from other typical capetisis.
Next, never referred to in literature except by Gray himself, comes his gamhianns, of no
date but prior to 1865. The skull of this reputed species also is in the British Museum
1_S4 THE CARNIVORES OF WEST AFRICA
and, apart from its somewhat larger size, docs not differ from typical capcnsis. The
name is a iwincn uihlimi; no description of the species seems ever to have been attempted
and it appears, with untraceable reterences, merely in the list of synonyms ot Aoiiyx
laliwdii Lesson given in Gray 1865: 130 and Gray 1869: 109.
Finally there is Rochebrune's supposed kiioiri from Senegal. The work in which
this is reputed to have been described is said (Thomas, 1889: 196) to have been privately
printed and hence never validly published. However, he apparently actually examined
Rochebrune's description and remarked, in a footnote, that the so-called "diagnosis"
surtered the usual lack of all diagnostic characters. This, therefore, is also a ncmcn
nudum ; and in this case there is, moreover, no biown existing specimen of the reputed
species whatsoever (Lonnberg, 1910: 3).
Whether or not there arc forms that might constitute valid West African races is
quite indeterniinable from the study material at present available. In the erection of
such races considerable caution would be called for in view of the knowii idiosyncratic
variation that occurs in otters — illustrated earlier in this account by Procter's obser-
vations oiLntra macnlicolUs. The situation, in fact, has changed very little in the half-
century since J. A. Allen (1924), with commendable understanding and restraint,
wrote of the then reputed forms of ^4(';i)'.v capcnsis: "As these five forms appear to have
been described in each case from a single specimen, without flesh measurements and
in some instances from poorly prepared material, none of them can be said to rest on
a very satisfactory basis. The differences in coloration indicated by the descriptions of
these forms are more than covered by the range of variation in the present Lang-
Chapin series of some twenty specimens from a single locality (Faradje), while the
individual difference in size is more than covered by the twelve adults. The status of
these various forms should be held more or less in abeyance until a good scries from
each type locality has been studied and compared. Under such circumstances, it seems
better not to add another name to the list . . . ."
Such skull measurements as are derivable from the present very poor West African
material in the British Museum are given in the table on page 159. There are no external
data.
Subgenus PARAONYX Hinton, 1921
Small-toothed Clawless Otters
The chief characters which distinguish this subgenus from Aoiiyx scnsti stricto are
sufficiently shown in the key on page 149. The main point of difference certainly lies
in the dentition, the contrast in size of the posterior cheekteeth being so marked,
as shown in figure 22, that in spite of the otherwise strong overall general resemblance
of the skulls it is easy to tell the two subgenera apart at a glance.
Three species have been described: couq^ica Lonnberg from lower Congo, philippsi
Hinton from Uganda, and micradon Pohle originally from lower Cameroim but
subsequently reported also within the limits set for this account. These have been
pretty generally accepted as independent species; but it appears to the present writer
AONYX 155
that the evidence in support of this, based as it is on insinifigcant differences of size
argued for the most part from single skulls, is very slender indeed. The three "species"
can at most be reckoned as races; and the value of even such a reduced status is, on the
few data now available, of doubtful worth or validity. Lonnberg's description ot
congica, the earliest form now assigned to Paraoiiyx, was based largely on a broken skull
from which very limited measurements could be taken. Because of tliis, PolJe, in
later erecting micmdon, was able to make only three cranial comparisons, though in
respect of teeth he made ten. None of the differences he cites seems to have much
importance. Hinton in diagnosing philippsi as the type species of his new genus Para-
onyx appears to have been quite unaware of PohJe's earlier creation. The three forms
are here treated as a single species, congica.
AONYX CONGICA Lonnberg Small-toothed Clawless Otter
Aonyx capensis congica Loiinbcrg, lyio, Ark. Zool. 7, No. 9: 1-8. Lower Congo.
Aoiiyx iniaoioii Pohle, 1920, Arch. Nalurgcscli. for 1919, sect. A, pt. 9, 85: 145-147. Bomsc, Nana River,
Cameroun. The specific name is from the Greek micros small, and odon tooth.
Paraoiiyx philippsi Hmton, 1921, Ann. Mag. nat. Hist. (9) 7: 196-200. Lake Bunyonyi, British Ruanda
(= Uganda). Type in the Biitish Museum, No. 21. i. 22.1; skin in good condition, skull with the
arches broken and partly missing. Called after Capuin J. E. Philhpps, M.C., District Commissioner
at Kigezi.
Distribution and general. No specimen actually assigned to congica or emanating
from the Congo exists in the British Museum; but there arc four skulls and five skins
of philippsi, all from Uganda. The position as regards the form most closely associated
with western Africa, microdou, is in some ways the best. The existence o£ Paraoiiyx in
West Africa itself was not suspected until in 1938 Dr. M. D. W. Jeffreys obtained
several skulls from the Nun marshes. Two others were procured later by the present
writer from the same area, so that there arc now 9 skiills and a skin in the British
Museum collection, a much better representation than there is of the other forms and
of the more widely distributed Aonyx capensis.
It seems impossible today to fix with precision the whereabouts ot the village of
Bomse whence micivdon was originally described; but it was probably at about 5°55N.,
I5°I5E. The Nana River is a tributary of the Sangha and flows eventually into the
Congo. It is thus just extralimital to the area covered by this present work. The Nun,
or Noun, River flows into the Sanaga and thus lies well outside the Congo basin to the
north. Its upper reaches pass through the Ndop plain and are there extensively marshy,
the centre of this swampy area being at about 5°55N., io°25E., that is to say some
40 km directly east of Bamenda. The surrounding area is today open country with
scattered trees of a rather Guinea woodland type, though in the past it was very probably
closed forest, rehcts of this vet remaining. The altitude of the Ndop plain, though lower
than that of the siurroundmg mountains, is probably still of the order 1300 metres.
No specimens assigned to microdon from elsewhere in the area herein treated as
West Africa seem ever to have been recorded; but Pohle cites a number of localities
of occurrence in lower Cameroun. These, however, are for skins imsupported by
156
THE CARNIVORES OF WEST AFRICA
skulls, and their identification as what we now call Paraonyx rather thaii as typical
Aonyx is, as Pohle himsclt pointed out, somewhat uncertain.
Through the courtesy of Dr. P.J. H. van Brce of the Zoologisch Museum Amster-
dam it has been possible to examine a skull procured by him from Loa-Loa near
Makokou, North-cast Gaboon. This lies in a geographical situation distinct from those
of the other specimens imder consideration, since it is neither in the Shari and Niger
Fig. 23. Aonyx (Paraotiyx) amgica: skull, B.M. No, 1938.9.29.4, 5e.\ ?, ■■ i ; Literal view
drainage systems, taken in this work as constituting the major part ot West Africa,
nor in the Congo basin but in the westwardly sloping area emptying into the Atlantic
via the Ogowe River. The locality is some 700 km south of the Nun marshes. Dr. van
Brce's specimen ditfers in certain respects from the iiiicwdoii material from this last
area; but it is less toothworn, and the basisphenoid suture is still detectable, as it is not
in the British Museum material. Moreover, it is a female, whereas none of the London
examples is sexed. These facts may account in some measure for the differences which
arc apparent: there is no sagittal crest, no sharp supraorbital processes, the rostrum at
the level of the canines is appreciably narrower, the mastoid processes are far less bulky,
and the coronoid process more slender. But even it these distinctions held for further
specimens from this area they are racial rather than spiecitic.
AONYX
157
F.G. a4. Ao.y. iParaony.) co„sica: skull, B.M. No. Z938.9.39.4, sex ?. x .; paUal & dorsal
I.SS THE CARNIVOKIiS OF WEST AFRICA
Description. In general overall appearance Pciraoiiyx is extremely similar to Aciiyx.
The body bnild is much the same, possibly a little larger; the pelage colour and pattern
basically the same, that is to say deep sepia-brown on back and belly but the chin,
throat, cheeks and chest wholly creamy-white without spots. The most obvious
external dirtcrcncc lies ni the "trosting" of the head, neck and shoulders due to the
bristle-hairs being white-tipped, A little of this may be present in A. capcnsis but it is
usually either lacking or not very marked; in A. coti^ica, on the other hand, it forms a
conspicuous feature extendnig from just back of the eyes to a little past the shoulders.
In some cases it is pretty concentrated, in others well diffuse but none the less obvious.
The margins ot the small ears, too, are more conspicuously white. The forefeet arc
un webbed; the hindfeet only to the second joint, Hinton said that the facial vibrissac
were weakly developed in comparison with capensis; but there is not much evidence
that this is so except possibly in so far as those situated above the eyes are concerned.
Skull (hgs, 23 and 24). Apart trom the dentition, which is comparatively shown in
figure 22, there is not much to distinguish the mature Paiaonyx skull from mature
Aotsyx except tor a rather broader interorbital and intertemporal breadth. The rami,
instead of being more or less straight-sided, curve in towards one another.
Attention has already been abundantly drawn to the relatively small size of the
posterior cheekteeth, the most important diagnostic feature of the subgenus; the average
crown width ot iii^ is only about <)h mm as contrasted with at least 50 per cent
more in capensis. The canines arc rather shorter and slighter; and in 8 out of 13 skulls/)^
was never developed.
Habits. Absolutely nothing is known ot the habits ot l\iraouyx. Lonnberg (1910)
and Hinton (1921), however, both made some interesting surmises from the nature
ot the tect and the reduced size of the posterior cheekteeth, supposing from the one
that this species would probably prove to be yet more terrestrial than capensis; and
from the other that it was not well adapted to dealing with hard-shelled Crustacea but
rather with small terrestrial vertebrates and eggs. In this connexion it is of interest to
note that the Ndop plain, which microdon inJiabits, has numerous small streams and
other water-courses but no largish rivers such as otters normallv like to frequent and
swim in and which would be much better stocked with fish. The swampy nature of the
terrain makes it ideal for amphibian lite, and it would thus ccrtainlv seem possible that
this otter consumes a high proportion ot frogs and relatively little tish.
Taxonomy. Something has already been said ot this. There seems to be no good
reason to suppose that three independent species exist. The question boils down to
whether the named forms do in truth constitute valid races. The data are slender and a
little confused. There is no specimen ot coiigica in the British Museum from which to
make direct measurements and comparisons; in respect ot tins, then, there exist only
Lonnberg's figures from a broken skull (incidentally, misquoted in Hinton, 1921).
On this very fhmsy evidence it does, however, seem to be slightly the largest. The
table on page 159 gives the impression that microdon is a little larger than pliilippsi;
but the 3 skulls of the latter are all females; the microdon skulls are unscxed but from the
appearance of the sagittal crest nearly all seem to be males. The evidence, therefore,
IS quite inconclusive and a third name is, at an\' rate tor the time being, best omitted.
LUTRINAE — MEASUREMENTS
159
Table 8: Numerical data for subfamily Lulrinac
Lutra
iiiacuUcollis
Vegetation
Various
Number in mean
4
Condylobasal length
105-7
Basilar length
94-4
Palatilar length
41-4
Zygomatic breadth
62-0
Upper cheekteeth breadth
32-6
Interorbital breadth
i6-o
Postorbital constriction
i6-6
Braincase breadth
5I-I
Toothrow [c — ml)
33-0
p* length
II-7
ml length
8-1
ml breadth
9-5
mi length
13-3
ma length
4-5
Head & body
648
Tail
378
Hindfoot
109
Ear
18
RATIOS (per cent)
Tail/head & body
58
Zygom. br./condylob. 1.
59
Biaincase/condylob. 1.
48
Braincase/zygom. br.
83
Palatilar l./condyloh. 1.
39
Interorb./postorb.
96
p'^jc—m^
35-5
p^jm^
144
mi/;n2
296
Aoiiyx capciisis
calabariais gambiaims
Type Type
Forest PGuinea
I
I
(130)
(140)
114-4
I25-I
59-0
65-4
86-7
93-8
(47)
(49)
26-3
32-3
24-5
26-1
66-5
69-0
42-4
46-6
14-4
(u-7)
13-2
13-3
14-5
iS-6
19-8
I9-I
(6-6)
6-2
(67)
(51)
77
(45)
107
34-0
109
(300)
(f'7)
(49)
74
(47)
123
31-5
no
308
A. (Parac
inicrodon
?Guine3
9
125-0
113-2
54-1
93-4
39-5
32-1
31-9
70-2
37-9
9-8
8-6
10-3
12-4
4--I
75
5strum is usually long, but less so in the Hcrpcstinae; the anterior nares wide.
The palate is mostly rather narrow but broadens somewhat posteriorly and is con-
tinued in a narrow- parallel-sided extension considerably to the rear of the toothiows.
The bullae arc large and bulbous, showing anteriorly a more or less clear constriction
indicating the internal septum dividing the structure into tw-o chambers; but Nandiiiia
is extremely unusual in having the posterior one of these merely cartilaginous and thus
disappearing from all but very carefully prepared skulls. The paroccipital processes are
sometimes virtually absent or indistinct, but arc mostly well developed and closely
adherent to the posterior aspect of the bullae. In old skulls the fusion of all sutures is
very complete.
Flower & Lydekker (1891) wrote that the second lower incisor was raised above the
level of the first and third; but, at least as far as West Africa is concerned, this is wholly
untrue. The six incisors of both jaws form a compact more or less straight transverse
row, the outer ones being somewhat or pronouncedly larger than the others; in some
cases the lower incisors arc bilobed. The premolars arc ot the usual form; the upper
caniassial has an anterior lobe despite Flower & Lydekker 's assertion to the contrary,
though mostly small and sometimes, with wear, indistuict. The upper molars are
mostly narrowly transverse and are usually markedly smaller than this tooth, but m
some of the mongooses j/A does approach />' in size; j;i-, when present, is generally
much smaller, and in XiVidiiiiii is reduced to a peg. The dental formula is very variable
even within a single subfamily, the total number of teeth being 36, 38 or 40. Since the
incisors and canines are always '^ this is due to diversity in the premolars and molars
of both jaws, these cheekteeth numbering 7-^, r^, j-,, or ^.
Habits. Since these are vastly different in the different subfamilies, or even genera,
little that is ot general application to the family can be said here. The Viverridae are
by nature fierce, both in the pursuit ot their pre\' and in self-defence; but some ot them
it captured at an early age prove themselves to be interesting if not charming pets.
Many people have kept, and have derived pleasure and even benefit from, the civet,
genets and several different kinds of mongoose; but some of these, if not all, exhibit
far more independence of their homes and keepers than members of the Canoidea or
even of the Felidae or true cats.
The Viverridae are nearly all essentially nocturnal or crepuscular but some of the
mongooses can be come across by day. They arc all basically carnivorous, consuming
various kinds of smaller creatures, rodents, birds, reptiles, amphibians, myriapods,
crabs and insects; but many, if not all, also consume fruits. They mostly live in small
holes or crevices; but, in fact, extremely little beyond broad generalities is known of the
way of life of the majority of them.
Locomotion in certain African viverrids has been photographically investigated,
analysed and sometimes figured bv Taylor (1970) in relation to these species occurring
VIVERHIDAE 163
ill West Africa : Civcttictis civcttii, Gciictta spp., Nainhnia hinotata, Atilax paludinosus,
Herpestes ichneumon, Galerclla sanguinca, Ichnewnia albicaiuhi, and Miingos iitungo. The
motions studied arc walking, trotting, running, galloping, jumping, climbing, swim-
ming and burrowing. He found that the only form of locomotion common to all
was walking; trotting is the gait most usual to the bigger open-country viverrids,
probably in association with the larger territories there customary.
Taxonomy. Of the three major groups into which the Feloidea arc traditionallv
divided there is little disagreement regarding the clearly separate identity of the Felidae
and the Hyacnidac and, at the same time, the close affinity existing between the animals
of which each of these families is composed. The Viverridae, however, are more open
to doubt. The six apparently natural groups of animals, comprising nearly 40 genera,
therein lumped together may seem a somewhat heterogeneous collection to form a
single family. Pocock, basing his ideas largely on external characters, gradually veered
away from the standard conception of the Viverridae as a close-knit unit and came
ultimately to elevate several of its six subfamilies to full family rank. All three West
African groups were concerned in this more disassociated conception of Pocock's,
the civets and genets becoming the Viverridae in a restricted sense, the mongooses
the Herpcstidae (originally Mungotidae), and the West African palm-civet the Nan-
diniidae, independent of the Asiatic palm-civets with which it had previously been
associated.
Simpson's (1945) classification, however, rejects this and retains the wider conception
of the Viverridae; and this grouping is adopted herein. It has a long history of accep-
tance by taxonomists, dating back, to all intents and purposes, a century and a half to
Gray. During this period the majority of systematic mammalogists, no matter what their
renaming or rearrangement of major cadres may have been, have seen little to disagree
with in the close association with one another of all the small carnivores under dis-
cussion.
As regards the taxonomy of the Viverridae in general it must be added that Diicker
(1957) from observations on instinct and behaviour reached the quite divergent con-
clusion that the Herpestinac are, in fact, related rather to the Canoidea than to the
Feloidea, and most particularly to the primitive Mustelidae.
The three subfamilies may be separated by the following kcv, external distinction
being easy but cranial differentiation more difficult.
KEY TO THE SUBFAMILIES OF VIVERRIDAE
(previous key page 160)
I. Coat speckled, or unicolorous blackish, sometimes with a transverse pattern of
bands, but never spotted; tail bushy but never ringed; posterior part of
the bulla with one exception (Galerclla) much more inflated than the
anterior part and subglobular or rather taller than it is long; auditors
meatus with posterior and anterior lips which meet below in a horizontal
V, thus forming a narrow gap which often extends as a slit into the floor
l64 THE CARNIVORES OF WEST AFRICA
ot the bulla; postorbital aiidjugal processes long and nearly or actually
joining to form a complete circumorbital ring . Herpestinae {pa'^c 239)
Coat spotted or blotched, tail at least partially ringed; the whole bulla inflated,
oval and longer than it is tall; or [Paradoxurinac) partly cartilaginous
and usually mostly missing in prepared skulls; the auditory meatus a
shallow rmg without projecting bony lips; postorbital processes short
(except Paradoxurinae) and far removed from a very short jugal process,
leaving a widely incomplete circumorbital ring ..... 3
3. Two pale spots on the shoulders (sometimes indistinct); coat colour deep red-
dish brown; bulla cartilaginous or missing; postorbital process well
developed ....... Paradoxurinae [pngc 229)
No pale shoulder spots; coat reddish but not deep; bulla normal; postorbital
process short ....... Viverrinae (/Jiijjt' 164)
Subflimilv VIVERRINAE Gill, 1872
Civet, Genets, Lmsangs
Distribution. The first of the three subfannlies to be dealt with in this work con-
tains S genera. These are divided by G. G. Simpson (194.S) into 2 tribes, the Viverrini
and the Prionodontini, of which only the former is of West African interest, the
latter, with 2 genera, being wholly Asiatic. The Viverrini also comprise 2 genera that
are wholly Asiatic, besides 3 wholly African and one that occurs ni both continents
and extends its range into extreme south-western Europe as well. These figures,
however, are subject to dispute according to the breadth ot conception of some of the
genera. Very considerable doubt and controversy e.xist also in regard to speciation in
Genctta: but in this present account the 3 relevant genera of the subfanuly are held to
cover a total of 12 species.
There are a few apparently very rare and extremely local viverrines but the majority
of the genera are common; and no part of West Africa, from coast to Subdcsert,
is without some representative ot the subfamily. Extralimitally the group is widespread
throughout the whole continent.
General characters. The West Atrican animals included in the Viverrinae are
very diverse in size, ranging from the bulky civet, scaling perhaps 17 kg, to the slender
linsang, which can scarcely weigh more than 2 kg at most. All members of the sub-
family dealt with herein have a prominent spotted or blotched pattern to their coats,
never being merely speckled like the Herpestinae; and their mostly long or very long
tails arc at least partly ringed, and with few exceptions more or less cylindrical in
form. The face is sharp, the ears mostly fairly large, rounded and conspicuously up-
standing above the level of the crown — in this, as in many other ways, the subfamily
differing clearly from the Herpestinae. The ears differ further in always possessing a
marginal bursa on the pinna. The limbs are short or shortish, ending in 5 digits armed
with curved claws that are mostly to some extent retractile but in the civet scarcely
at all. With the exception ot the pads, and sometimes the area immediately around
VIVERRINAE 165
them, the soles of the feet are hairy to the heel. The pelage is composed of fme underfiir,
often very dense, and of bristle-hairs of roiuid or very slightly flat section, which in
some genera arc very long and dominate the coat, in others of subordinate hnportancc.
The scent glands in the Vivcrrinae are perineal, situated between the scrotum and
the prepuce in the male, and between the anus and the vulva in the female. They are,
besides being of different form, thus quite differently sited from those of each of the
two other West African subfamilies. The glands open externally into a medial pocket
or pockets; and in one case, Civittictis, the waxy secretion collecting therein has been
commercially exploited for many centuries.
Skulls. The skulls are narrow; even in the case of the very large and in some ways
exceptional Civctlktis the braincase is little broader than that of markedly smaller
herpestines. The rostrum is long and narrow, ver)' distinctly longer and narrower
than that of any mongoose except Liberiictis. The orbital ring is far from complete,
the postorbital processes being short, the jugal processes often little more than rudi-
mentar)'. The postdental palate is short, nowhere near so long as it is broad. The large
bullae are long and ovoid; the anterior chamber well-inflated, not depressed and
reduced in size as in almost all Herpestinae.
There may be 38 or 40 teeth, the cheekteeth numbering ^ or ^. Again with the
exception of Cii'cttictis the dentition is by comparison with the mongooses relatively
light, the anterior premolars being narrow, almost linear. It is clearly of a sectorial
nature, not of an insectivorous type found in many members of the other subfamily.
Habits. Although nearly all the Viverrinae are common animals their lives in the
wild have not been much observed. This is because they are nocturnal, solitarv and
secretive, spending much of their time in trees or dense undergrowth. All but the
civet are arboreal, though they often come down to the ground to hunt. They are
basically mainly carnivorous, the genets fairly strictly so, probably preferring small
mammals and birds to other things; but the civet at least, and possibly genets too,
consume reptiles, eggs, termites, beetle grubs and so forth if opportiuiitv offers. They
also from time to time eat fallen fruits.
So far as is known all viverrines are hole-dwellers, either on the ground or in trees.
Breeding habits and periods var)' and the little that is known of them will be found
under the different genera, below. Both the civet and the genets have been kept as
pets; but while they tolerate domestication during early life they remain more aloof
than the mongooses and soon seek their independence in the wild.
Taxonomy. There is little doubt of the distinctness of the Viverrinae from the
Paradoxurinac and Herpestinae, though the relationship with the former is much
closer than with the latter. In fact, as explained later in the introduction to the Herpes-
tinae, it is open to considerable doubt whether the two groups are really as close to
each other as G. G. Simpson's classification puts them. General form, pelage, feet,
claws, scent glands, skull characters and teeth are all quite distinctly different.
Within the subfamily the genera are clear, and there has not been that confusion
which arose in the Herpestinae. But there is argument regarding the closeness of
affinity between African and Asiatic representatives of the subfamily which affects
I66 Till (AHNlVdlUS 1)1 WIS! AIUHA
two ot tlic throe genera covered by tliis present aeeouiit. (jrciiiciis and I'oiaua. Tiie
tormcr is \\idelv held to be synonymous with or at most a subgenus ot the Indian
I "ivcna; and the latter to be closely allied to tjie Asiatic Imsangs, Prioiiodon and Panhcils.
The present audior follows Pocock in considering Cii'ctliais to merit generic standing
of its own tor reasons given later in the account ot this genus; and he further believes
superficial resemblances between Atrican and Asiatic linsaiigs to be misleading and
probably more the outcome ot convergence than of particularly close relationship.
KEY TO THE GENERA OF VIVEKRINAE
(previous kcv page 163)
1. Size large, head ^ body about 800 mm; tail only about halt as long; a white
or pale patch, sometimes obscure, bordered with black on the side ot the
neck, and a black mask across die face trom cheek to cheek. Adult
skull about 140 mm; postorbital constriction not marked; dentition
powerful ........ Civettictis {pdi^c id?)
Size much smaller, head & body not more than 550 mm, otten much less;
tail at least three-quarters as long as, or longer than, head & body; no
conspicuous black and white pattern on the side ot the neck or mask
across the face. Adult skull rarely as much as 100 mm, mostly mucli less;
postorbital constriction well marked; dentition light .... 3
2. Very slender; head & body about 380 mm; tur very short and velvety; mostly
without any clear dark spinal stripe; spots always small; tail otten widi
taint intermediate rings. Adult skull about 70 mm; cheekteeth 5-^;, the
posterior lower molar being minute .... Poiana [pa^c 21'j)
Of heavier build and mostly larger size, head & body 400 to 550 mm; pelage
longer and coarser, otten with a distinct dark spinal stripe; the markings
may be small spots but are more trequcntly larger blotches; tail without
intermediate rings. Skull 75 to 90 mm; cheekteeth ^ Gcrietta (pd{;c 177)
Cenus CIVETTICTIS Pocock, 191 5
Atrican Civets
I'lrom Liim.icus, 175X, Syni-iiui i\\iliii\h\ lotli cJ., I: 43; m pait, ot v.irious .uitliors, cspcci.illy bctorc
1915. Type species ("both by elimination and selection", Tlioma'., lyii) I'/Vcnii ci7)c(//ii l.inn.ieus,
Bengal. I'ivcria was the Latin name for a ferret.
Civettictis Pocock, 1915, PiOi. :oo\. Sec. LoniL: H4- Ivpe specie-. \'iviiiii
■3
lyO THE CARNIVORKS OF WEST AFRICA
T]ic dorsal pelage is coarse and rather wiry; it varies in length, being in some speci-
mens fairly long and in others unexpectedly short. It is composed of rather imdulatc
and tangled undertur in which are mixed fairlv abundant, verv slightly flat bristle-
hairs, strong distally but tapering basally. These bristles are markedly longer than the
imdertur, which they dominate not only by reason of this but because, also, of their
much greater diameter throughout most of their length. They average about 40 mm
in length but sometimes attain 50 mm, the underfur measuring only some 20 to 30
nmi; and the\' may be wholly black, or white with longer or shorter black tips. Along
the mid-dorsal line ot the back from the shoulders or the hinder part of the neck to
the root of the tail runs a long, black, erectile crest, the stout bristles of which arc
everywhere appreciably longer than the main pelage but increase in length posteriorly
where, in the region of the hindquarters, they may attain 100 mm or more. These
spinal bristles are dark througliout most of their length but always have longer or
shorter glossy jet-black terminal portions. When, in anger or alarm, this crest is erected
the animal assumes an appearance of even greater size than it really has.
The coloration of the African civet is widely variable. The dorsal ground-colour
ranges between near-white, through creamy, to a slightly reddish-buff. On this is
superimposed a pattern of dark spots or blotches, sometimes deep-brown but generally
black. The size, shape and independence of these markings vary. Most typically the
spots are large, of rather irregular roundish or quadrate shape, clear-cut, and nearly
all separate from each other. They are not set in more or less regular longitudinal
lines, as in some of the genets, but something in the nature of half-a-dozen rows can
be made out on either side of the spinal crest. Occasionally the spots, especially those
near the medial line, tend to coalesce into longitudinal bands; but in some specimens
they display a tendency to rmi together laterally; and in some thc\- are relatively ill-
defined. Over the torequarters the bold spotted pattern fades out and the pelage becomes
an intimate mixture ot light and dark hairs, any maculation in this area being very
obscure apart from a number of spots low on the shoulder.
On the side of the neck is a pronounced longitudinal black band which is separated
below by a conspicuous white band from a second black one which broadens and
passes below across the throat. The underparts are somewhat variable, the chest being
always black or blackish; but the belly may be white, or blackish, or a mixture of the
two. The face is strikingly marked with a black mask which crosses from check to
cheek and just encloses the eyes. It is bounded anteriorly by a white area around the
rhinarium and on the lips, and posteriorly by a whitish frontal area between the eyes
and the ears. The continuity ot the mark across the face is interrupted in some speci-
mens, though rarely, by a medial whitish band joining the anterior white to the frontal
grey area. The ears are rounded, low but none the less conspicuous with their w hite
rims. A bursa is always present; its convex posterior flap is continuous above and below
w^ith the rim of the pinna; its anterior flap is widely but not deeply emarginate.
The upper portions of both fore and hind limbs are obscurely spotted; but the
lower parts and feet are wholly black. There are five digits on each foot, the claws
fairly long and only moderately ciurved, and very slightly retractile. In both feet tlie
central plantar pad has the small pollical lobe separate from the main body ot the pad
CIVETTICTIS 171
(Pocock, 1915a). In the forefoot the metacarpal pad is bilobed and is joined, in most
cases, by two narrow strips of naked skin to the anterior naked sole area that lies
between the central and the subdigital pads. The hindfoot, also, has this anterior sole
area naked. The tail is little more than half the length of head & body. Dorsally it is
wholly black; but the basal half has about 5 partial white rings, passing from side to
side below. The tail is coarse-haired with very long bristles.
The perineal glands have been dissected, figured and described by Chatin (1874)
and Pocock (1915a). Since these two accoiuits differ somewhat in their details the
following is founded mainly on that of Pocock. He characterized the organs of the
male and female as "tolerably similar". The glands are situated between the scrotum
and the prepuce in the former, and between the anus and the vulva in the latter. Exter-
nally they appear as paired swellings separated medially by a slit-like orifice about
25 mm in length. When these two glandular lobes are drawn apart a moderately large
oval orifice can be discerned on the inner face of each, and this leads into a large hair-
lined sac or pouch which extends forwards, backwards and upwards within the gland.
The inner walls of these two sacs secrete into them a very thick whitish or )-ellowish
substance with a powerful musky odour, the "civet" of the perfiimery trade; and this
then makes its way tlirough the two oval orifices into the intervening space between
the two glandular lobes, wliich thus becomes a storage reservoir conununicating with
the outside world by the slit-like orifice first mentioned. Besides these perineal glands
tliere is a pair of anal glands situated on tlie wall of the rectum. These secrete a foetid
yellow liquid which imparts to the faeces a characteristic odour which may be of
service in the demarcation of territory but, in view of the regular defaecation habit
mentioned later, is more likely to be of defensive significance, as in the case of other
small carnivores.
Complctch^ black specimens are recorded from time to time; and in these it is
impossible to detect any trace of the basic pattern, at any incidence of light, as one is
often able to do in other melanos. One British Museum skin, referred to by Sanderson
(1940), is quite different in its coloration from all the rest. Li this, the ground-colour
of the entire pelage, including the normally white areas of the neck and face, is a
bright ochre-red. This may, of course, be natural, due to some form of erythrism; and
if it is it is, indeed, remarkable. But the skin is obviously a locally prepared one, and
Sanderson thought its luiique appearance probably due to its having been cured in
smoke, a not uncommon practice. The colour, actually, is not so much that yellowish
tone produced by smoke as, rather, one which could be caused by steeping in palm-oil.
This would have been a not improbable proceeding in the area from which the speci-
men came (Ogoja, extreme eastern Nigeria); and the soft pliability of the skin, as
contrasted with the usual brittle stiffiiess of African sun-dried pelts, lends some support
to this idea. The fur would, of course, have been subsequently washed to rid it of oil,
but the residual red-dyeing effect of the palm-oil in the unpigmented hairs would
resist this. Against all this is another fact: that though the long glossy black crest and
tail are quite typical, the spots are smaller and more obscure than usual, as if there were
something fundamentally exceptional about the whole skin.
Skull (figs. 26 and 27). The skull is strongly built, long and rather narrow, with a
172
THE CARNIVORF.S OF WF.ST AIIUCA
Fig. 26. Cii'cttictis civctta: skull, B.M. No. 59.^41, V,
lateral ■
long ovoid braincase and not very marked intcrorbital and intertemporal constrictions.
The supraorbital processes, though clear, are blunt, deflected, and not very extensive.
All adult skulls, male and female, have a well-developed sagittal crest, joining a sharp,
flange-like supraoccipital crest posteriorly in a T, The zygomatic arch is strong, the
juga! process pronomiced but blunt and short. The bullae are moderatelv large and
oval, the posterior section iar exxecding in size the anterior. The paroccipital processes
are closelv applied to the whole posterior aspect ot the bullae and extend ventrally
beyond their limit in long, strong points. The mandible has deep, strongly built rami;
and, indeed, the whole skull and dentition constitute an instrument fashioned for
powerful biting.
The dental torniula is
3 14:
40. The incisors are strong and set in a transverse
curve: 111 both jaws the outer ones arc larger than the inner; but this is most pro-
noimccd in the upper jaw, where fl is separated trom the others by a small gap, is con-
siderably larger and, being pointed and curved, begins to resemble a small canine rather
than the normal chisel-edged incisor. The canines themselves, though strong, arc
not particularly large, and their outer faces lack the fme, shallow furrows characteristic
of the other West African viverrids. All the cheekteeth, from ;)i to \n~ and pi to »i-.>,
are strongly built; m^ and ;ii- areparticular]\' large tor this subfamily; m\ is exceptionally
broad posteriorly, and the whole tooth has more of a crushing than sectorial function.
The posterior lower molar, also, is uncomnionlv well-developed, with a flat, far less
acutely cuspidate, grinding surface than is usu.il.
CIVETTICTIS
173
One British Museum skull. No. 10.6.19.1 from Okigwi, eastern Nigeria, has an
extra, large, premolar on each side of the lower jaw, set between p4 and 0/1.
Habits. Everyone is agreed that the African civet is an extremely secretive animal.
This fact combined with its more or less strict nocturnal habits is responsible for the
almost complete lack of field observations. It may seem strange that this should be
Fig. 27. Civctlkus civctia: skull, B.M. No. 59.941, j:.
pah
&d
orsal views
SO in a species that is pretty common in museum collections and which tairly regularly
appears in zoos; but it is a not very intelligent animal and one that is readily trapped;
and so, though not often observed in nature, becomes easily killed or captured. Some-
times a civet may be seen trotting up the road in the headlights ot a car: and occasionally
in the very early morning one may dart swittK' across a path, possibly disturbed by a
174 T"I- CARNIVORES OY- WEST AIIUIA
Jog troni its tcinporar\ resting place in the thick undergrowth. But civets can also
walk stealthily; and even it one should be about in the halt-light it is not easy to detect
Its grev spotted coat aganist checkered vegetation. With so little known about their
habits it is impossible to be certain, but civets are generally believed to lead entirely
solitarv lives except at breeding time. They are certainlv almost completely terrestrial,
their shape and teet being little adapted to climbing in the manner of cats, though
where there are positive tootliolds thev can scramble up short distances, particularly
betore thev have attained their adult bulk. Thev are said to be able swimmers, not
atraid ot taking to the water. Normally, during daylight hours they lie curled up in a
temporarv "torm" in any dense vegetation; but when breeding they use a tlxcd nest,
usually a hole in the ground made bv some other animal, or amongst tangled roots.
It seems probable that such nests are not lined in any wav.
When it comes to the question ot tood, information is rather more certain because
ot the data yielded by droppings, stomach contents, and behaviour in captivit}'.
African civets seem to be more or less omnivorous. Antelope meat, cane rat, guiiica-
towl, eggshell, carrion, termites, beetles, grass and leaves have been found in stomachs
(Bothnia, 1965), carrion being predominant. But thev are also said to eat other kinds
ot insects and their grubs; rats, snakes, lizards, frogs, crabs and millipedes. T. S. Jones,
in a private comnuuiication, savs that civets not uncommonly hunt in mangrove
swamps, presumably to obtain crabs and mud-skippers. Me has trapped them widi
stock-fish or dead rats; and Verhcyeii (1951), indeed, gives them almost the character
of complete scavengers. At the right seasons they consume an exceptional quantity
of berries and other truits. T. S. Jones observed that thc\- caused a good deal of trouble
amongst young low palms just starting to truit at Njala (Sierra Leone) by gnawing
otf the oilv mesocarp. Astley Mabcrly ([9S5) records that they commonly cat the
truits ot Briddia and Zizyi'liiis; and since civets mostly deposit their droppings in one
fixed spot until they torm a heap, such places otten abomid in dense seedling growths
of these trees. Verheyen (1951) found one such excremcntal pile to measure 60 cm in
diameter and N to is cm thick. Civets, like other viverrids, arc known to raid poultry
houses at night; and Shortridge (1934) recorded that they sometimes killed )-ouiig
lambs. The widely catholic tastes in tood of African civets, and their piartialit}' to truit,
has been well demonstrated in captivity; specimens, within the present writer's know-
ledge, consuming almost anything put down for them, including bowls of pawpaw
and banana. That some food should jiave excitant properties is of interest; Mallinson
(1968) records that it has been found inadvisable to feed freshK' killed chicken or rabbit
to viverrids with newly born yoiuig; female civets, and others, so fed have been
found to become highly excited and subsequently to turn on and kill their litters.
Some ot the older writers on natural history stated that the amoiuit of "civet" secreted
bv animals maintained in captivit\' tor this purpose depended on the richness and
abiuidance of tood given.
The African civet can become verv tame and docile in captivitN' it caught \ouiig
enough; but Menzies (1966) foimd those of 6 weeks old already savage and to remain
quite intractable. However, without question, reaition to .ittempts at domestication
will varv with different indniduals. One lu-t civet kept, or at least encouraged, about
CIVETTICTIS 175
the house used to follow its owners like a dog for evening walks; and at dinner time
would circulate roiuid and imder the table giving unsuspecting guests playful nips in
their fingers should they by chance hold their hands down. Eventually, as sexual
maturity is reached, civets, like so majiy imconfuied pets, disappear into the bush
never to return. An African civet has been known to live for almost 14 years in cap-
tivity.
Civets make a variety of sounds. They hiss when alarmed and utter a deep warning
growl when they feel they are likely to be attacked. If they do get into a fight with one
another there is a great deal of excited growling and yapping very like a dog-fight.
Some years ago residents in the environs of Lagos often heard such quarrels at night,
being sometimes awakened by the tremendous caterwauling and yapping.
There is little positive information with regard to breeding; but Mallinson (1968)
has recently published a few facts gathered from animals in captivity. From two
observed copulations he deduced that the period of gestation might be either 45 or
60 days. Nine births, covering 20 young, showed the litter size to lie between i and 4,
the commonest being 2. The mothers frequently killed, and sometimes ate, their
new-born young; and it \sas foimd less worrying for the female and less likely to lead
to traged)' if the father were removed. Walker (1964) stated that there are generally
two litters a year; but Mallinson, on the contrary, found with 5 females that the
average interval between litters was approximately 12 months, and never less than
288 days. Immature specimens in the British Museum suggest that there is no favoured
season for breeding in West Africa since ver)' juvenile animals have been taken, in the
forest belt, in mid-October, December and January, that is to say the beginning and
middle of the dry-season ; and also in mid-June and early July, well into the rains.
According to Vcrheyen (195 1) the newly-born young remain only about a week in
the nest.
For many centuries the civets have been chiefly famous as the source of a widelv-
used ingredient of perfumes. This is an off-white or yellowish, very thick, greasy
substance with a powerful musky odour secreted into the perineal pouch as already
explained earlier in this account. Like many other basic materials, vegetable as well as
animal, employed in the production of scents "civet" in concentrated form is apt to
be nauseating or even highly repulsive; but when used in minute traces becomes not
only tolerable but attractive. The trade, which was once extensive in Europe, North
Africa, the near and middle East, has now considerabl)' diminished, at least in Europe ;
but formerly African civets were kept imder cruel conditions in very small cages in
which they could not turn round, so that the animal could be easily controlled and was
constantly available for the product to be scraped, with little effort, from the glandular
pocket. This took place about twice a week, the animal being forced back against the
back of the cage and its tail and hindlegs seized and held through the bars while the
"civet" was collected, using a small spoon. The yield was about 2 or 3 grammes a
time. Abyssinia was recently the chief source of supply, the substance being exported
packed in hollow antelope horns; but earlier, Amsterdam was the centre of a big
trade, both for the import of the substance to Europe and for its production on the
spot by captive animals. The Dutch trader William Bosnian, who was resident m
I7f> Till: CAKNIVORIS or WF.ST A Ml 1 C A
r.ible y: Numencil dat.i tor Cii'cliiiti< uiif/ii
West
A tVic.i
general
Vegetation Various
Number in mean 12
Condylobasal Icngtli 14s
Basilar length 134
Palatilar length 73-6
Z\gomatic bieadth 77-1
Upper cheekteeth breadth 47-4
Interorbital breadth 29-2
Postorbital constriction H'O
Braincase breadth 43-0
Toothrow (f— »!-) 57'7
p^ length 1 3 '0
ml breadth 13 '7
mi length I3'5
mi length S-2
Head i\ body S26
Tail ' 432
Hindfoot 123
Ear 57
KATIOS (per cent)
Tail/head & bod)- 5-
Zygom. br./condylob. 1. 53
Braincase/condylob. 1. 30
Braincase/zygom. br. 56
Palatilar l./condylob. 1. 51
hiterorb./postorb. 122
p^c-
--'>
"l/"'2 ~ '"3 165
West Africa towards tlic end ot tlic 17th century and wrote a nuniber ot letters des-
cribing the various countries, tlicir customs and animals, li.nd, in translation, this to
sav of the civet "at present so well known in Holland that 1 need oiiK' acquaint you
tliat tliev are brought to be sold to us verv yoiuig. and then we give about eight or
nine Shillings sterling for one. A large share of Trouble and caretul Attendance is
requisite to breed tliein up: Their Food is Pap boiled or made ot Millet, with a little
Flesh or Fish. They produce Civet when even very young; ot which that ot the Males
IS better than that of the Females, because the latter cannot avoid urining into the
Civet Bag, which spoils it".
Besides its obvious wortli to the scent trade, "civet" was also tornierix- imicli valued
for a wide varictv of reputed, but now long discredited, medicinal properties. It was
emploved as a diaphoretic; or as an emollient in such eruptive skin conditions as small-
pox, measles and scabies; as a balm against apoplew; as either a soporitic or an aphro-
ciisiac; or even as an ingredient ot tooth powder.
GENETTA 177
Taxonomy. Other races besides the noniiiiate one have been nained, G. M. Allen
(1939) recognising two, cotij^ka Cabrera, from north-eastern Congo, and schivarzi
Cabrera, troni eastern Africa. If these arc accepted, then the correct citation of the
West African civet must be Civcttictis civetta civctta (Schrcber). But the species shows
such a degree of nidividual variation that there is a tendency today amongst authors
to be sceptical ot the existence of identifiable races.
Genus GENETTA G. Cuvier, 1816
Genets
Gciktla Okeii, 18 16, LcUrhnch Ar Nalnrgcsclikliu; 3, 2: loio. Type species Vivcrra genelta Linnaeus. Oken's
Lehrhnch is ruled to be unavailable by Opinion No. 417 of 1956 ot the International Commission on
Zoological Nomenclature. This name is a Latinization of the Old French (jo/rt/c, itself derived from the
Arabic name for these animals, janiail.
Gaiclia G. Cuvier, iSi6'/jVc Sherboni), Le Rcgne Animale. I: I>i6. Type species \lverra (fcuciiii Linn-
aeus.
Distribution and general. Genet is a simple enough name; nevertheless it is one
that is not often used by English-speaking people in West Africa, who usually refer
to these animals as bush cats. Not that the average person often, or ever, sees one of
these nocturnal creatures in the flesh; but he may sometimes become aware of them
through a raid on the fowlhouse, or occasionally through being brought a motherless
baby ior sale as a pet. Genets are, none the less, widespread and fairly common, occur-
ring through practically the whole of Africa except the Sahara, and, in the region dealt
with in this book, ranging from the Subdescrt to the coast. They are known to occur
at high altitudes in eastern Africa but were not taken at am- great height on the Camer-
oun Mountain by Eisentraut (1963).
Genets, though all conforming to a general casily-recognisable pattern, occur in a
wide number of different forms. How many of these, and indeed how many basic
species, are valid is a much-disputed question. In this present work nine different species
arc described but, as will be gathered from the taxonomic section which follows later,
some of these may not be valid or, as the outcome of a greatly-needed pan-African
review of the genus, may eventually be shown to be merely western representatives
of other, extralimital, species.
General description. The genets (Plates 3 and 4) are all smallish mammals with
pointed muzzles and large ovoid ears in which a large bursa seems to be always present
ill West African species. The posterior flap of this arises, both above and below, behind
the pinna, the shape of the anterior flap being variable in the different species. The
slender, spotted body is some 450 to 550 mm long, the ringed tail somewhat shorter.
Genets stand about 120 to 150 mm at the shoulder. An average weight of an adult is
around 2 kg. The pelage, both of the body and the tail, consists of a mixture of terete
or slightly flat-sectioned bristle-hairs and fine woolly luiderfur, the absolute and
relative lengths of these varying from form to form. There is often, but not always,
a media! dorsal stripe from about the shoulders to the root of the tail, dark, mostly
lyS rill ' AUNivoius in west aiuk a
lil.ick, toiisistiiig ut soincwli.it. or iinicli. lojigcr briitlcs tliaji tlic rest ot tlic dorsal
pelage, and apparciul\' erectile. On cither side of this spinal stripe run more or less
parallel rows ot spots, the number i^t such rows varying from 4 to about 7, though the
hues arc apt to become contused on the Hanks and difficult to coiuit precisely. The
spots themselves vary in number from comparatively few to many; their spacing from
relativTly wide to close. They range considerably in size from tairly large to pretty
small; in shape from quadrate to oval or roiuidcd; and from full independence of one
another to longitudinal coalescence into longer or shorter bands, more or less parallel
to the mid-dorsal crest. Such concurrence usually concerns only the line or two lines
nearest the spine, and commonK' only the region ot the lower back. The spiots may be
wholly ot one colour, black or some shade ot reddish-brown ; or they may be annulate,
having black margins with larger or smaller brown centres.
The back of the neck is basically marked with 5 more or less parallel longitudinal
lines; a fme central one which posteriorly becomes the mid-dorsal stripe; on either
side ot this a slightly broader, but still narrow, line which joins the innermost series
ot spots; and, on the outside ot these, two broader, and otten very pronounced lines
which posteriorK' curl round and down over the shoulders. But some or all of these
nuchal lines may, ui certain specimens, become wholly or partly obscure. The belly
is usuallv paler than the dorsal ground-colour and may be almost white, well-spotted
or with very tew spots. The thighs are spotted, the uppier piart ot the toreleg often so;
the lower parts ot both legs mav carrv small spots or be practically plain. Thev ma\' be
light or almost black, this, at least as tar as the torelegs are concerned, being otten
idiosxncratic and hence taxonomically unreliable, although in the past it has often
been assumed to be diagnostic. There are tive toes on each toot, digit J being separated
from the others; each is armed with arcuate, tine, verv sharp claws, which are retractile
but not so completelv as in the cats. Beneath each digit is a naked hemispherical or
ovoid pad; a larger, composite one surrounding a central depression is situated 111 the
middle of the toot; and another posterior to this, very long and biturcate on the
hindfoot. l1ic taec carries certain general markings, sometimes very clear, sometimes
obscure though almost aKva\s detectable. The most prominent is a pale, otten white,
spot below the eves; this is separated by a dark patch trom a similar pale or whitish
region on the upper lips aiouiid the rhinarium. There is a pale area between the eyes,
parted mediallv b\' a dark line; and another pale spot above each eve.
The tails, though subject to some individual variation, are more constant 111 their
basic char.Kters than the doi^al markings are, and sufticientU' distinctive to torm the
easiest and most dependable diagnostic character. The nature ot the tail derives firstly
trom the actual length ot the hair with which it is clothed and secondK from the
relative length of the bristle hairs and underfur. In {^ciicttii, tor example, the tormer are
not oiiK- ver\' long in themselves but also tar exceed in length the undertur, which
plavs a verv seccindar\- role, the whole structure being ot relatively rough appearance.
In scrvaliiid. on the other hand, the bn-.tlcs .ire short, and the abundant undertur only a
little shorter, plaving a major role in producing a sott, smooth, subcylindrical structure,
hi addition to this, the number and nature ot the annulations are usetullv diagnostic,
though bv no means so unswervingU' constant as was tormerK- thought.
GF.NKTTA 179
C'lCiicts arc tuniislK'(.i w itli mciu glands secreting a nmsk\' ddoiir. Tlrcsc arc situated
in tiic male between the scrotum and prepuce, or in tlic female between the anus and
vulva. Pocock (1915a) furnished a full description of these, both external and internal.
Briefly, in his own words, they "consist of two elongated eminences covered with hair
both extcnially and intcrnall)-. When luidisturbed the two lobes are closely apposed,
their line of contact being marked by a longitudinal sulcus whicli is Y-shaped anteriorly,
that is to sa\-. just bclnnd the vulva or prepuce". Internally, the glands arc mostly
tripartite, being imperfectly divided into three compartments by transverse ridges of
integument. This description applies to the males oi tigriiia, partliiia, nibiginosa, gciietta,
(kiiigolana (^ senegalciisis) and fcliiia, and to the females of the first three; but the
females of the second three have a simpler, undivided pouch. Pocock regarded this as
indicating two groupings of relationship within the genus.
Skull. Unfortunately Gciictta skulls and teeth exhibit very little constant diflercnces
of shape and so are not a great deal of help in distinguishing forms. Some of the charac-
ters picked upon by Schwarz (1930) are misleading, being not certainly applicable to
all members of a species-group.
The Gciurra skull in general (tig. 29) is long in comparison with its zygomatic
breadth, with a long, ovoid braincase, and pronoiuiccd postorbital processes separating
marked intertemporal and interorbital constrictions. Yet though the whole skull is
long and tapering the actual rostrum itself is remarkably short. Posteriorly, in the
fashion common to the Feloidea, the braincase contracts and curves broadly out again
to meet a very well-developed, flange-like supraoccipital crest. There may or may not
be a sagittal crest over the main body of the cranium; but there is always a wcll-
devclopcd, sharp posterior portion joining the supraoccipital crest in a T. The forward
continuation or absence of this over the braincase has been regarded as a valid specific
character (Schwarz, 1930); the evidence from 29 mature West African skulls in the
British Museum indicates that this mav be broadly so but that its development is a
function of age, and its absence save from really old specimens thus possibly mis-
leading. There is, for West Africa, no evidence relating to sex, comparison being im-
possible since there are no old female skulls o( pardiim, in which species the crest is
normally present in old males. A superficial look at some dozens of Gciietta skulls
from other regions revealed only three reputed females with developed crest. The
bullae are fairly large, long ovoidal in shape. There is some evidence pointing to diff-
erent relative developments of the anterior and posterior portions in different species,
the anterior chamber aroiuid the meatus being largest in gciicna and iliicrryi. less in
pardiiui and least in cristata (fig. 28).
The ratio of the postorbital constriction to the interorbital breadth is fairly reliably
diagnostic but not absolutely constant. In {ifiwtliudcs 19 skulls had the former appreciably
less than the latter; but in 2 skulls the breadths were subequal. The atim/i'im group also
has the postorbital width less, one externally quite typical cristata specimen, No. 10. 6. 1.2
(Oban, Nigeria) alone having it greater. In a few exceptional cases the postorbital
constriction is extremely marked. In one gciicltoides specimen. No. 46.397 (Oda,
GhaiKi), it is as little as .S-4 mm in contrast toa mean of 11-5 mm; the difference between
it and the interorbital breadth being about 6 nnn as compared with a mean of the
iSo
Till. ( AHNIVOKHS OI" WFST AIUK.A
Fig. :;S. GcMifdi; left bullnL-, 3: .1. ('• 'yiuiM (? ii/rii), B.M. No. i;.ii.2.34,
b. (,'. i,'nii7;i'ir/i.f, li.M. No. 4'i.39'', ;-, c. G. iTisiiUii. B.M. No. 39.323, ;
order ot z niui. For such atypical incaMircmciit^ there is no apparent explanation.
The postorbital constriction is wider than the interorbital breadth in the gciu'tta group
and tliienyi. Citing the postorbital processes as diagnostic Schwarz stated that they were
absent or sliglit in scrvaliihi; this assertion would appear to be whollv untrue, except of
juveniles, in both scii'ijUiuj and cristam.
The dental formula is j-pj-i = 40. The mcisors form tightly packed transverse
rows, the outer teeth, both above and below, appreciably larger than the others, the
cutting edges ot the lower set benig slightly bitid or trihd. The canines are strong and
arc chiefly notable tor bearing verv narrow, shallow, longitudinal striations on their
outer faces. The lower canines are abruptly curved in the middle ot their posterior
taccs and thus have the appearance of being bent upwards. The premolars are ot the
usual form except that jfi in some cases has an additional cusp at the base of its inner
tace. When present it is, most commonly, well developed and quite obvious; but it is
sometimes reduced m size. Tliis tcature is in some measure diagnostic but is not com-
pletely reliable: in 16 :iduh gciicticiidcs skulls this supplementary cusp was present and
large in 9. small in 3, and absent from 4. Other species show something of the same
irregularity ot occurrence except tliienyi where the cusp occurred in all 6 available
skulls. There is nothing especially remarkable about the remainder of the teeth except
that 111- IS always the smallest m the two jaws and may sometimes be very much
reduced. Thomas & Wroughton (1910), writing ot stuhhnatmi, detected a marked
sexual ditference m the size of the upper carnassials, the temales being i mm or more
less in length. The comparative material trom West Atrica available in the British
GENETTA l8l
Museum is too sparse to furnish reliable data relating to this; but such as it is it tends
to confirm these authors' fmdings.
Habits. With the confusion that has existed over the different species of Geuetta
it is not practicable to relate the few field observations that exist precisely to named
forms, and the general account which follows, a compilation from various sources,
must take the place of the individual notes normally appropriate to specific sections.
It can fairly safely be assumed that all genets are broadly similar in their habits except
for such differences of detail as may arise from dissimilarity of habitat, high-forest
species being possibly rather more arboreal in their way of life than those from more
arid inland vegetation. It is, in some way, curious that so widespread and not uncommon
creatures should have had so little recorded of them in nature. This is partly due to
their secretive nocturnal lives; but they have long been kept in zoos and, within the
present writer's experience, as domestic pets; yet though there are fairly plcntifid
records of their littering nothing beyond the most meagre observations on other
aspects of their behaviour and way of life has, apparently, ever appeared in print.
Although exceptions, of course, always occur genets are pretty strictly nocrumal
and are therefore rarely seen except in the headlights of a car by or those who range
the bush at night with lamps. That they are from time to time seen and shot by local
hunters is obvious; and they may occasionally be accidentally flushed from some
da)time resting place in a tree or dense undergrowth; but the general run of skins and
skulls in the British Museum gives the impression that the animals must for the most
part have been taken in traps rather than killed by gun-fure. Experience with animals
maintained about the house shows that they do, under these circumstances, put in an
appearance about dusk; but even should they happen to be on the move in the forest
while there is still daylight their excellently disruptive coloration combined with
their stealthy caution would nearly always render theni exceedingly difficult to detect.
Since authentic first-hand records of these animals in the \\dld are almost entirely
lacking it is virtually impossible to make defmite assertions. However, they appear to
be rather solitary creatiu'es except possibly at mating periods, when they doubtless
forage in pairs. Even, at a later stage, the mother and her young do not openK' associate
for long, as they do in many other animals; for young genets, once past the earlv
helpless state that confmes them to the nest, quickly become capable of leading an
independent existence.
Genets are to a fairly considerable extent arboreal, their sharp, curved claws being
almost as well adapted as those of cats to scrambling up tree trunks. Some of their
foraging is regularly carried out in trees, for birds, possibly their eggs and certainly
their fledglings, for arboreal rodents, bats or the helpless young of larger tree-living
species. They are commonly found in oil palms. However, a good deal, possibly even
the major part, of their hunting is on the ground for groimd rodents and reptiles.
Like any other animals endowed with the power of swift climbing, genets take to
trees as a ready and dependable means of escape from threat on the groimd. Yet it
would seem that, the first instinctive urge apart, they appear on reflection to feel safer,
and with wider chances of escape, on the ground rather than limited to the trimk
and branches of a single tree; for after their first hurried rush up the bole they frequently
TS2 the carnivores of west AFRICA
make tor the ground again. Sanderson (1940) records that he onl\- once saw a genet
{crisutta) in a low tree; alarmed at his presence it returned to the ground in an attempt
to make its escape. He formed the opinion that the species was quite definitely terres-
trial. Stevenson-Hamilton (1947), too, said of a South African species that when pursued
it almost invariablv took to a tree but on being approached generally leapt to the ground
again to make oH with bounds ot remarkable length. He cited an instance of a genet
leaping from a height of 5 metres and landing 7 metres away.
In consonance with their at least partial arboreal existence they have been recorded
as sheltering or breeding in hollow trees or holes in trees. Loveridgc (in Lawrence &
Lovcridgc, 1953) tells ot shooting a genet in Mozambique "as it lay curled up beside a
hole high up on the trunk of a great baobab". Nevertheless, the majority of breeding
observations — and they arc not many — refer, rather, to holes in the groiuid. Possibly
this may be due to the greater likelihood of accidentally coming across a nesting
place in or on the ground than at some distance up a tree. Yet it is strange that those
collectors who have spent a considerable time in West Africa "smoking out" or felling
hollow tree stems in the search for bats or flying-squirrels seem never to have recorded
the incidental discovery of genets. Possibly the sort of restricted holes that might appeal
to genets as favourable breeding sites, such as small natural cavities wheie branches
have fallen away, or woodpecker borings, or honibill nests, are overlooked. The
tccKnique of smoking nevertheless commonly succeeds in South Africa (Stevenson-
Hamilton, 1947).
However, holes are not always essential and, in the dry-season at least, nests may
sometimes be exterior. One such was come across in February by an agricultural
labourer near Freetown and described to T. S. Jones as "a bundle of sticks". From this a
female genet had made her escape carrying one of her young in her mouth; the second
baby was then captured in the nest and subsequently successfully reared. That such a
breeding shelter was constructed by the genet itself seems improbable, and it is most
likely to have been a fortuitous conglomeration ot rubbish or, very possibly, a deserted
squirrel drey. Jones also came across another unusual oft-thc-ground refuge, namely
the roof of a house in Samaru (north Nigeria) from which he observed a family of
4 or s genets make its exit down a verandah post v^'hen it was nearly dark. This he
was told happened ever)- evening and that an adult pair dwelt permanently in the roof
(private comminncation). This home would appear to have been naturally discovered
and occupied by genuinely wild genets; but the present writer knew a more domes-
ticated single animal that, having been captured yomig and hand fed, also discovered
the roof as a warm shelter and spent its days concealed in a corner of the grass thatch,
descending regularly at dusk to the sitting-room to be fed.
flenets, then, at least sometimes, occupy holes in trees; and according to Stevenson-
Hamilton thev may rest stretched out along a branch. But the majority of records
refer to sleeping or breeding quarters on the ground; and in observations made on
captive genets in which they were afforded a choice of sleeping quarters Carpenter
(1970) certainly found them to prefer a hollowed out ant-heap on the ground to a
covered box fixed high up. In suitable terrain terrestrial shelters may be amongst
rocks; but thev are more usuallv in the smaller kinds of "earths", either naturallv
GENETTA I83
occurring holes or those made or improved by other animals — ground squirrels,
giant rats, porcupines, termites and so forth. They are almost certainly never self-
constructed, the genet legs and paws being ill-adapted to excavation. Besides earthen
holes, those amongst tree roots, or actually in rotted bases of trunks, or fallen, hollow
logs arc used; and temporary sleeping sites, though probably not breeding quarters,
are made ni thick grass, reeds or other dense low vegetation. The only actual first-hand
description of a nest appears to be that of Loveridge (1922) who came across one in a
hole in the ground consisting of a circular chamber with two bolt holes, unlined but
perfectly clean.
If events in captivity can be used as a guide, breeding takes place pretty regularly
twice a year. This, according to Verheyen (195 1) is probably at the beginning and
end of the rains — a supposition that is to a certain extent confirmed by juvenile West
African specimens oi genettoidcs in the British Museum collected in March, April,
June, July, September and early December. The usual litter size is 2 or 3 ; but i is not
imcommon, and 4 have been noted on several occasions, having occurred in no less
than three successive births in the Duisburg Zoo in Germany (Remy & Condc, 1962).
These latter authors have gathered together a considerable number of records, cliiefly
relating to the European genet; but the longest-continued observations of living
animals arc those made by Volf (1959 and 1964) in the Prague Zoo. He records that a
single female genet (geiictta) produced 32 young in 7J years, of which 20 were born
in the last 4 years of her life, the litters being almost without exception two a year.
This animal attained sexual maturity at the age of about 4 years and died, of old age,
at 13 years Such an age for genets in captivity is not uncommon, the record being
about two months short of 15 years (Crandall, 1964).
The period of gestation has been put at 10 to 11 weeks (Volf, 1959); but Remy &
Conde (1962) mention an American figure (Smithsonian) of only a little over 8 weeks.
A well-authenticated period observed in Paris was 68 days, that is, just on 10 weeks.
The young are born blind and with the auditory meatus covered; Volf (1959) observed
the ears to open on the 5th day and the eyes on the 8th. According to the same author
the newly-boni kittens are well-haired though this covering is of a deeper, wholly
greyish, hue than that of adults, to whose rather paler and warmer colourmg they do
not approximate until after the second month. In the British Museum there is a very
dark, almost melanistic juvenile oi genettoides. Writing of the European genet Volf
said that at birth there are only 3 rings near the base of the tail, the whole dorsal aspect
of the tail being dark, the underside paler. With advancing growth the pale colour
encroaches on the dark and at the same time forms more rings, which reach 6 in
number at three weeks. Weight at birth in this species is from 61 to 82 grams. This
increases vciy rapidly, especially in the first few days, doubling itself in 12 days and
quadrupling in a month. The growth curve is more or less a straight line; and at two
years the fully adult weight is of the order of 2 kilograms. This may increase slightly
with greater maturit)'. It has been found in captivity that immediately after birth the
male must be removed for two or three weeks as he may otlierwise kill the young.
But according to Verheyen (195 1) in the wild the male actually provides food for the
lS4 THE CARNMVORES Of- WEST AFRICA
resting and nursing motlacr. Killings of their ncvvlv-borji kittens arc, for unknown
reasons, not infrequently carried out bv the females.
Volf (1959) provides data tor the eiuption ot teeth, given below. It is, however,
not clear whether reference is to milk or permanent dentition; it may be presumed to
be the former, yet the premolars which emerge subsequently to the molars on the
44th and .SI St days would seem to be replacements of an earlier milk set. Volf observed
tlie upper incisors to appear on the 23rd day, at which date, also, crawling commenced
and the ability to raise the head. All the incisors and both upper and lower canines
enipted by the 30th day; the molars appeared on the 37th day; the upper premolars
on the 44th, and the lower premolars on the STst. The young animals attempted at
5 weeks to detend themselves bv spitting; and after N weeks bv biting. At this age
they were fully active and weaning took place, and thev were capable ot taking care
of themselves. A certain amoimt ot solid tood is in tact accepted trom about 6 weeks.
Loveridge (1922) foimd that a babv he attempted to bring up would not look at meat
but only drank milk sweetened with sugar to the consistency of treacle; and it became
very fond of jam and would eat egg, after shelling. Mrs. Soniajetfrey (private com-
munication) toiuid that a baby genet could be successtully reared on tinned imsweetened
milk with an occasional mixed vitamin tablet. This animal showed an interest in red
meat when it weighed 255 grammes and ate its first mouse three weeks later when it
weighed 310 grammes.
Although the shift to solid food normally takes place at about 2 months. }-oiuig
genets, if given the opportunity may continue to suck up to the age ot 6 months. In
captivity, Volf records, the yoiuig of one litter were usuallv removed trom the mother
before her next pregnancy. Where this, on one occasion, was not done the mother,
through over-long continued suckling, reached a state of exiiaustion in which the
foetuses failed to develop properly and the yoiuig were either still-born or died soon
after birth. It is interesting to note that although Volf estimated that the iiciicna tcmalc
which mothered 32 young must have been about 4 years old before she became sexually
mature, he found the oifspring to attain this state after about only 2 years.
It has sometimes been held that genets, unlike many other flesh-eaters, are very
strictly carnivorous in their diet, firmly rejecting all kinds of vegetable toods. This is
not so. Dr. Gordon Corbet shot a genet in Kenya that had its whole intestine crammed
with a mass of orange-coloured berries, besides the remains of a .spider and insects
(private commimication). From the description it would seem that the truits were almost
eertanily those of the so-called Jujube Tree, Ziziplms iiuwriiiaihi Lam., widely spread
ill the more arid woodlands of tropical Africa. Another very interesting observation
concernmg teeding on vegetable material has recently been made by Sonia Jeffrey in
Ghana (private communication). She found that something like 70 per cent of the
contents of one stomach consisted of rotten wood, some of the pieces being up to 18 mm
long. She supposed that this indicated that the animal had really been alter some more
tasty delicacy buried in the wood. Such an explanation is very possible since beetle
grubs, sometimes ot large size, frequently abound in dead trees, as woodpeckers well
know. The fact that genets are often connected with palm trees leads to the speculation
GENETTA 185
that they may, at least sometimes, be attracted not only by the fruits but also by the
very large grubs of the giant palm-weevil foimd in decaying stems.
As concerns the more usual, carnivorous, foods Shortridge (1934) wrote that genets
prefer fresh meat; but Lovcridge found tinned corned beef to be quite acceptable;
and Stevenson-Hamilton (1947) says they arc partial to carrion and readily enter
baited traps. Besides rats, gcrbils, squirrels and hares they take insects, especially locusts
and beetles, and will tackle birds of the size of guinea-fowl and domestic breeds. They
also eat lizards; while Loveridgc (in Lawrence & Loveridge, 1953) found the remains
of a burrowing snake and a large yellow scorpion in the stomach of one. Whether
they eat amphibians and molluscs seems to be imrecorded. They reputedly eat shrews;
but Sonia Jeffrey kept a young genet that would eat most kinds of meat but would
not look at a shrew. Although one of her captive specimens would not touch a freshly
killed snake another one readily ate them. Stomach contents of genets examined by
her included the legs of skinks and insects, and orthoptera wings ; and one contained
two mice \\hich may, from the red colour of the fur, have been Lophiiwmys sikapiisi.
One of her live specimens pounced on and ate a bird that flew into the house.
In their attacks on fowl-runs they show, according to Stevenson-Hamilton, con-
siderable cunning, generally taking a single bird at a time, returning for another when
this has been consumed. A bird of this size is attacked at the throat, the blood sucked,
and then the upper part of the breast taken. Carpenter (1970) found some evidence
that males were more often poultry thieves than females; a:id that once a genet had
acquired a taste for domestic fowls it tended to disregard more natural prey. In the
absence of such an acquired taste rodents were nearly always preferred as food to birds,
being pounced upon and killed forthwith, while birds were only killed and taken later if
the rodent supply were insufticient. On two widely separate occasions Carpenter foimd
genets to capture bats in roofs, the genera concerned being Eptesiais, Scotophilus and
Rhiuolophus. Over a period of five nights one genet took an average of six bats a night.
Genets are not normally offensive; but when alarmed spit and bare their teeth,
and if hard-pressed can give a severe bite. Like other animals they vary a good deal in
temperament and it is difficult to generalize. They can be tamed, especially when
captured very yoiuig; and some become fairly trusting and even milldy friendly,
though never to the extent of mongooses and other small mammals. They seem to be
ver)' much one-man pets and to resent strangers. The majorit)', at least imdcr con-
ditions of semi-freedom in Africa, as a rule soon manifest independence; and though
they may appear in the evening to take what food is offered they mostly remain rather
coldly self-sutiicient, exhibiting the aloofness of a cat with none of its friendly domes-
ticity. Genets are also doubtful assets; for while it is true that they may keep the house
free of rats and possibly snakes they have no loyalty when it comes to a question of
robbing their owners of meat from the pantry or effecting a midnight entr)' to the
fowl-house.
Genets emit a musky odour, though not so strong as that of some other viverrines.
Gray (1830), in his description oi iiiaaihua as a living animal in the Tower of London
menagerie, said that it had an exceedingly strong musky smell and was continualK-
placing its two hindlcgs against the wall of its cage, and pressing the subanal glands
l86 THE CARNIVORES OF WEST AFRICA
against it, leaving two chocolate brown musky streaks. These were far larger than
those of a similarly captive common civet, which performed the same action; and it
seemed, theretore, that more secretion was emitted by the genet than by the bigger
animal. Both were obviously attempting to demarcate their territories; and it may be
assumed that something of this sort is regularly carried out under natural conditions,
though, in the wild, urination and defaecation arc probably the most usual method
iUid adequate to the purpose. Genets have not usually been reckoned amongst the
animals that use fixed latrine sites but Carpenter (1970), working in Natal, found these
to be not uncommon, generally in dry river beds or thick bush, some of them obviously
in use for a considerable time. As regards territorial behaviour and the instinct and
ability to return to the home range the same author has made some very interesting
observations and deductions in relation to the rusty-spotted genet in South Africa.
From these it would appear that females are more attached to a given territory than
males, which are more inclined to wander. By repeated trapping and marking, these
genets have been shown to return to their original home ground trom distances up to
about 30 kilometres.
Taxonomy. Forty years ago Schwarz (1930) in his introductory remarks to what
has proved to be the last published general review ot this genus expressed the opinion
that its systematics were obscure. This is, to say the least, somewhat ot an under-
statement. As it stands today Gciwtta is a confusion of forms, with the definitions of the
classic species uncertain and the status of later creations — at times recognised as valid
species, at others bandied about as mere races betwixt one reputed species and another —
a tangled skein with no clear guiding threads giving hope of any gcnerallv acceptable
unravelment Over the years no less than 44 discrete species have been described in
Africa, apart from several forms rated from the start as racial. Most of these have now
bceji either reduced in status or synonymised, and G. M. Allen in his Checklist of
African Mammals (1939) recognised only 6 species and 27 subspecies. This is far from
being the last word.
There is something inherently wrong in a situation wherein authors so frequently
disagree over the status or validity of forms and their opinions regarding their essential
diagnostic characters; or have changed their own minds in these matters; and in
which nmseum labels so manifestly bear witness to uncertainty in determination.
Two plain facts emerge: the original descriptions have, in the light of greater know-
ledge and more abundant material, often been pitifully inadequate; and the taxonomic
significance of the characters chiefly relied upon for diagnosis has been greatly over-
valued since most suffer the disability of considerable idiosyncratic variation. As long
ago as 1910 Thomas & Wroughton, writing o( stithlmanni and siiahelica, two of Mat-
schie's creations, said: ". . . we have a series of over a dozen, the extreme individuals of
which are easily separable; but these extremes are linked up by the intervening in-
dividuals in such a way that after most careful examination of both skins and skulls
we have been obliged to acknowledge that we cannot fmd any constant character by
which these forms may be separated . . .".
It is possible to compile a long list of characters that have been used taxonomically,
but thev can be briefly categorised as the nature of the pelage, its colour in various
GBNBTTA I87
situations on the body, and the colour, size, number and shape of the markings. These,
and others, can build up into a very large number of combinations; and if, as has been
the tendency, each of these is regarded as constituting a valid form there is considerable
scope for taxonomic chaos. The difficulty is to decide which characters can, in fact,
be relied upon, and to what degree. This can only be determined from long series of
restricted provenance. One such, and the best for which published information exists,
is the series of 46 specimens, 24 adults and 22 immature, collected by the American
Museum expedition to the north-east Congo, and assigned to G. pardina ficldiana.
How careful the taxonoinist must be, and how far more limited than was originally
supposed are his morphological resources in Gaiclta, are indicated by J. A. Allen's
(1924) analysis of this collection: "This species, like its congeners, presents, when
adult, a wide range of purely individual variation, not only in size and coloration but in
cranial characters and in the teeth, especially in the size and form of m^ ... In coloration
the variation from the norm is toward, on the one hand, an extreme gray phase with
blackish markings, on the other, a rufous phase with deep brownish buff instead of a
gray ground color and dark brown markings (black strongly mixed with rufous).
The dark tail-rings are black or blackish in both; the light tail-rings are much lighter
in the gray extreme than in the rufous extreme, being white or whitish in the former
and strongly suffused above with pale rufous in the latter. The light tail-rings are
usually seven, but vary in number from six to eight, besides the terminal half ring,
broken by the black of the upper side of the apical portion of the tail. The light rings
are usually much broader on the sides and under surface of the tail than on the mid-
dorsal line, where in some specimens they are nearly obsolete, especially beyond the
fourth firom the base. The light rings are occasionally as wide as the adjoining dark
ones, but usually somewhat narrower, and frequently only about half the breadch of
the dark ones. The black tail tip varies in extent (measured from the last full ring on
the dorsal side) from 70 to 150 mm (in one specimen 220 mm) ... It is unnecessary
to describe in detail the irregularities in the size, number of rows and the arrangement
of the spots on the sides of the body, since they are more or less different in each speci-
men, and often different on the two sides of the same specimen. Neither is it necessary
to more than note that the relative width of the light and dark tail-rings is exceedingly
unstable and hence has no taxonomic value. Yet such inconstant features were once
made the basis of an elaborate synopsis of the species of the genus Getietta, noteworthy
mainly for its puerility and pernicious results ..." The synopsis referred to, in which
over thirty different species are distinguished, was that of Matschie (1902). Apart
from such idiosyncratic variation as demonstrated by this Congo, and other, series,
the intrinsic mutability and untrustworthincss of external characters may be further
judged from the experience of Schwarz (1930) who, in 20 years' study of the genet
question, had observed animals in captivity to manifest, as the result of enviroimiental
change, such alteration of appearance as might rank of specific worth.
Matschie aside, since the majority of forms have been named purely firom external
characters and firequently from single specimens, not always abult, it will be appreciated
that there has been considerable scope for confusion and misunberstanding. Moreover,
there has been some tendency to assume that because a specimen berives firom the
l88 THt CARNIVORES OF WEST AFRICA
locality broadly luuncd as that troni which the type was reputed to come it was in
tact that species, and on this basis invest the species with characters not truly pertijient
to it. And authors have not infrequently fallen into the trap of assuming that characters
present in one or two specimens, such as black forclesrs or an inner cusp on p'^, arc
diagnostically valid.
The tindings of Schwarz in his 1930 review ot the genus, have, to all intents and
purposes, formed the basis of all subsequent accounts. Unfortunately his diagnostic
characters do not always stand up to examination, being, in the light of more abundant
material, sometimes misleading, sometimes frankly inaccurate. These matters are
dealt with in the course of the species descriptions which follow. Meanwhile, it mav
be said that it seems almost impossible to build any lasting structure on the existing
shaky foiuidations — shaky because of the ill-definition of manv of the reputed forms
including some at least of the early classic species. An attempt has been made in the
present account to go back to first beginnings and to draw attention to some of the
difficulties and misconceptions which have since arisen. In the consequent arrangement
ot West African forms use is made ot species-groups, not trom any great conviction
but as a convenient way of indicating probable relationships at the upper levels whilst
leaving the field clear at lower levels for the subsequent indication of colour and other
variations which imdoubtedly often exist in the groups here designated species and
which may later be thought worthy of nominal distinction. Two subgenera, besides
the nominate one, have recently been proposed, Psciido^ciictta Dekeyser and Paragcncita
Kulm. The former has here been rejected; the latter retained, though no cranial material,
on which its validity depends, has been examined. This leaves the provisional organiza-
tion of the genus in "West Africa as 8 discrete species divided between 4 species-groups
in the typical subgenus, with a 9th, single, aberrant species in the second subgenus.
The following facts and opinions are here given for what they are worth in casting
possible light on relationships in this genus. Sympatry, and hence presumed specific
isolation, is, so far as can be assumed from non-existent or limited habitat data, demon-
strated by British Museum specimens between senegalensis, pardina and thiciryi in the
Wukruni Hills (Nigeria); between pardina and cristata in the Mamfc district of Camer-
oun; ajid between poaisis and johiistciii in eastern Liberia. However, crosses are known
to be possible in the genus, though no investigation has been made into the fertility
of the offspring. Zuckerman (1953), recording the known cases of deliberate hybridiza-
tion in zoos, lists a cross, in 1858, between gcnetta and sencgaleiisis; in 1859 between
(>cuctta and tiq^riua, 3 young being born; and in 1885 between gcnctta and pardina,
resulting in the birth of a single kitten. Finally, Pocock (1915a), from study of the
anatomy of the scent glands, came to the conclusion that two groups of related forms
could be recognised: furst tigrina, pardina and rnhifjiiiosa; and second (,'t7)(7fi;, dongolana
(= senciialensis) and fclina.
It may be as well to offer a word ot warning. In view of the diversity of opinion
held over the years amongst authors, zoo superintendents or museum taxonomists
regarding the identity of different forms, the naming of some, if not all, of the above
species should be regarded with a certain amount ot caution. This applies to the
literature m general.
GENETTA
KEY TO THE SPECIES OF GENETTA
(Previous key page i66)
1. A medial dorsal stripe present, frequently longer-haired than the neigh-
bouring pelage, mostly intense black and nearly always (except pociisis)
contrasting by reason of its continuity and/or colour with the, usually,
browner and broken rows of spots. Condylobasal length of adult skulls
over So mm. (Exceptional skins giving rise to doubt will not conform to
the following alternative key characters) ...... 2
Spinal stripe either obscure or, more generally, present but never black, very
narrow but nevertheless nearly always more or less clearly split longi-
tudinally into two by a streak of pale hairs ; the whole, usually clear-cut,
dorsal pattern of spots or lines individually very narrow (mostly under
10 mm wide), and generally ginger-brown on a huffish ground. Cond)'lo-
basal length of adult skull under 80 mm . . thierryi (page 214)
2. The back of the neck with a small crest ot erect hairs, or the hairs on its anterior
part directed forwards and parted posteriorly from the backwardly
directed remainder by a whorl. Tail soft and woolly, the bristle-hairs not
much exceeding the underfur in length, clearly ringed, both above and
below, from root to tip. Skull with no sagittal crest on the anterior part
of the braincasc; the postorbital constriction nearly always less than the
uiterorbital breadth; an internal cusp mostly present on p^ . . . 3
No nuchal crest or other abnormality of tlie fur of the neck ... 4
3. Back of the neck with a short erect crest; background colour of the dorsal
pelage mostlv pale huffish. ..... cristata {page 198)
Back of the neck without a crest but with the hairs directed forwards, and a
whorl; background colour ot the dorsal pelage a warm golden-brown
bini (page 200)
4. Tail densely woolly and soft, the abundant underfur dominating and being
almost as long as the rather narrow, relatively inconspicuous bristle-hairs;
ringed pretty clearly to the tip. Mandible rather flat, with the incisors and
canines directed forwards; p* about 6-6 mm long, in^ about 5-6 mm
broad ........ johnstoni (page 217)
If the tail is ringed clearly to the tip it is not sott and woolly. Mandible not
flattish, teeth larger .......... 5
5. Tail ringed to the tip and clad with ven,' long bristle-hairs which completely
dominate and far exceed in length the relatively insignificant underfur,
and is hence rather harsh. The postorbital constriction greater than the
interorbital breadth; a complete sagittal crest mostly absent ... 6
The soft tail, in which the abiuidant underfur is not completely dominated
by the bristle-hairs, has at least 75 mm, and generally far more, of the
distal end entirely black or with only slight indications below of pale rings.
Skull with the postorbital constriction almost invariably less than the in-
terorbital breadth; a complete sagittal crest present at least in old males 7
190 THE CARNIVORES OF WEST AFRICA
6. Larger and grcvcr; head & body about 490 nun; condylobasal length about
93 mm; dorsal ground-colour rather cold whitish-grey; pale rings of the
tail with relativelv little sandy-brown adulterating the white; top of the
hindtoot very pale grey, sharply demarcated from the intense black of
the underside, which spreads round the back of the heel genetta {page 191 )
Smaller and butl'ier; head & body about 440 mm; condylobasal length about
S5 mm; dorsal ground-colour a warmer buft; pale rings of the tail
almost entirely sandy-brown above, pure white below; hindtoot almost
pure w^hite above, black below but without the black spreading con-
spicuously romid the back of the leg . . . senegalensis {pdgc ig})
7. Size small, head & body estimated at about 450 mm; spots fairly large and
separate, brownish on a pale whitish-buff ground-colour; tail clearly
ringed white and black with only a relatively short all-black distal
portion ......... pardiiia (page 209)
Size larger, head & body about 500 mm; tail much more obscurely ringed,
the pale annulations being often much adulterated with darker colour,
even those near the base; a long, sometimes very long, distal portion
wholly black or indistinctly marked below with partial pale rings. . 8
N. Spots large and few, wholly black or annulate, mostly separate, in about
4 rows, on a dull butfy-grey ground-colour getiettoides (piigc 210)
Spots small and numerous, in about 5 or 6 rows, wholly black, those adjacent
to the spinal stripe coalescing into longitudinal bands; ground-colour
much as in ijt';R'»()i'(/«; tail predominantly black . . poensis (pa(;c 212)
A. gennetta Group
General. There are two members of this group occurring in West Africa: what
appears to be qciictta itself and saicgalcnsis. The latter is undoubtedly closely related to
the former, sharing with it a number of fundamental characters; but it is almost
certainly wrong to regard it as nothing more than a race. Many years ago, Riippell
(11S36) expressed the firm opinion that a single species ranged from southern Europe
to the Cape, differences in the size, shape and colour of the spots being nothing else
but climatic variations of one and the same animal, the skulls all being the same.
While it is not possible to make the sweeping inclusions that Riippell visualised there
IS nevertheless a basic truth in his assertion ; and though they are of no immediate
concern to West Africa it is perhaps worth recording the opinion that both the extra-
limital ligritia and fcliiia, in so far as they can be identified from their poor diagnoses,
are more closely allied with the group now under consideration than with any other.
At any rate, specimens from southern Africa in the British Museum which have in
the past been ascribed to fi'liua seem, with very little question, to he (leiictta.
Dcscriprion. The characters which define this group, and which very readily
differentiate its members at sight from the others are these. The dorsal pelage is very
long, though in the tropical forms it is less so but nevertheless still appreciably longer
than in the other groups; most typically it is 30 mm or more long; but if the forefmger
Senegal Genet {Ccncu, ^cncf.la,^i^: Hansa tienet (Ccucu., ,hn-rry,): Crested Genet (Cccm mst.,.,)
GENETTA GENETTA IPI
is run up backwards through the pelage the latter is seen to be at least as long as, and
usually considerably longer than the depth of this digit. From about the mid-back to
the root of the tail is a contijiuous, yet-longer-haired, spinal stripe, almost invariably
black but sometimes dark browai. This is erectile.
The tail is vcr)' long-haired, generally noticeably wider at the root than at the tip;
it is almost always clearly amiulatcd throughout its length, the margins ot the rings
not sharply dcfmed owing to the partial overlap of the long bristle-hairs of the next
proximal ring: there arc 8 to lO dark rings, and the tip in West African forms is white
or with only a slight sprinkling of black. The dorsal aspect of the tail has the pale rings,
either wholly or at least along a medial line, washed with red-brown; but below they
arc more or less pure white. The general appearance is of a rather rough, wir)' structure
due to the fact that the very long bristle-hairs far exceed in length the relatively in-
significant underfur. In this the group differs abruptly from pardina and servalina.
Neither the colour nor the spots are diagnostic of the group. The groimd-colour
varies from a pale creamy white to a colder light grey; the spots may be light brown
or almost black of uniform colour or, in the case of the darker ones, slightly dusted
over with red-brown. They vary in size from very small (lo x 5 mm) to moderately
large (40 x 30 mm), oblong or roundish, set in five rows on each side of the spine,
the fifth, or lowest, row being frequently much less well defmed or incomplete. The
spots are for the most part clearly separate, but sometimes, especially over the rump
and in the line or two adjacent to the spine, tend to run together longitudinally.
The forelegs are, in West Africa, pale above but sometimes darkened on their inner
and lower surfaces. At least the foot and sometimes the whole hindleg is pale on the
upper side and dark below.
Skull. In the skull, the intertemporal constriction is either about equal to or, more
generally, greater than the interorbital breadth. There may or may not be a sagttal
crest, p* generally has a large inner cusp, but this is sometimes lacking.
Taxonomy. Two forms are thought to exist in West Africa. Though most authors
concur in thinking that the various forms o( genetta are closely related there is some
disagreement regarding the taxonomic level of naming. Schwarz (1930) dealt with
them as conspecific but separable into (imdefmed) subspecific groups; thus ajra and
senegaletisis, though assigned to different groups, become nomenclaturally G. genetta
afra and G. genetta seiiegalensis. On the other hand Cabral (1966), while admitting that
this may be the case, has suggested that the G. geiwtta group is in fact a superspecies
comprising three independent species, the above forms becoming ascribed to G.
genetta and G. sencgalensis. Something of this latter view is adopted here, using the term
species-groups rather than superspecies.
The two West African species may be separated by the key given on page 189.
It should be noted that the characters there given are not necessarily of extralimital
application.
GENETTA GENETTA (Linnaeus) European Genet
Viverra genetta Linnaeus, 1758, Systema Naturae, loth ed. i: 45. Spain.
Genetta afra F. Cuvier, 1825, in GeofFroy, E. & Cuvier, F. Hisloire NatureUe des Mammijcres . . . , pt. 52,
192 nil; CARNIVORES OF WEST AFRICA
pi. 195 and pt. 51, text. Barbary. The specific name is Latin tor African. Here taken as a valid race.
Gi'iictta i'»(i;(iri.': Lesson, 1S27, Mamie! df Manimalc^iv, 011 hisloire natmcUc des Mammijires: 173. A renaming
of V. ^vih'llu Linnaeus. I 'i/Zijiir/s is the Latin for common.
Giitctta barbnra Hamilton Smith, 1842, in Jurdittc's Naturalist' s Library: 35(13 of Mammalia): 171. Barbary,
which the Latin specific name indicates. This is most probably svnonymoiis with afra, a form of which
Hamilton Smith was, apparently, ignorant.
There are other names ot purely extralimital mterest.
This genet, most typically from the Mediterranean region and originally described
trom south-western Europe is, in all likelihood, an introduction into the latter area
trom northern Africa. The species is of disputed range within the latter continent.
Ellermaii, Morrison-Scott & Hayman (1953), for example, following Schwarz (1930)
regarded Iclina and other forms from the south as races of (jtvu'Wii, whereas Roberts
(1951) excluded this species altogether from his account of South African mammals,
according it no mention whatsoever.
However, the existence oi genetta as a species in the southern halt ot the continent
is ot no immediate concern to this present study. Its range in northern Africa is. It is
generally agreed that gciictta, in the form afra, occurs throughout the Mediterranean
fringe of Africa trom Libya to Morocco; and it is known also that its range extends
down the Atlantic coast to the south-west at least as far as Mogador (Cabrera, 1932).
How much further it ranges down this coast does not seem to be recorded.
The assumption in this present work that it extends at least as far as Thies in Senegal
IS based on a single British Museum specimen, No. 9.1 1.2.34. f" view of the pro-
venance of this specimen it was labelled saicoalcfisis. In fact, it bears extremely little
resemblance to Cuvier & Geoftroy's plate and description trom which Fischer derived
his diagnosis of this species; on the other hand it does correspond fairly well to their
plate of afra and to other British Museum specimens of this deriving from northern
Africa.
Genetta genetta afra F. Cuvier North Atrican Genet
The range ot pattern and size ot this animal m western Africa cannot, ot course,
be told trom the single above mentioned specimen available. The pelage is long and
rather harsh, its superficial ground-colour pale grey adulterated with very light brown
and a little black, both derived, rcspectivelv, trom the subterminal zones and the
tips of the bristle-hairs. The markings superimposed on this background arc deep
blackish-brown with a slight over-wash ot yellowish-brown hairs. There is a very
deep brown, almost black, spinal crest of long hairs from just short of the shoulders
to the root of the tail. On either side of this the spots are set in four clear longitudinal
rows with a faint indication of an incomplete tifth. The spots of the row next to the
spinal crest are rather irregular in shape, on the whole rather narrow, and tend to join;
all the rest of the spots are to all intents and purposes independent. Those ot the second
row are the largest, the anterior four being 15 to 20 mm wide and the biggest some
40 mm long. The base of the fur is darkish grey; and there is a great abmidance of very
long, fine underfur (about 13 to 15 mm) amongst which the long (38 to 42 mm)
GENETTA SENEGALENSIS 193
bristles tliat form the outer contour of the coat stand fairly widely spaced at the base
though they come together closely distally to form a continuous cover to the whole
body.
The tail is rather shorter than the body, long and coarse haired, with eight dark rings,
the basal two narrow and rather indistinct ajid a very small black tip. The bristles
measure about 50 to 60 mm near the root; the undcrfur only 14 to 17 mm. The long
hairs of one ring overlie the bases of the hairs of the next more distal ring for some
considerable length so that the lines of demarcation between the rings arc not at all
sharp.
The forelegs are pale, only slightly spotted; the hindfeet are whitish above in very
sharp contrast to the black of the outer aspect of the toot and the back of the ankle
joint. The face has the usual genet markings: a white patch under the eyes separated
by a blackish patch from the white area beside and below the rhinarium. The chin is
white centrally but is bordered by blackish lower lips.
Skull. This in the single specimen available, a moderate-aged male, has a low, sharp
sagittal crest extending the whole length of the cranium. The intertemporal con-
striction is a trifle wider than the interorbital breadth, the postorbital processes short,
with sharp points, p^ has a large, sharp, internal cusp; the antcro-extemal cusp o£ p* is
bifid; 1112 is fairly large, quadrate, with 4 clear cusps. The bulla is shown in fig. 28a.
The measurements are given in the table on page 219.
GENETTA SENEGALENSIS (J. B. Fischer) Senegal Genet
Viveira sciu'galcmis]. B. Fischer, 1829, Synopsis Mammatiuitr. 170. Senegal.
Vivcrra don^olana Heniprich & Ehrenberg, 1833 [fide Sherbom), Symholac Physkae scti Icoiies et Des-
criptioncs Mainmaliiim . . . , dec. 2, folio k: 2. Dongola, Sudan.
Viverra kptura Reichenbach, 1836, Rcgnuin Aiiimalc, I: 23, f. 270.
This has been variously regarded as a discrete species or as merely a subspecies of
genetta, a matter that has been discussed above. The exact, or even approximate,
locality in Senegal from which this animal was first described is imknown.
Taxonomy. Fischer seems fairly obviously never to have seen this genet but to
have framed his diagnosis from Cuvier's description and plate (in Geotiroy & Cuvier's
Histoire Naturelle dcs Alammiferes . . .) published in December 1821. It was called by
them the Genette du Senegal but not given a binomial Latin designation. The scientific
name and description must therefore officially date from Fischer, 1829, though, as in
the parallel case of G. tigrina, the diagnosis is not first-hand but extracted from an
earlier work.
Since Fischer's diagnosis is nothing more than a Latin description ot Gcoifro)' &
Cuvier's plate the characters originally ascribed to senegalcnsis can best be gathered from
this last and its accompanying notes. The species is shown as a considerably paler
animal than ajra, almost cream in colour, the black spots being in only 4 rows either
side of the spinal stripe. Each of the innermost two rows consists of 4 spots only,
long and very narrow and quite separate, the spots of the two outer rows being shorter
and more rounded. The tail is shown as having 10 black rings and a white tip; the
194 THE CARNIVORES Of WEST AFRICA
hiiidlcg as having a sharply cut oft' black mark above ajid on the angle. The chin was
said to be black; the coat not remarkably long; but the hairs of the tail were, making
it look bigger than it really was. In Cuvier's original description of the Senegal genet
(Livraison XXV) the tail was given as the same length as the body (i.e. about 500 mm) ;
but later, in the second description of ajra (Livraison LIl) it was mentioned incidentally
as being about 175 mm longer, the proportions in the two species being thus very
different.
Although at one time and another a number of specimens in the British Museum
have been ascribed to scncgalensis there is none that at all convincingly corresponds to
the very pale animal with widely separate, sharply defined, dead black, linear markings
illustrated by Gcoftroy & Cuvier. The nearest seems to be a doiigolana specimen
from Shendy (Sudan), No. 1939. 1768, in which the spots are brown and roundish
but slightly confluent, making them appear long and narrow The tail-rings are 10,
not very clear-cut; the tip white, though mixed with a little blackish. The black mark
on the hindleg is fairly abruptly defined. The pelage is not markedly long but there is a
long shiny black crest from the mid-back.
Hemprich & Ehrenberg's description ot dong^clatm is limited in scope, and a good
deal ot it is irrelevant in being a comparison with the purely Madagascan Fossa. How-
ever, they state that doiigolana is a whitish, not a grey, form, with a blackish line from
the middle ot the back and black spots overlain with cimiamon, there being no mention
ot their shape, size or arrangement. The tail is described as differing verv widely from
gcnctta and "'singularly short", the number of rings unstated.
The specimen. No. 1939. 1768, mentioned above as most closely representing sene-
galaisis corresponds pretty well to this sketchy description and there seems little doubt
that the two torms are really one and the same.
Distribution. This being so, we have a range tor scncgaknsis troni Senegal to the
Nile and beyond (Setzer, 1956) to the Red Sea, that is to say transcontinentally through
the Sahel and Subdesert zones of vegetation. Actually, as will shortly be seen, it occurs
also further south in tlie Sudan and Doka zones. Specimens exist in the British Museum
from Aoudcras (Air — Subdesert) ; Fort Lamy (Chad — Sahel woodland) ; Zuarangu
(Ghana — Sudan woodland) ; Famiso and Kabvvir (Nigeria — Sudan woodland) ; Zaria
district and Wukrum Hills (Nigeria — Doka woodland). It is also probably this species
which A. J. Hopson (private communication) observed to be trequent in creeper-hung
trees in the Yobe Valley, near Yo, Lake Chad (Sahel).
Description. The Senegal genet, as represented by these specimens (Plate 3),
is a distinctly pale animal with a background colour ot whitish, cream or buftish,
and spots that arc mostly light or medium brown but more rarely blackish-brown.
They vary in size but are never large, rarely having a maximum diameter ot more
than 20 mm and usually less. One unusual example from the Wukrum Hills has very
small spots, bearing the same relationship of pattern to the other skins as the servaline
does to the serval. They arc also very variable in shape but could be described in
general as irregularly roimd or oval, occasionally oblong; and though here and there,
chiefly over the rump, joined together, they are in general sufticiently widely separated
for there to be as much or more ground-colour as spots showing. In this last, as in
GENETTA SENEOALENSIS
195
Fig. 29. Cenetla seiiegatensis: skull, B.M. No. 1939. 1766, (J, X i
Other ways, they differ from the small-spotted servalina in which the spots dominate
the ground-colour.
The pelage, although considerably shorter than in genctta, is nevertheless still long
by comparison with other West African species, the bristles, in typical non-moulting
specimens, measuring about 25 to 30 mm over the rump, the underfur about 12 to
14 mm. There is a long-bristled spinal crest from just posterior of the shoulders to the
196 THE CABNIVOHES OF WEST AFIUC:A
root ot tlic t.ul. rhis IS almost al\va)s black but in some cases merely deep brown.
The distinctive character of the tail unites all these skins and shows their affinity to
i^ctictru. It is long-haired, the coarse, terete bristles measure trom 45 to ss mm near the
root, the uiiderhir 13 to 15 mm. As in all this group, the annulation, though mostly
quite clear, is not sharp-margined because of the overlap ot the long bristles. For this
reason the basal two or three rings are difficult to determine; and this, in some measure,
accounts for differing opinions regarding the number of rings. The most usual number
of dark rings is 10, though in some cases there seem to be only 9. The tip of the tail is
white but often has a tew black-tipped hairs mixed with it. The dark rings may be
black or deep brown; the pale rings are always pure white below, but dorsally they
generally have a good deal of light-brown mixed with them, especially medially,
or are entirely golden-brown. The length of the tail seems to be irregular. Only 5 skins
have field measurements; and in these, two tails measure slightly less than head &
body, one appreciably less, and two appreciably more.
The forelegs are pale above and blackish below. The hindteet are white above;
the legs arc blackish below and on the outside and back ot the lower leg (tibia) but not
usually entirely aroiuid the ankle as mgeiictta ajra. The usual genet white tacial markings
are present — below the eye, around the rhinarium and between the eyes. The chin is
always white medially, bordered by a broader or narrower black area running along
the lower lips.
From the limited figures available and trom the general appearance ot the skins this
genet would seem to be smaller than the animals living in northeni Atrica and Europe
((;cuiitd). The actual mean measurements ot British Museum specimens will be tound
in the table on page 219.
Skull (tig. 29). Seven skulls, of which only tive are mature, exist 111 London. Though
the latter are pretty even in size they show, as throughout this genus, contusingly
irregular characters, the various inconsistencies being distributed over all the specimens,
irrespective of sex, and not confined to one or two exceptional examples. In two temales
and one male there is no trace of a sagittal crest other than a posterior portion adjoining
the supraoccipital crest; in one other oldish male there is a clear, but very low, crest
across the whole length of the cranium; and in the third, oldish, male such a rudi-
mentary crest is partly present. In tour ot the specimens p'^ has a large internal cusp;
but from the fifth, a medium-aged tcmale, all trace of this is completely lacking. In
this skull, also, in- is very much reduced in size. The development ot the postorbital
processes varies trom long points to fairly short ones. One lower jaw has a large
quadrangular, four-cusped iiij.: but in the remainder this tooth is triangular and thiee-
cusped.
B. servalina Group
The animals included here are usually regarded as a single species covering tour
described subspecies; but the two forms occurring in West Atrica seem to be sufficiently
distinct to constitute two discrete though, as in the case of (jc/dVAi, very closely connected
species. This will be dealt with more fully below in the taxonomic section.
GENETTA SERVALINA 197
Description. There is little room for doubt in tlie determination of the scrvaline
genets. The characters, especially in comparison with genetta are very distinctive.,
The dorsal pelage is short compared witli that of typical genetta, though not markedly
shorter than in seiiegalciisis; but it has none ot the harslmcss of the genetta group, being
instead rather silky. This is because ot the quite different form of the bristle-hairs (20
to 25 mm long) which, in this group, arc only expanded for the distal one-third of their
length, tapering abruptly proximally to a long narrow pedicel not readily distinguish-
able by eye from the underfur, than which it is not a great deal wider. The distal
portion is of oval section, not terete. The luidcrfur is from 15 to 18 mm long. The
ground colour in this group is, in general, redder than in genetta because the pale-grey-
ended underfur that forms its basis is overlain by bristles that have wide subterminal
bands of red-brown or even bright orange. This reddening always shows but varies
in extent, being sometimes in young skins and at moult considerably reduced. The
pattern is of very numerous, small-sized, dense black spots not always very clearly
disposed in longitudinal lines; but where they can be coimted they form at least seven
such on each side of the body. The spots are generally well under 20 mm diameter;
and they are mostly separate though occasionally they may coalesce near the spine.
There may or may not be a spinal crest.
The tail is the other very characteristic feature of this group. It is subcylindrical,
relatively short-haired and soft-furred, and very distinctly annulated from root to tip,
the edges of the rings being fairly sharply cut. The bristle-hairs do not greatly exceed
in length the abundant underfur. Whereas in genetta the tail broadly matches the body
colour, in seri'alina it, as a rule, torms something of a contrast by reason of the fairly
pure white of its light rings. The dark rings, which are generally black or very deep
brown, are usually 10 to 12 in number, the tip being white, though as a rule not so
purely white as the remainder ot the light rings. The alternate rings are subequal in
breadth or, more commonly, the white narrower than the black. The tail is shorter
than the head & body. The legs may be black or pale; the usual genet pale facial
markings below and between the eyes and around the rhinarium are present but
mostlv more obscure than in genetta.
Skull. There are not enough mature skulls to make comparisons with the genetta
group particularly useful. The characters are variable. In the six skulls available the
sagittal crest is restricted to the extreme posterior segment in four and, across the main
body of the braincasc, is incipient in two, a medium-aged male and a very old female.
The internal cusp on p^ is large in two examples; and in the others ranges from vestigial
through a mere bulge to a low tubercle. Both /))- and "12 vary a good deal in size,
and the latter may be quadrate with four cusps or triangular with three. The postorbital
processes may be long fine points or blunt and dctlccted. From such variable material
it is scarcely possible to derive any useful diagnostic characten. The most constant
feature is that the intertemporal breadth is narrower than the interorbital ; but in one
case it is appreciably wider. In the genetta group examples it is always wider.
Taxonomy. The group is usually regarded as a clear-cut one. Today it is universally
held to be monospecific although in the past three discrete species were named. In this
ipS THE CARNIVORES OF WEST AFRICA
present accoiuit it is suggested that there arc three vaHd species, two ot thcin occurring
within the area dealt with in this work.
The group first became known when in 1H55 Puchcran described sen'aliiia from
Gaboon. His brief Latin diagnosis was not in itself very precise; but since it makes it
clear that the spots arc extremely numerous, the limbs almost completely black, and
the tail long with broad black and narrower white rings the species is readily recog-
nisable and has never been questioned. At the same time Pucheran named a second
specimen from the same area aiihryana, the description being vague, the only apparent
difference from scrvalina lying in the ground-colour. As tliis last is alwa\s variable the
two names have for long rightly been regarded as synonymous.
In 1902 Thomas described hcttoni trom Kenya as a discrete species on the score ot
smaller size and annulate spots. This, which has proved to be a very common form in
eastern Africa, is now regarded as nothing more than a race oi scrvalina. The annulation
of the dorsal spots is not constant; but the markedly small size of the skull and the
shortness of the rostrum and palate probably justify Thomas's original evaluation as a
species. The same cannot be said ot Lonnberg's i\itcusa from the Congo, the author
himself being doubtful ot its validity as a race of seri'aliiia. Ot more immediate concern
to West Atnca is the third named race ot this species, cristata Hayman. Paler ground-
colour and pale forelegs apart, there are certain features of this animal which appear
to the present writer to entitle it to consideration as a discrete species. These arc the
possession not only of a black spinal crest, absent from true scrvalina, but of an erectile
nuchal crest as well. The distribution, so far as we at present know, lies between the
Cross River and the Sanaga; whereas scrvalina itselt is tound, in West Africa, only to
the south of the latter river, that is to say entirely extralimitally.
The evidence that scrvalina occurs as far west as the River Niger, as stated m Schwarz
(1930) and Rosevear (1953) is slender. No record has been traced to account for
Schwarz's assertion; the green spot shown on map No. iSia in Rosevear relates to a
rather poor skin collected by the present writer near Benin and of doubttul standing.
Though from its general appearance, including the typical short-turred, wholly-
annulated tail, it obviously belongs to this present group it is certainly neither scrvalina
nor cristata. The ground-colour is a richer orange-browai than in these forms; the spots
are appreciably larger and less numerous — so much so that it was originally thought
to be a "niaailata" . This was later altered to scrvalina by R. W. Hayman. The forelegs
are black, much as in scrvalina, but there is a very clear, black spinal crest on the lower
back, not occurring in that form, though no nuchal crest as in cristata The tail has only
nine black rings, the tip (the 9th) being black Whether this is a species or merely a
race is open to question. It is here treated as a new species in its own right.
The two, consequent. West African forms can be readily told apart by means ot the
key on page 189.
GENETTA CRISTATA Hayman Crested Genet
Gcuctta seri'ttlina criitaia H.iynun, 1940, Trans, zool. Soc. Land. 24: 6Sf)-6S,s. Okoiyoiig, Mamfc Division,
Cimcroun. Type 111 the British Museum, No. 39.323, ^; skin in good condition, skull good except for
GENETTA CRISTATA 199
the loss of four upper teeth. The subspecific name was given with reference to both the nuchal and
spinal crests.
Distribution and general. This genet (Plate 3) is known in the British Museum
from 5 adult and 9 yoiuig or juvenile specimens, all from the Cross River — Cameroun
forests. Only 3 of the adult specimens arc complete with both skin and skull. The
majority of these come from various places in the Mamfe Division, collected by Sander-
son (1940): Mamfe, Binjong, Bashauo, Olulu and Okoiyong; but there are others
from the Rumpi Hills (Kuniba Division, Cameroun) and Oban (south-eastern Nigeria).
In addition to these, Eisentraut (1963) records 2 specimens from Mount Cameroiui at
Buea and Malendc. The whole area of known distribution, therefore, lies between the
Cross River basin and the Cameroun Mountain, a heavily forested and, imtil recently,
secluded region, about 100 miles long by the same wide, well north of the Sanaga
River, south of which the species is replaced by servalina.
Description. In the servalina group the ground-colour of the dorsal pelage is some-
what variable from specimen to specimen ; but in so far as a generalisation can be made
it is rather paler in aistata than in servalina itself. It is buffish-grey ; the orange-
brown subterminal rings of the individual hairs adulterate this colour to a greater or
lesser extent in different examples, sometimes scarcely at all except romid the anterior
edges of the black spots. The latter vary a little in size and shape but most typically
may be regarded as roimd, and 10 or 15 mm in diameter, though sometimes they tend
to become rather longer than broad. They are, in general, independent but there is in
some specimens a greater tendency to unite than in others; and almost always they
torm short lines near the spinal crest in the region of the hindquarters. The bristle-hairs
of the lower back are about 20 to 22 mm long; the underfur some 10 to 13 mm.
In servalina the belly is dark, almost blackish-brown; the chin and throat are lighter
but nevertheless darkish grey and, especially in the latter, washed with golden-brown ;
cristata, on the other hand, is considerably paler below; medium buffy-brown on the
belly but the chin and throat cold whitish-grey with little or no warm over-wash.
The black spinal crest becomes noticeably long from the mid-back to the root of the
tail; but it is, in a shortened form, really continuous with the nuchal crest. This latter
is much shorter than the main spinal crest being, indeed, little longer than the surroimd-
ing tur from which it stands out by reason of its ercctness instead of being adpressed
and directed backwards. Though it is most conveniently referred to as the nuchal
crest it, in fact, has its origin, in a very low form, on the front of the face from about
between the eyes, whence it continues over the crown to the neck where, the fur being
longer, it becomes far more obvioas. It is caused by the direction of the hairs, which
converge towards the medial line. On the neck it involves the central black stripe and
the brown-ringed contiguous hairs, and is thus of mixed colour, not wholly black like
the spinal crest. It is of interest to note that rudimentary traces of the frontal crest
arc to be found in typical servalina.
The tail may have fewer black rings than servalina, the type having only 8 clear
rings, thougli others, including all the juveniles, count up to 10; but the reckoning
of the terminal rings is open to argument, there being a very obscure narrow darkening
200 THE CARNMVORr.S OV WEST AIKICA
that could or could not be regarded as a ring. The tail-tip is really white, but because
of this indecisive blackening becomes greyed. The forefeet arc in their upper portion
light-coloured and spotted similarly to tjic back, but become blackish towards the toes
and on their outer and lower aspects. The hindlegs are somewhat similar but often
carrv a distinct pale, greyisli, patch over the metatarsal region above.
Skull. There is no particular distinction between aistata and sevvaliiia. From the
very few examples available for comparison it is just possible the inner cusp of /(^ is
better developed in the former than in the latter, as in the three available specimens
it is large in t\\ o and represented by a prominent swelling in the tliird. All skulls have
a posterior sagittal flange; but a crest across the lengtli of the braiiicase is rudimentary
in two, and absent from two, an old male and a fairly old female. This old male is
the only specimen in which the postorbital breadtii is greater than the interorbital.
The transverse breadths of /ii- and ;»2 are rather greater than m scivaUna. The bulla is
illustrated in fig. 28c.
Habits. Sanderson (1940) characterised the habitat as scrub, low tangled vegetation
and bare ground below trees. He considered the species as detmitely terrestrial, only
one specimen having been seen in a low tree, and that returned to the groiuid to make
its escape.
GENETTA BINI spec, not: Benin Genet
The erection of this species from one very incomplete specimen seems to be justified
bv its apparent difference from other known forms, from its extreme western pro-
venance, and the wide gap of 750 kilometres separating it from its nearest recorded
neighbour, across the two fauiial barriers of the lower Niger and Cross Rivers. It is,
in fact, described below from a, in some respects, poorish skin devoid of anv supporting
skull.
Description. The type, B.M. No. 50.315, o (Plate 4), was collected for the present
writer in December 193S by R. II. Hide, a forest officer, in the Ohosu Forest Reserve
some 65 kilometres north-west of Benin City, western Nigeria, altitude 250 metres.
Although appreciably different in detail, the overall resemblance of the pelage and
tail patterns to scrviiliiia make it quite certain that it belongs to that group. The ground-
colour of the dorsal pelage is of a much richer red-brown than in the two other more
commonly known western species, or in the more golden-brown of iiitciisa from eastern
Congo. This is because the subterminal bands of the bristles are both broader and more
deeply coloured than in the other species. The pelage length is much the same as m
cristata.
This new proposed form (I'late 4) diHers from sciraliini 111 the piossession of a long
black spinal crest from the mid-back to the tail; and from crinata by the absence of a
nuchal crest. The fur on the back of the neck, however, is directed forward and meets
the backwardly directed hair of the frontal region in a line across the crown between
the ears. Where the differently orientated hairs of the neck and back meet there would
appear to be something in the nature of a whorl; but tlie only skin is so poorly made
that the position here is a little obscure.
GENETTA PARDINA
The greatest difference between this new and the other species, however, Hes in
the size, shape and number of the spots. They arc much more irregular, less round,
in shape, and appreciably larger and fewer in number. Many of them are subquadrate
or triangular, 20 mm or more across, those of the middle lines being larger than those
near the spine or lower down. Between the shoulder and the hip there are not more
than 7 or 8 in a row as compared with 10 to 12 in the hitherto known forms. A few
merge over the rump and one or two are confluent with the spinal crest. Owing to
the make-up of the skin and the virtual absence of the belly it is not possible to say
precisely how many rows there arc, but it is probably 6.
There are 9 dark rings on the tail including the black tip. The pale rings, which
are only about half the width of the black ones, although white below, arc dorsally
distinctly more coloured than in the other species of this group, especially the proximal
ones, being greyish with a fair mixture of reddish and black hairs. The bristles of the
proximal black rings are rather longer than normal and overlap the light rings more
than is usual. The chin and throat are medium grey, fairly well spotted. Although
most of the legs are present there is only one foot, the left forefoot. The upper arms
are deep grey and spotted on top but black beneath like the whole of the remainder of
leg and foot. The lower parts of the hindlcgs are blackish all roiuid.
Nothing is recorded of this animal except that it lives in closed high forest, and the
only specimen known smelt strongly of musk. The name Bini, pronounced bee-nee,
is the name of the ethnic group inhabiting Benin and the surroiuiding district.
C. pardina Group
In all the confusion that exists in the genus Genctta the greatest, without question,
concerns the three classic species, tigrina (1778), jAina (1811) and paidina (1832). The
last is of closest concern to West Africa; and though it has in some measure, through
the attribution of the same forms to one or the other, become involved with the first
an endeavour must be made initially to sort out the question o( pardina alone.
With the pardina group we reach a remarkably mixed assortment of forms, far and
away the most variable in visible characters. Wliether the group embraces more than
a single species is an open question. The animals lying at opposite ends of it are assuredly
markedly different in appearance; yet there seems to be no clear-cut division between
them. Great play has been made by taxonomists with spot size and colour; with the
number of tail annulations and the relative widths of the light and dark rings; and with
the colour of the feet. There is little doubt that these characters, and others less often
taken into account, are subject to considerable variation, possibly to some slight
degree local but for the most part individual; though from the largely disjoint study
material in the British Museum — scarcely any two from the same locality, apart
from questions of age and sex — it is hardly possible to do more than conjecture that
this must be so. However, the 46 specimens collected by the American Museum
Congo Expedition, by reason of their restricted provenance, cast a more certain light
on the matter. J. A. Allen's detailed comments on variation of supposedly diagnostic
characters as cxliibitcd by this scries have already been quoted on page 187.
THE CARNIVORES OF WEST AFRICA
Description. In ilic Uci: ot tlic \^■idc inconstancy thus revealed it sccnis difficult,
it not impossible, to find dependable characters that might serve to defuic pauUiia.
In truth the matter boils downi to a similarity of tail structure and markings throughout
an otherwise capricious group, serving to differentiate the animals included here on
the one har.d clearly (i-oni (jaicttci and on the otlier, less emphatically, trom scnudina.
This tail is of the relatively short, backwardly-directed-haired, subcylindrical type
charactcric o£ scri'aliiia, differing manitestly trom the ver)' long-, coarse-haired type
ot du't^cnctla group in its extremely soft and silky nature. It differs from that oiscrvaliiia
in being appreciably more melanistic, the pale rings being not only few er in number
but, as a rule, of far less width than the black ones. Further, the terminal portion of the
tail, sometimes only about 75 mm, sometimes very much more, is almost whollv
black, particularly above, there being, not uncommonly, faint traces of white partial-
rings on its underside. It is, in this species, more convenient to count the pale rings
than the dark; but often there arc only halt-rings, or less, visible on the underside but
not the top. The count varies, therefore, on the two aspects, dorsal and ventral, of the
tail. Taking the latter, there are usually 6 to S pale rings or partial rings, but in excep-
tional specimens there may be one less or more. Even in the basal portion the rnigs are
often fir more obscure above than below. Their lesser number, the (as a rule) narro\\-
ness and relative obscurity ot the pale rings, and the long black end to the tail serve
as a distinction between even the small-spotted forms of this species and the scrvahiia
group, in which the rings have a sharp clarit)', are subcqual in width, and continue,
above and below, to the distal end.
Taxonomy. The mix-up owcv paniiua stems partly trom Matschie (1902) and partly
from Schwarz (1930), both, in an obscure situation, having long been regarded as the
ultimate authorities; and what they have said, surmised or hinted at has, often blindly,
been followed. Added to this is, at least as far as the British Museum is concerned,
a plethora of specimens trom the rain-forest belt and a paucity trom the drier inland
areas. Now, Isidore Geoffroy, clearly as the result of specific enquiries, described
pardiua as emanating trom the interior of Senegal, that is to say from one of the more
arid types of woodland. Sudan or Saliel, certainly not from the closed forest. There is a
temptation today, and both Matschie and Schwarz succumbed to it, to criticise early
notions ot West African geography and to amend reputed provenances to suit what
seemed to them to be the tacts. It cannot be denied that this is sometimes justitied;
but there is no reason whatsoever to suppose that Geotiroy was in this particular case
misled. Senegal had tor long been intimately connected witli France and in Geoffrey's
day was certainly closer, and a far more likely source of livhig animals, than any more
heavily forested part of West Africa.
Matschie showed himself to be in two nriiids regarding the vegetation and distri-
bution proper to the species. On page 1135 of liis paper he connected pardiua with the
coastal belt of Togo; but on page 1142 he gave as the range first north Cameroun,
secondly Togo; and implied possible synonymy with pociisis, a forest form. Schwarz,
too, who first identit'ied piudiua with tnacidata (a synonymy since necessarik reversed)
was not very clear regarding the ecological background proper to the species. In a
footnote on page 277 he said, in translation: "Gciictia uiaaiLua is p.itentK' the same av
GENETTA PARDINA 203
Genetta pardina I. Geoffroy, so that the second, more used, name must disappear as
clearly a synonym. Matschie (1902) has already shown that maculata is no north African
form as the original description says . . . (It) doubtless emanates from upper Guinea
and, since it was imported live, can very well, like the type of G. pardina, stem from
Senegal, which was at that time one of the most important export coimtries for African
animals". In other words lie agreed to its relatively dry-zone origin. Yet in his main
text, near the top of the same page, Schwarz quite unequivocally rated maculata (as he
called the species) an upper Guinea closed-forest ("Waldgcbiet") animal, ranging from
the Niger to the Gambia — thus vegetationally, and possibly distributionally, running
cotuitcr to Gcoftroy's "interior of Senegal". J. A. Allen had obviously adopted an
entirely different view of the distribution oi pardina in that he had ascribed the major
part of his north-eastern Congo material to the species. Further, in another footnote,
on the previous page, while allowing the species a certain degree of range in climate,
Schwarz none the less still confmed it to the high-forest: "While among the forms of
the moist forest G. maculata produces a small-spotted type 'pocnsis', the spots in the
marginal areas oi the forest arc large and the groimd-colour pale — 'maculata' {'pardina'),
'genetloidcs' ; intermediate types 'diibia', 'johnstoni' " . Attractive and reasonable as this
hypothesis may seem, it will be seen shortly that Schwarz was quite wrong in con-
necting spot size with humidity.
Relatively recently an extremely interesting set of 10 flat skins, in only very fair
condition and luifortimately without skulls, has been received bv the British Museum
from Mr. J. D. Carter. They all emanate from a very restricted area some 30 kilo-
metres long, in the Pctico-Zo locality ot the Wukrmn Hills, 25 kilometres north-east
to 25 kilometres north-west of Lau, which lies on the upper Bcnue River in Nigeria
at about ii°30 East. This area is in the Sudan woodland zone. The interest of this
scries lies in the fact that it shows that no less than three quite distinct, though very
similarly coloured, taxonomic forms, scncgalcnsis, pardina and tliierryi, live sympatri-
cally; that eight of the skins which arc pardina display a wide rajige of size, shape and
colour of spots, some of them being close enough to Geoffroy's description and illus-
tration to make it clear that the type specimen o£ pardina may well have come from
the Sudan zone and that Schwarz's limitation of the species distribution to the closed
forest v»'as misleading. It must, 111 all fairness, be added that one exceptional skin.
No. 39.132, from a small island in the mouth of the Volta River, near Ada, is not dis-
similar from these Sudan zone specimens, though of a slightly colder tone, and quite
distinct in its paler ground-colour from all southern Ghana forest material. But although
so near the coast and thus expectcdly in the wet-forest zone, this area is, in fact, one of
remarkably low rainfall, not greatly exceeding that of the much more inland Sudan
zone, and the vegetation of a dry littoral kind, together with some mangroves.
Because of the wide range of pattern and colouring in the pardina group the present
writer is rather sceptical of the usefulness, or the feasibility without considerable further
evidence, of drawing nomcnclatural distinctions between the various forms. The
extremes are very distinct and could support independent names; and it thus seems
unjustifiable to apply precisely the same term to the pale, creamy-butf, large-spotted
animal ot the dry open country and to the relatively dark, greyish, small-spotted
204 THE CARNIVORES OF WEST AFRICA
inhabitant ot tlic dense rain-torcst. But, though there arc considerable laciuiac ni the
study material, there is some reason to suppose that the forms arc wholly clinal, and
in the attempt to differentiate everything one could be faced with devising an almost
endless string of names, the limit of whose application was anything but clear-cut.
In the face of this the question of what to do \\ ith puniiiui is a difficult one. The
course adopted in tliis work is to regard it as a species-group, calling the pale open
comitr)- forms pardiua. the large-spotted forest form <^aicttoiiIc.<, and the small-spotted
iorcst form fiicnsis. This is more a matter of convenience than anything else and can
at once be shown to be illogical since, while differentiating between the extreme
forest forms, it tails to do the same for the dry-zone forms; and, as regards the forest
forms themselves, whether a specimen is large-spotted or small-spotted must often
remain purely a matter of personal opinion. But it has this merit: that it does not
burden an obscure situation with a mass of new names that more abundant material
and deeper continental-wide research would almost certainly show to be unnecessary
and inept.
This decision having been made it is necessarv to examine and discuss the standing ot
certain named forms ui more detail in order to explain the difficulties which have arisen
over some and the reasons tor the rejection of others from the West African list.
The earliest of these is viaailata Gray, 1S30, which since Schwarz (1930) has been gener-
ally accepted as synonymous with paniiua. Owing to doubts and contradictions in
Gray's diagnosis the position is, in tact, not absolutely straightforward. I lis English
description is of a grey-brown animal with a brown erectile crest from the shoulders
to the tail, and spots in 3 rows on each side, the 1 innermost square, those next to the
spine largest and nearest together, the lowest series roundish; below these are scattered
black-browii spots. There would thus appear to be 3 clearly defined rows ot spots
on each side with further spots, not disposed in lines, below them, a not unusual
arrangement in the genets, giving rise to the common diagnosis "spots in 4 or s rows
on each side". But Gray's Latin diagnosis gives the spots as in 6 rows with scattered
round spots on the flajiks. hi view ot the English description this could logically be
taken as meaning a total derived from two sides of 3 each. The accompanying illus-
tration, however, clearly shows 6 well-defined rows of independent spots on a single
side; and since Gray must presumably have seen the drawing before publication it
might be taken that this is what he, in fact, meant — though it scarcely accords with
his verbal description ot 3 series of spots with scattered spots on the sides ot the bell)'.
Neither does the illustration bear out the written description ot the tail. This states
that there arc 7 black bands increasing in breadth towards the end. The picture, how-
ever, shows 8 together with a black tip making, in all, 9 black bands, which show no
appreciable increase of width.
Schwarz, synonymising maculata znA pardiua, drew attention to the fact that Matschie
had already expressed the view that the former was no North African animal, as Gray
had stated, and had noticed, also, the discrepancies between Gray's diagnosis and his
illustration. Schwarz, ignoring the confusion over the spots and contming himself to the
tail, expressed the view tjiat the written description "mav be regarded as authoritative
as it speaks ot 7 dark tail-rings, which .igrecs with the upper (iumea genet. Apart
GENETTA PARDINA 205
from the inexactness of the drawing of the tail the picture shows a long-legged, long-
faced genet that doubtless emanates from upper Guinea . . .". It will be seen that
Schwarz accepted or rejected the illustration to suit his own theory and his own
defmitions. He further supported his view by a pure guess that the country of origin of
Gray's specimen w^as really Senegal.
With doubt cast on the body pattern and on the tail there seems little enough in
the type description that may be picked upon as usefully diagnostic apart from large,
quadrangular, more or less separate spots. One thing is certainly lacking from both
verbal and pictorial descriptions : any reference to the markedly melanistic tail with the
extra-long black tip and pale half-rings which arc the essential characteristic of all
parditia. The illustration is, too, of a distinctly more tapering and fmely-tipped tail
than is general in that species. The true country of origin of Gray's specimen is, in fact,
not without relevancy; tor his figure is, apart from an extra line of spots on the flank,
not a bad representation o(^ genetta afra, which, if the animal came from North Africa
as he said, it could well be.
The name maailata has been rejected as unavailable because of prior occupation.
Were this not so it could justifiably be abandoned on the grounds of obscurity of
definition. This latter also renders its correct position in synonymy uncertain, but in
order to avoid introducing unnecessary confusion it has herein been retained in its
customary place under paidlua in the synonymic list which follows later on page 209.
It is perhaps of general interest, as a commentary on Gray's contused diagnosis,
to quote here a view of his reliability written not from the purely zoological angle
but as a pointer to his character: "He published a steady stream of brief zoological
papers with such haste that in a number of instances he changed his mind and published
a second paper correcting his first; indeed, in some cases he soon published a third
paper amending his second". (Paden, 1964: 13).
The forms pociisis and gcncttoidcs will be commented on in their own appropriate
sections later. The status of a few other names must be examined here. The first calling
for attention is ficUiaiia Du Chaillu, i860, because from time to time West African
specimens have been referred, directly or indirectly, to it or it has been stated to occur
in that region — as, for example, in Rosevear (1953). Du Chaillu's description is not
very helpful but makes it probable that his animal belonged, in fact, to the pardiim
group. This is fairly clear from the tail, which was said to have 7 dark rings, the first
incomplete below, the last indistinct; the last 5 inches of the tail brownish-black.
Other points in the description tend to confirm this. The size of the spots is not given :
but, indirectly, by mentioning that the two lateral rows on each side of the spinal
band arc broken up into five or six smaller longitudinal spots he indicates that these
must in fact be largish, as in genettoides.
J. A. Allen (1924) treated ficldiana as a race ot pardiiia; but Sch\\ arz {1930) regarded
it as a subspecies of tigrina, bemg followed 111 this by G. M. Allen (1939). In view of the
character of the tail this latter classification seems improbable. Not much is revealed
by skulls in Gciietia; but there is a Du Chaillu skull of an old animal in London (No.
1645a, sex?), without a skin, that conforms pretty closely to the general riui of the
pardiiia group except that it is on the whole rather below average size and has an
:o6 Till" rAUMVdliKS (II- WIST AIRKA
:i'
uiubually narrow iiucrtcmporal auisuiction. However, it is matched m this lattc
by one of the laigc-spottcd oencnoiilcs from Ghana (No. 46.397). Tlic standing of
Ucld'uiiui is only incidental to West Africa but is regarded herein as jiertaimng to the
Ihudiiia species-group, the name being iirobablv synonymous widi ocucttoidcs.
Matschie's duhia, the exact provenance ot whicli in West Atrica is unknown, is also
almost certainly fn'itdtoides. No set description ot diihiii was given by Matschic and
diagnosis ot the form must be built up from his scattered key characters. The most
significant ot these are: spots large, in 5 rows; tail short-haired, the hairs not more
than 2s mm long at the root; the tail itselt very short, two-thirds of the head & bodv
length (tliere is a misprint in the key), with only 6 pale rings, which arc at most as
wide as the dark. Tails are. 111 the pardiiux group, normally rather shorter than the
head & body length (about S'i per cent); but it is doubtful whether such an extremely
short-tailed animal exists except as a treak or the result ot accident. The torin dulna.
theretore, isjiere regarded as, in iact.frciicttoidcs. Schwarz, also, put the two in synonymy.
Cabrera s liistilctrls. described trom Fernando Poo in 1921 because no trace ot pociisis
Itselt had so tar come to light on the island, must next be considered. Schw'arz grouped
this with rifJiiiia; but Cabrera himselt likened it in general to pardina, oi which he
considered it to be an island representative. He described it as having few spots, large
and annulatcd for the first two rows, which are more irregular and imperfect than 111
pdidiihi. many of them formed trom two black spots enclosing an intermediate, ill-
detmcd reddisji space; the base of the pelage infused with cinnamon, the flanks and
tjiighs very bright; the feet grey. No mention is made ot the tail. There is no reason
to suppose that Cabrera's estimate ot taxononuc relationship was wrong and that
Schwarz was more justified in allocating the form to ti(iriiia. The form is, therefore,
in this work s\nonynnscd whh f^ciicttoidcs; but it is possible that with tunher collecting
It may be found to merit some more independent status within the pardina group.
No form is attributed in this present work tl 11 in (ieriii.iii was iicH alwa\s \-er\' acctir.ite, .is when
GENETTA PARDINA 207
he said there was a black stripe troin head to tail whereas Vosmaer quite clearly des-
cribed it as running from the middle of the back. It may be as well to add here that
the plate accompanying Schreber's type diagnosis is, as far as outline is concerned,
an exact copy of Vosmacr's or perhaps made from the same block; but the colouring,
in the British Museum version at least, is of a very markedly darker brown than in
any ot tour different copies of Vosmaer.
Very few characters of any taxonomic value arc, in tact, given by either author.
Schrcber stated that there were many irregular browii spots; this he derived purely
from the illustration, Vosmaer having made no reference to them. This picture, which
carries the subscription that it was drawn trom life, depicts them as extremely irregular
in shape, that is to say neither roundish nor squarish but with far more jagged outlines
than are tound in any existing genet skin. This is explicable m that, if the drawing
was truly made from a living annnal, the tur may well have been in a rough, wind-
blown state, not brushed smooth as in a prepared specimen. This would argue a long,
loose pelage; but it is difticult to make the markings ot any existing specimen ot genet
take on the size, appearance and positioning ot the blotches depicted by Vosmaer's artist.
The tail was described as ringed with black and white and having a black or brownish-
black tip; but the number of rings was not stated, nor does the illustration make this
clear, the only two, very narrow, pale rings visible, near the distal end, run at a slope
such as is foimd in no genet. The dark tip is relatively short, narrow and pointed.
The overall impression given is that ot a long-haired structure, matching the similar
inference respecting the dorsal pelage made above. Other, more minor, points cannot
be touched on here.
Clearly there is very little that is reliable in the way ot characters to be squeezed
out of the original diagnosis. When we come to consider what has been made ot
lif^rina since by Matschie (1902), Thomas (1906, 1908), Schwarz (1930) and Roberts
(195 1) we get a pretty mixed picture, sometimes in direct conflict with the type des-
cription and/or its illustration. Each has attributed to the species characters of his own
devising. Matschie, for example, defmitel)' stated that the hairs at the tail root were
at least 30 cm {sic) long; that, despite the obvious taper in the illustration, it was about
as broad at the base as at the tip; that the last quarter of its length was black with only
traces ot narrow half-rings below; that the tirst two dark tail rings, neither visible
in the illustration nor described, were at least as wide as the succeeding pale rings.
This may be tigrina Matschie but it certain!}' is not tigrina as described by Schreber.
Again, Thomas & Schwann (1906) stated, directly and indirectly in their key, that the
whole midersurtace and forelimbs \\ ere dark-brown or black — which trom Schreber
and Vosmaer was patently untrue; and, in tact, Thomas & Wroughton (1908)
modified this to forefeet black — the original description ("feet with a lot ot brown")
and the illustration belying this misleading assertion. Schwarz endo\\ed tigrina with a
number of new characteristics: short legs, short face; pear-shaped bullae, strong post-
orbital processes and a distinct sagittal crest — all very inconstant characters in Gencltti
skulls. He stated the pelage to be variable but never shaggy, with large, mostly brown-
centred spots; and the tail to have 8 to 9 dark rings. Other uncertainties and contra-
dictions in diagnosis concern the niiier cusp on /)■'.
20S TIIF. CARNIVOKFS OT WEST AIRUA
The characters which most authors concur in tor n^i;n'»ci and whicii do not conflict
with Schreber's and Vosmacr's descriptions, and which also accord with specimens
trom the reputed region of provenance. South Africa, are: that the pelage is longish,
with large, mostly independent spots in about 4 longitudinal rows, and a dark, probably
erectile stripe on the lower halt of the back; the tail with about cS dark rings and a dark
tip; the legs and feet with a good deal ot dark colour on them, especially the back ot
the hindlcg contiguous to the heel, the hindfoot having at least some grey on its upper
side. However, in view of the doubt and conflict of interpretation that ti^rina has
engendered a good case could be made for the abandonment of the name. But whatever
attitude may be adopted towards this, it seems clear that, in so far as it is possible to
gather, the animal intended by Vosmaer and Schreber is probably more akin to the
long-haired ficnctta group than to the relatively short- and soft-furred pcvdliui; and
that as regards specimens from eastern Nigeria and Cameroun, as well as Allen's
eastern Congo series, Schwarzs hypothesis about distribution is entirely at fault,
these animals being certainly pardiiia.
The question ot iiihi'^iiicsii Pucheran, iHa, is otten involved with that ot rif^riiid,
as to whether they are either one and the same, the former a race of the latter, or
specifically discrete. Though ot no tirst-hand concern to West Africa, the following,
which emerged incidentally from the wide investigations necessarily made, may be
added here as an appendix to the above. Pucheran's curt Latin diagnosis would appear
to be too vague to be of any real use whatsoever; whitish-grey washed with tawny;
the spots of the back almost totally rusty; the tail furnished at the base with four rusty
rings then four black. That is all. It is too obscure, 111 the light of now known colour
variability, to render riihi{iiiio/(i the spots were large and the forefeet black, whereas in riihigiiwsa they
were of medium size and the forefeet pale.
Skull. The general form of this in the pardiua group is ver)- much as in other genets.
Possible points of distinction, such as the existence ot an inner cusp on p-^, or ot a sagittal
crest, are as variable as in other groups. One dittereiuial character alone is practically
constant; in fully adult skulls the postorbital constriction is, with few cxcep>tions, less,
and sometimes markedly less, than the interorbital breadth, this difference being on
the average greater than it is m the scwaliua group. No significant variation of external
appearance can be detected in exceptional examples. On the average the excess of
interorbital over postorbital breadth is ot the order of 2 mm. At its greatest it may
reach 5 or even 6 mm, the latter width dropping to as little as 8-0 mm, the cranial
constriction becoming a very marked feature of the skull. One skull in which this
occurs. No. 46.397, is a quite typical large-spotted f^cncttoidcs from Oda (Ghana);
the other. No. 1645a from Gaboon, has no skin.
Of the two adult skulls 111 which the intertemporal breadth is greater than the
interorbital, one has an excess of oiiK' or mm. In the other. No. 27.8. 12. i (Ghana),
the excess is 41 mm; ,ind there .ire iiileresting ditterences 111 the teeth. />'. )»' and m-
GENETTA PARDINA 209
being smaller than normal, especially the last which is only half the usual size and lacking
from one side. A similar state of affairs with regard to iifi exists also in a Mamfe skull ;
but neither skin is in any wav out of the usual run o^ ^cnettoides save that the Ghana
animal was in full moult.
Two other skulls have unusual features but both arc without skins by which to check
their standing. No. 61.42, from Musaia (Sierra Leone) is narrower across the upper
camassials than any other specimen and also has an appreciably smaller p^ and m'.
And No. 68.493 has remarkably long, backwardly directed postorbital processes,
measuring 27-1 mm across in contrast to a mean of 19-5 mm. These points are men-
tioned since they have at one time or another in single skulls been regarded as having
some taxonomic significance. There is no clear reason to suppose that they have;
the material available for study is quite inadequate to provide sound data; and they
are connected with no readily detectable external differences. Other characters which
liave been used taxonomically yield, as regards British Museum specimens, the follow-
ing data. An iinier cusp on p^ is of irregular occurrence and is independent of age or
sex. hi adult skulls of the pardina group it is present and large in 9 specimens, small in 3,
and lacking from 4. As for a sagittal crest: it is present in 8 old skulls, 3 of them ^,
5 of unknown sex; and it is absent from 4 medium-age skulls {zSS, 2??). There are
no adult female skulls from which to derive data. Finally, the nature of the postorbital
processes has been suggested as of possible taxonomic significance. However, they
show no fLxed form, in length, pointedness or direction, varying from short, blunt
and deflected to sharp and long-pointed irrespective of other characters.
The three West African species included in this species-group may be separated by
the key on page 189.
GENETTA PARDINA I. Geotiroy Parduie Genet
Viverra maciilala Gra\', 1S30, Spicilegia Zoologka .... 2: 9, pi. 9. Type localit)- given as Nortli Africa
but in tlie opinions of Matscfiie (1902) and Schwarz (1930) this should really be West Africa. This
was described from a living animal in the Tower of London and there docs not appear to be any
preserved type. The specific name is the Latin word for spotted. Preoccupied b\' I'ii'crra inaailaia
Kerr, 1792 (= Dasyiirops iiiaailattis Kerr). See p. 205 herein.
Gcnctia pardina I. Geoffroy, 1832, Mag. de Zoo}. 2iid year, class I, pi. S and accompanying text. Interior
of Senegal. The name is a diminutive of the Latin pardiis lepoard.
Cciietta pamherina Hamilton Smith, 1842, in Jardhie's Naliiralisl's Library: 35 (13 of Mammalia): 171
No type locality given. This, from the Greek panther leopard, would appear to be nothing more than
a renaming, or misnaming, of pardina \. Geoffroy resulting from the French name also given b\'
Geoffroy at the head of his description, G.(cncttc) panthcrine.
Distribution. As imdcrstood in this present work, and as defined above, pardina
occurs in the drier open woodlands from Cameroun to Senegal. How far east it ranges
in these belts has not been determined; but the present writer has little doubt that
East African forms from as far afield as Tanzania, and usually nomenclaturally associated
with tigrina, belong, in fact, to the pardina group, if not pardina itself. West African
specimens ot pardina exist in the British Museum from the Wukrum Hills (Northern
Nigeria. Sudan woodland) and Fort Lamy (Chad, Sahel woodland).
::to iin: (aumvores or wrsr ai kh a
Description. The scries ot K flat. p.irtK' incomplete skins trom the Wukriuii Hilis
has alrcidy been referred to. No two skins are precisely alike in their colouring and
marking. Six have the ground-colour a sort ot creamy- or sandy-buff; the seventh is
rather grcvcr; and the eighth somewhat more whitish — though, nevertheless, still
palely washed with sandy tips. There is an erectile spinal crest trom the shoulders to
the tail, black in most cases but rusty in two. On each side ot this are four well defmed
lines of spots, ranging, in different skins, between almost entirely black to almost
entirely red-brown, some being obscurely black-ringed with rust\ centres. The sizes
and shapes of these markings arc highly variable. In some skuis thev are wholly indepen-
dent, in some coalesced longitudinally into short lines; some are roughly quadrate,
some more oval, some rounded; and in one skin they are almost Imear, the width ot
the blotches being only some 7 or S mm, as contrasted with twice this or more in other
skins. The dense soft undcrfur is about 17 mm long, the tine-shafted bristle-hairs
about 24 mm.
The tails arc relatively short-, sott-haircd and exhibit dorsally 5 to 7 pale rings and a
black end portion 75 to 150 mm long, this terminal part interrupted laterally and/or
below by 2, occasionally 3, partial white rings. The legs and feet have been mutilated;
but are pale rather than black, diough the inside of the hindlcg appears to be dark,
at least in some cases.
The Fort Lamy skin differs in being of a more yellowish-brown colour and a less
well-detincd pattern of whoUv brown spots. The fulvous colour, which extends also to
the rings of the tail and to the undcrsurtacc of the skin itself, seems as if it might be due
to drying over a smoky fire.
Skull. There are no skulls.
Habits. No special notes regarding habits accompany the skins. There is no reason
to suppose that open-countrj' genets differ materially 111 these from their closed-forest
kindred except in so far as they probably more trequendy make use of rock crevices
tor shelter and breeding than high-forest forms do; and ground birds, francolins and
guinea-fowl, doubtless play a more prominent role in their dietary.
GENETTA GENETTOIDES Tcmminck Pel's Genet
I 'nrrr.i •jcnctioiilcs Tcmminck, 1S53, Esipiiacs zoologiipus sin In cote tie Ciiific: Sy. River lioutry and
Hiiuina, Gh.ina. According tu ]cntink (1887 and 1S92) there are four r\'pes in the llijksmuseum van
Natmirlijkc Historic, Leydcn: 2 mounted ++ with skulls separate, tVom the River Boutry; and 2
mounted animals ot unknown sexes from Elmina. Tcmminck coined this name from Gcnclta .ind the
(ircek termination -o-cidcs implying resemblance, trom tides lorm.
(jcncttci ficldiaiia Du Chaillu, i860, Pioc. BpsIoii Sec. nat. Hist., 7: 302. hiterior ot Ciabooii. This was
named in honour of his "distinguished follow citizen", Cyrus W. Field.
( H-nctta duhia Matschie, 1902, V'crh. des V intanationaleii Zoologcn-Concircssts zii Berlin, igoi: 1141. West
Alric.i, locality unknown. The Latin word diibia h.is several meanings, of which doubtful was that
most likely in Matschic's mind in reference more to the species' uncerM'ii standing (p. 1137) than to
Its indefmite provenance.
Cmclta insiilaris Cabrera, iy2i, Bolii R. Soc. ap. Hist, iinl., 21: 261. Rcbola, Island ot Fern.indo Poo.
rhc specific name is from the Latin iiisiiln island.
Plate 4
Pel's Genet (Gvm-tla_icncmmi,<): Small-spotted (ienct (G.wiu, i>oo,„>): Uen.n Genet (Cauv,, him)
GENETTA GENETTOIDES
Distriburion. This form (Plate 4) iiiliabits the closed-forest and neighbouring Guinea
woodland, plentifully from Gaboon to Sierra Leone and thence, less commonly, to
Gambia. The localities from which specimens exist in the British Museum are too
numerous to cite; but, closed forest apart, it may be worth mentioning that the most
inland places which have yielded specimens are Katsina Ala (Northern Nigeria),
Musaia (Sierra Leone) and Fouta Jallon (Guinea). These Guinea zone specimens may
possibly come from high-forest remnants or extensions ("fringing forest") rather
than from the more open woodland itself; but on this point there appears to be no
positive information.
Description. As typically described this form comes from the Ghana coastal forests.
One such skin in the British Museum, No. 57.9.16.3, is, in fact, from its history, almost
certainly one of Pel's original specimens and therefore a paratype of Temminck's
genettoides. Temminck wrote at some length of this genet, regarding it as very closely
related to genetta itself He gave no clear account of the dorsal markings, laying his
greatest emphasis on the nature of the pelage. From tliis and from his description of
the tail — not at all bushy, the hairs being half as long as in gaictta, the annulation
irregidar, the black rings wider than the white, the terminal third black with brown
half-rings below — it is abundantly clear that he was wrong in his estimation of relation-
ship and that the form without doubt belongs to the pardina group.
It has already been explained that pattern and colour are very variable and that
there seems to be no clear-cut division between this form and pociisis. In the most
typical gencttoides the spots are fairly large, separate, oval or quadrate, either wholly
black or with varying amoimts of ginger-brown hairs at the centre, making them to a
greater or less degree annulate. In size, those nearest the black spinal stripe are 30 to
40 mm long and 15 to 25 mm wide. They arc in 4 rows, with, often, a small part
of a fifth; and there are about 5 of these large blotches in the innermost row between
the shoulder and the hip. The legs may be either pale or blackish; but neither this nor
the colour and size of die spots is regarded as having any real taxonomic significance.
The hindfect are dark above with a more or less pronounced pale streak running along
the inner edge from about the base of the 2nd digit.
Between this and pocnsis at the other end of the scale, with small-sized spots too
numerous to count, there are all manner of intermediate stages of size, colour, shape
and independence. Some are deep black and remarkably clear-cut; other brownish
and confused. These variations show no connexion with localities. Two corresponding
to the deep-black clear-cut category come, the one from Sapoba (Benin) west of the
Niger, the other from Ikot Mbo (Calabar) well to the east; and from Ejura (Ghana)
come fairly large-spotted and very small-spotted skins. The last, however, is pretty
young, and it is possible that its markings would increase in size with age.
The pelage ingcncttoidcs is of medium length and fairly soft, and consists of abundant,
very fme, rather sinuous underfur 12 to 14 mm long together with finely stalked,
terminally broadened and flattened bristle-hairs measuring 19 to 21 mm. Ground-
colour is usually of a mediumly-dark grey washed with pale brovwi; but one veiy-
large-amiulate-spotted skin from a small island in the mouth of the Volta River near
Ada (Ghana, strand and mangrove vegetation, of very low rainfall) is a pale whitisii-
;i2 Tllr. ( AUNIVOKTS OI- WPST ArillCA
grey, not dissiiiiiljr lu toiio tioiu opcn-countrv panliua itsL-lt but witliout this hitter's
warmer creani-buft tint.
Alh of course, have the typical jhirdiihi group tail, short-haired, snaooth, subcyhu-
drical, and silky in texture. Tliis last is because the bristle-hairs, which are only some
19 to 2.Z mm iu lengdi, are slender and taper to a very fme base. The abundant iindertiir
IS about 9 to II mm long and plays a prominent role in determining the te.xture and
shape ot the structure. All this, wliile vastly different from the extremely long, coarse
bristles oi i^cncliii, is not luihke savalina; but m. ^cmttoidcs, as in others of the pardina
group, black is the dominant colour since the pale rings (mostly wliitish) are generally
much narrower than the dark and only completely encircle the tail in the basal halt.
The black end portion is usually at least 75 mm in length and often much more, with
2 or 3 halt-rings (or shorter) below, or sometimes with no trace whatsoever of white.
Skull. The bulla is shown in fig. 2Sb. Notliing appears to distinguish the larger-
spotted (jci/c/A'/f/c^' trom the smaller-spotted pocnsis. No comparison, cither of form or
of size, can be made with open-coiuitry pardina itself since no skulls of diis last are
available for study.
Taxonomy. Sutticient has already been said regarding this in the introductory
remarks to the group.
Habits. There are no collectors' tield notes turnishing any usetul information
peculiar 10 f^cncttoidcs. One specimen was trapped in a farm, one shot at night; and the
species is said to be tairly common in Sierra Leone. The first of these collectors' remarks
runs coimter to Sanderson (1940) who said that tins species in Camerouii (under the
name G. lifirina fieldiana) inhabited low trees m both high deciduous forest and ram
forest, occasionally entering old secondary forest, but never recently cleared land,
i.e. "cultivated land and tertiary growth". Common observation also casts doubt on
what Sanderson wrote, since these genets are tairly rehably known to come into open
cultivated compounds and firms for the purpose of stealing fowls.
GENETTA POENSIS Waterhouse Small-spotted Genet
Ot-ncttti piH'iisis W-itcrliousc, I1S3S, Proi. zool. Soi. Loud.: 59. Island ot Fcinando P(')o; but doubt was
cast oil this provenance by Pocock (njoSb). Type in the British Museum, No. ss. 12. 24. 41:!, sex ?;
skin only, in fairly good condition, but legs incomplete.
Description. The doubtful validity of this species (Plate 4) but its descriptive
usefulness have already been brieH\' discussed above. It remains to enter into this matter
a little more fully.
The species /)oi';).w.s has suttered niiire tips and downs than possibly any other form ot
(jcnctta. Waterhouse himself thought that his newly proposed species was nearest to
pardithi: and Matschie (1930), possibly acting on this opinion and seemingly never
having seen the skni or read the description with much care, s ynonymised the two.
Pocock (1908b) was astonished at this as "it would be hard to fmd two more dis-
similar species in the genus". This is very true, as reference to Geoffroy's plate and
diagnosis abundantly shows, and if these alone are taken as the sole criteria o{ pardina.
GENETTA POENSIS 2I3
The markings ofpoetisis (Plate 4) are widely different from the large, squarish, indepen-
dent blotches of fully typical pardina. They are very abundant, narrow, of much the
same width as the spinal stripe or even less {i.e. rarely more than about 10 mm), the
two rows on either side of this being largely united into similar almost black bands
but in fact comprising some 8 or more spots between the shoulder and hip. Besides
these partial stripes there are about four more longitudinal rows of roundish or oblong,
more or less independent spots on each side.
Spots apart, one of the first striking differences between the type skin of pocnsis
and other specimens of the pardina group is its golden-yellow groiuid-colour. Indeed,
with this colour, the abundant narrow spots and the forelegs being black, the skin at
first sight closely recalls scrvalina, but is at once distinguished by its typical pardina
tail with its lengthy black distal portion and indisrinct or partial pale rings. The authen-
ticity of tliis colour, however, has been called into question. Pocock (1908b), acting
on a hint from Oldficld Thomas, thought that the yellowish tone might well be due to
drying over a smoky fire. It is now not possible to be sure. The only West African
specimen in the British Museum at aU resembling it in its warm colour is the animal
herein named hitii: But there is a marked difference in that the reddish colour of the
latter is bright and hvcly whereas that o( the pocnsis type is dull, pervasive and seems
very likely to be adventitious. This is, in a way, confirmed by other specimens closely
conforming to the pocnsis pattern, Nos. 1938.7.25.5 and 57.12. 5.1, in which the ground-
colour agrees well with that of the ordinary run of high-forest genets of the pardina
group. The same may be said of No. 39.686 from west of the upper Cavally River in
north-east Liberia. But this specimen, which was figured by Pocock (1908), has some-
what aberrant features. In it the rows of spots adjacent to the spine are more con-
current than usual and also in places join the crest itself so that the medial dorsal pattern
is rather darker and more confused than in other examples. The fur of the back of the
neck is not in good condition but it appears that the linear pattern is there not so clearlv
defined as in the r)'pe and Kumasi specimens. Added to this is the fact that the tail is
almost completely melanistic, the pale annulations consisting solely of 5 rather obscure,
basal half-rings. Moreover it is very short, recalling what Matschie must have meant for
duhia; but it is by no meaiis certain that it is all there.
Pocock's scepticism regarding the identity oH poensis with pardina — or with ^c;)f«-
oidcs, which he would at that time have regarded as pardina — is excusable. Yet,
in the light of more abundant material than was available to Pocock, it seems to the
present writer that the two may well be demonstrated in the future to be the same,
pocnsis being at the small-spotted end of a pattern range. Nevertheless, until that time,
and in the present state of doubt, it seems worth while retaining the name even if
only for its succinct convenience in description. It nmst be noted, however, that were
it to become established that transition from large to small spots was in fact of this
nature, or that spot size was simply a matter of individual idiosyncrasy within a single
species, then the name pocnsis would have to replace gcnettoidcs, which it antedates.
Pocock was also inclined to doubt the island provenance o( pocnsis. In this, in view
of the early date of its collection and the imprecise conception of West African geo-
graphy at the time, he may well be justified. At any rate, nothing siirdlar has come to
2\i THE CAUMVOIiFS 1)1 V^' 1 S T AFRICA
light smcc troni rcniando Poo. One ot the skins referred to above as most closely
conforiniiig to the ty'pc came from Ghana, the other is, unfortunately, also an early
one and labelled merely "West Africa". It will be seen from the clinal nature of the
paniliui forms that it must often be dirticult to assign smaller-spotted specimens un-
hesitatingly to a specific form. Apart from the type and the other skins just mentioned.
It would seem that a few specimens from the closed forests of Ghana, Western Nigeria,
and possibly Cameroun, might be regarded as pccnsis.
Skull. There is no type skull; but that belonging to the skin which, apart from
colour, very closely agrees with Waterhouse's diagnosis o£ pociisis, No. 1938.7.25.5,
collected at about 64 kilometres north of Kumasi (Ghana), is far larger than any other
of the pardind group, being between 7 and 8 per cent larger in many measurements but
with a toodirow rather more than 20 per cent longer than average. It is tempting to
regard this as validly diagnostic and as confirming the specific distinctness oi. pocmis;
but it would be very ill-advised to jump to conclusions on the strength of this single
specimen. This is the more so as the only other adult skin closely resembling pociisis
(No. 57. 12.5. i) has a skull that differs quite appreciably in form and is of average size
wnth an average toothrow.
Habits. Nothing whatsoever has been recorded of habits specially pertaining to
piU-iisis.
D. thierryi group
The relationship to other genets of the species included imder this heading is obscure;
but, for various reasons, it seems to merit separation from the species-groups so far
dealt with. Since there is only one such species known it caiuiot logically be considered
as part of a species group, and it is so dealt with here solelv for the purpose of main-
taining uniformity of treatment.
GENETTA THIERRYI Matschic Hausa Genet
Gfm'/M thicriyi Matschic, iy02, Will, ties V intcriiatioiialcii ZooL-'i;cii-Ccii}<;rcsscs zn Berlin, tgor. 1142.
Hinterland of Togo from latitude 9'N. onwards. Specimen No. 19015 in the Iiistitut fiir Spcziellc
Zoologie und Znologisches Museum der Humboldt-Univcrsitat zu BerHn is herein, below on page
215, designated as lectotvpe. There appears to be no record ot exactly who Thierrj' was, in honour
of whom this was named; but since he was an Obcrleum.mt it is probable that he was an army officer
posted to Togo.
Psiudi.-'gi.ncua villiersi Dckeyser, 1949, Bull. Mis. natn. Hist. imi. Aims, (2) 21 : 421-424. Mcssirah, Senegal.
Type in the Museum National d'Histoire Naturelle, Pans, No. C.G.i9s2, No. 4 (I.F.A.N. 48.5.32
(453)), stuffed skin and skull; 6 paratypes in the Institiit Francais d'Afrique Noire, Dakar. Dr. Villiers
.ifter whom this was named is a well-known West African collector and zooloaist.
m
Taxonomy. It is necessary to discuss this fust before any pronouncement can be
ade respecting distribution and other matters. So ftr as published Hterature reveals
there does not appear to have been any critical re-exannnation of Matschie's material
since his erection of the species at the beginning of this century. There is therefore no
first-hand description additional to the rather inadequate accoiuit which can be put
together from his kevs. The characters there given vield the following: the spots are
GENETTA THIERRYI 215
small and very narrow, sometimes several united into a long stripe; colour is not
mentioned, but there are 4 complete rows with at least 9 spots in the first row, and
those of the tliird row further apart than their length. Both fore and liindfcct are
pale. The tail is short-haired, the hairs not longer than 25 mm at the base; some up-
standing hairs at the root; there are 10 pale rings, the last few visible only on the
underside; the light rings at most as broad as the dark ones.
By the kindness of Dr. Rcnate Angcrmann of the histitut fiir Spczielle Zoologie und
Zoologischcs Museum dcr Humboldt-Universitat zu Berlin it has now been possible
to study some of Thierr)''s specimens used by Matschie. There is no type o( thicrryi,
nor did Matschie ever label any of these skins with names. According to Dr. Angermann
(i)j lilt.) there is "a series of 10, all of them prepared in the same manner, all without
skulls and without precise dates or locality ("Togo")". Of the 3 skins examined in
London 2 belong, without question, to the pardina group; the third. No. 19015, agrees,
as near as can be told, with Matscluc's key characters for thicrryi and also with several
skins in the British Museum from the interior of Ghana — that is, from the area neigh-
bouring that from which Matschie described tliis species. The present writer, and
R. W. Hayman (1935) previously, had independently come to the conclusion that these
British Muscimi specimens represented what Matschie intended by thkrryi, an opinion
which has been strengthened to virtual certainty by comparison with the Berlin
material. Dr. Angcrmann has \vritten that the majority of the series of 10, not sent for
study, agree with specimen No. 19015, which is here designated as lectotype.
Before proceeding further with this question it must be said here that it does not
appear to the present author that there is any good reason to bcHeve that villiersi
Dekeyser is materially different from thierryi, or that Pseudogenetta Dekeyser, of which
genus villiersi was made the type species, merits separation from Ccnetta even as a
subgenus. This latter opinion was held also by Kuhn (i960).
Distribution. On this basis, specimens o{ thicrryi exist in the British Museum from
Senegal (Bakcl and an unspecified locality). Sierra Leone (Rokupr), Ghana (Ejura,
and Mole Reserve) and Nigeria (Wukrum Hills, north of Lau on the Bcnue). These
provide 1 1 skins and 7 adult or fairly adult skulls. The Berlin Thierry skins are labelled
as coming from Borugu, Togo, which would appear, from an old German colonial
map, to be the equivalent of Borgu, io°46N, o°34E, about 30 kilometres north of
Sansanne Mangu, just in the Sudan woodland zone. It is not known on what grounds
Matschie gave the distribution of the species as Dahomey and Togo westwards; these
specimens show that it ranges through the open woodlands of the Guinea, Sudan
and Sahel zones from at least the upper Senegal River to the western side of Northern
Nigeria. Further, and of considerable taxonomic interest, the last British Museum
specimen cited above demonstrates that three species, sau'g sp(.\'iiiicus ot riihdiJsoni collected b)' hini and nicasui'cd in the held, the mean length
ot all 6 specimens working out at 95 per cent of the mean head tV body measurement.
It is true that this does not accord with the figures given by J. A. Allen (1924) where
in all of 4 specimens, 3 of diem |uveiiilcs, the tail exceeds the head &: body. The greatest
excess of tail over body, 52 mm, m the Bates specimens occurs in his youngest animal,
a subadult: it is also a subadult that exhibits the largest difference, 53 mm, in Allen's
Congo series. The tail is clad with dense, more or less erect, iinderfur and only ver)'
slightly longer, soft, bristle-hairs. There are from to to 14 dark rings according to
species, somewhat narrower than the intervening pale ones, which mav sometimes by
faintly divided bv very narrow accessory dark rings.
There appears to be no pubhshed account of the detmite e.visteiice or absence of
scent glands; and no Uve or spirit material has ever existed in London from which this
question could be determined. But Pocock, who, was intensely interested in this
matter, said in a footnote (1933 : 970) that he h.id found I'vidence on made-up skins of
their existence in Poiivui.
Skull (tig. 30). Since no skulls ot lfii;liioiti exist in the British Museum the tollowmg
description applies more especially to richardsoiii. Apart from its considerably smaller
size the Pciiiiid skull differs most obviously from that oiGviutt.i 111 its lesser development
ot the posterior region. Although an occipital crest does exist it is relatively incon-
spicuous, often httle more than a ridge, and there is no very marked excavation of the
posterior braincase to rise again, as 111 Go/cffj, to elevated occipital flanges. Nor is
there any short posterior sagittal crest, as throughout that genus. It may be mentioned
here that in these matters the oriental Unsangs differ from the African, their skulls
more closely agreeing with those of the genets. The braincase is smooth and ovoid;
the postorbital processes may be sharply pointed, though never lengthy, or the\-
maybe poorly developed and blunt; and in 4 of the 7 London skulls this region is
fenestrated. The postorbital constriction is wider than the interorbital breadth, the
difference averaging rather less than 2 mm. The rostrum is narrow and pointed. The
zygomatic arch is strong and in most cases has a will-developed, pointed, jugal process;
but this appears to be a question of age since this section of the circumorbital ring is
more or less absent from the subadult skull. The bullae are tairly large, the posterior
portion far exceeding the anterior part, the two being conspicuously divided by a waist;
the meatus is large. The post-dental palate is relatively longer and narrower than in
(jciuriti. and the notches that separate it from the mam palate shallower and rather less
obvious than in that genus.
The dentition normally differs from that ot Gciictrii m having one less upper molar,
j-y-jrs — 3N; but III- may be occasionally present, though minute, as in B.M. No.
I. II. 21. 7, a medium-aged female; iii^ varies a good deal in the London specimens but
is on the whole more reduced than in Gaictia. In the iiiandibL-, which is relativeK
weak and flat, ;»l> is minute and seems, like its upper counterpart, to be on the way out.
The canines ot both jaws have niinute longitudinal furrows along their outer faces,
.1 character the\- sliare with Gcmtui aiiil Wiinlliil,:. and one so unusual th.it it must
assuredly indie.ilc phylogenetic atfinitv.
Habits. Exceedingly little is known of the linsangs iii life. Their apparent rarir\'
and purely local occurrence, their secretive and almost certainly nocturnal habits
have prevented their way of life from being observed save by a hmited number of
African hunters who, in the way of things, have never recorded what they know.
African linsangs, at least, seem never to have been kept in captivity, either in zoos or,
more locally, as domestic pets. Further, it is possible that more specimens have not
come to light because of some special value traditionally attached to the pelt. Over a
Fig. 30. Poiaiia richardsoiii: skull, B.M. No. yS. 3. 19.11, q, x i
century ago Allen & Thomson (1848) recorded: "This is a rather scarce animal at
Fernando Po, and as the skin is considered one of the most s.icred and valuable amulets
or charms of the Edeeyalis, they are unwilling to part with it. The small specimen in
the British Museum, presented by Dr. Thomson, was skinned from the mouth, and is a
proof of the mgenuity of that singular people". The special significance of these rare
skins noted by these authors in Fernando Poo seems to be reflected also in Liberia
where, according to Dr. Hans-Jiirg Kuhn in observations referred to below, the
^;.l Tlin CARNIVORKS OF WEST ATRICA
pi'lts arc usfd to make mcclicuic bags, that is to say containers tor materials regarded as
possessmg exceptional properties aiid worth. However, the mystical value of the
Pciiin.i skin doubtless varies with different individuals or may have suffered a decline,
since Seimimd, responsible for the greater part of British Museum collecting in Fer-
nando Poo, recorded that a t.iil and fragments of a pelt were "got m exchange for
rum '.
Habit notes boil down at present to two collectors. Hates c\ Kuhn. The former
(190s), who characterized Poiaiui richardsoni in Cameroun as a rather rare little beast,
said that it was foimd only in the forest, sleeping in the daytime on thick tangled vines,
and walking (? waking) onl)- when disturbed. A female brought to him in October
had miUc in two teats; and he was told that these animals produce two yoimg at a
birth. More recently, Kuhn, in an unpublished commimication quoted in Walker
(1964), as a result of intensive mammal collection and study in Liberia, has given the
most abundant information so far available. In this he states that the nests are round,
at least z metres and usually more from the ground, and ot green material, several
animals sleeping there for a tew days and then moving on to construct a fresh nest.
Further, although these animals have been recorded as using abandoned squirrel dreys
as shelters, he was informed by reliable Airican hunters that the reverse was the case,
squirrels taking over abandoned linsang nests. Kuhn says that the diet includes cola-
luits, insects, young birds and plant material. Doubtless it also covers small rodents
.md possibly reptiles as in the genets. This is die sum total of our present knowledge.
Taxonomy. It has often been assumed that the African Imsangs, Poiana, are closely
related to the Asiatic Imsangs of the genera Piionodon Horsfield and Pcirdictis Thomas
in spite ot being ver)' widely geographically separated with no known connecting
forms. Both groups are tropical rain-forest dwellers; and Misoime (1963 and 1965)
has shown that the existence ot any forest bridge between Africa and Asia later than the
Oligocene is improbable. The likelihood of either group's being an offshoot ot the
other, or both the relatively recent descendants of some common linsang stock is
remote; and the affinity between the African and Far Eastern genera is, in fact, con-
siderably more distant than often supposed. Poidita is without doubt more closely
akin to Gciutta than to Piioiwdoii and Pardictis. This is in some measure expressed in
Simpson (1945) by the allocation ot the two groups of linsangs to different tribes,
the Viverrmi and the I'rionodontim.
Resemblances which on the surface appear striking are on closer exannnation seen
to be not so ex.ict, and are assuredly fortuitous, the result rather of convergence than ot
artinirv'. The position cannot be more than sunnnarily glanced at here. Any corres-
pondence between the skulls is no more than frequently occurs ui this family; and,
111 tact, the supraoecipital region and the zygoma are of diverse forms in the t\vo
groups. Loss of the posterior molars, giving rise to similarity of dental foriiuilae, is ot
common e\olutionary occurrenee and of no overriding significance. Against this
numerical consonance, the forms of /ii and ot iiio ;ive distinctly different. Li the Asiatic
Imsangs b>>th of these teeth are somewhat more complex, p^, when not too worn,
Iviiig cle.irK tncuspidate; while ///a, in them, is fir more laterally compressed and more
sharply, luieveiily and line.illy cuspid. ite tli.in the lallier sqii.it qiiadi.ilc or ti langiilai
POIANA 22.S
tooth of Poiaiiii, bearing in side aspect a passable resemblance to a small premolar.
Finally, and quite significantly in a rare though slight character, the canines of the
oriental species show no trace of the fane furrows on their external faces so typical of
Poiana and Gcnetta. These cranial and dental disconformities are supported by external
ones of pattern, pelage and feet, that make it clear that close aft'uiity between the
African and Asiatic linsangs is more apparent than real. As Pocock (1908) indicated,
a character that would go far towards clinching the question of relationship, but
about which no information was available, would be the presence or absence of perineal
scent glands in Poiana since it is known tliat they are lacking from the Asiatic linsangs.
A quarter of a centmy after pointing this out Pocock (1933: 970 f.n.) had convinced
himself, by the re-examination of the same dried skins as were previously available to
him, that there was, in fact, evidence in them of the existence of such organs; and he
therefore concluded that the affinities of Poiana quite defmitely lay with Gemrta and
not with Prionodon.
Three forms of Poiana have been described: richardsoiii, ochracca and ki^htoni, the
two last as subspecies ot the first. No study skulls of these two exist, at least m London;
but from external characters it seems probable that whereas ochracea is merely a colour
variant of richardsoni, leigjitoni merits specific status. Only two of the three forms
occur in West Africa, ochracca Thomas & Wroughton, 1907, being described from the
Aruwimi River, eastern Congo. The two relevant species may be differentiated in the
follo^^^ng way.
KEY TO THE SPECIES OF POIANA
(previous key page 166)
Dorsal ground-colour dull reddish-browii; belly off-white; no continuous dark
spijial stripe; rmgs of the tail parallel-sided richardsoni [patic 22s)
Dorsal ground-colour bright bufi^'; belly pure white; a narrow, more or less
continuous dark spinal line present; dark rings of the tail rather chevron-
shaped. . leightoni (panic 227)
POIANA RICHARDSONI (Thomson) Richardson's Linsang
Genelta ricliardsoiiii Thomson, 1842, Ami. Mag. nat. Hist, (i) 10: 204. Fernando Poo. Type in the British
Museum, No. 42. 10.18. 1, sex unknown; unmounted young skin, in poor condition; no skull. This
was named in honour of Dr. John Richardson, Inspector of the Naval Hospital at Haslar. Thomson,
whose name was consistently misspelt Thompson by Gray in synonymies, was one of the medical
ofiicers to the 1841 expedition to explore the Niger and co-author of the published account. A note
in Gray's handwriting in the British Museum Register for 1842 indicates that it was he who actually
prepared the description "for Mr. Thompson".
Distribution and general. This is the better recorded of the two African linsangs,
the British Museum possessing 8 skins, one of which is juvenile, one fragmentar)%
and 7 skulls. These all come from the Bight of Biafra, that is to say more specificallv.
226 THE CAIiXlVDRVS Ol- WFST AFRICA
the island ot Fernando I'oo, Benito Jdwr m Spanish Ciuinca, and Bityc in lower
Camcroim, G. L. Bates being responsible for all the 6 mainland, and only perfect
adult, specimens. This area, of course lies wholly within the rain-forest belt. The
skin referred to by Pocock (1908b) as being labelled from Sierra Leone has iiot been
traced, nor any evidence fouird of its existence. However, one of the skulls. No. iof<4a,
was said to have come from Sierra Leone, the relevant note being in Gray's own
handwriting on the page so numbered of his draft catalogue but imder an 1854 Register
niunber that does not, in fact, exist. But, though purchased from a third party, it was
also stated to have been one of Eraser's collection — and, thus, with little doubt from
Fernando Poo. It bears a label to this last effect; and Gray himself must have come to
this decision since he quoted this skull m Gcrrard (1S62) and illustrated it, reversed left
to right. (1865a and 1869), making no mention at all of Sierra Leone as one oi^ Pciivui's
possible habitats. It does, hideed, seem highly luilikely that richardsoni docs, or ever
did, exist in Sierra Leone. PeiTct iv Acllen (1956) obtained a specimen from Foulassi,
Cameroun, which is not fir distant from Bitye; and Auerbach (1913) recorded one
from Yaoimde, also Cameroun but somewhat further north.
Description. Richardson's linsang (Plate 5) appears to be the larger of the two West
African species — though no measurements of Icigbtoni seem ever to have been pub-
lished; but it is still smaller than any known genet, to which animals it bears some
considerable superficial resemblance. Fiowcvcr, it has a yet more slender body; and it
is very readily recognisable by the complete absence of any continuous spinal stripe.
Its small, roimd or oval, fully independent black spots, and its very long, absolutely
c\lindrical, amiulated tail carrying 12 to 14 dark, parallel-sided rings.
The pelage of Richardson's linsang is short and soft, consisting of abundant, relatively
straight underfur, 10 to 11 mm long, with which are mixed comparatively few, only
slightly longer (13 to 15 mm) bristle-hairs. These are expanded subterminally into a
narrow, flattish blade but taper proxinially to a very long stalk almost as fuie as the
underfur. The base of all the pelage is medium grey, the visible pattern of the back
being produced by either brown or black distal portions of the hairs. The ground-
colour of this dorsal pattern is a sort of dull orangey-brown on which is superimposed a
pattern of black spots. Down the spine is a discontinuous series of narrow, and hence
rather linear, spots, on cither side of which are about 4 rows of roundish or oval spots,
well separated from each other. The back of the neck carries three, sometimes four,
more or less parallel longitudinal black marks; the medial one (or two) narrow, some-
times a discontinuous series of spots, and on cither side of tliis a far bolder, continuous
band. The undcrparts arc creamy or off-white, though the exact colour in hfe is difficult
to determine from old and hence rather soiled specimens.
The head is small, the ears big and rounded, the upper lips pale. The proximal parts
of both fore and hind limbs are spotted; the feet much the same colour as the back or a
little more sepia. The outside edge of the hindf'oot, in all but one specimen, is deep
blackish-brown. The long tail is remarkable in its narrow, almost perfectly cylindrical
nature and the regularit)- and clarity of its parallel-sided annulation from root to tip.
The colour of the light rings above is precisely that of the ground-colour of the dorsum ;
but below IS rather paler. The dark rings are deep-brown rather than black, that is to
Richardson's Linsang {Poiaiia richardsom); Two-spotted Palm Civet [Kaiidinia Iniioiata)
r o I A N A 227
say, noticeably paler than the dorsal niaculation. hi a number of specmiens, but not
all, a few, nuich narrower, relatively indistinct intermediate dark rings occur in a
few of the pale annulations. The composition of the more or less upstanding tail fur
is very similar to that of the back; abundant underfur 10 to it mm long, and bristle-
hairs 13 to 14 mm long; but widely scattered guard-hairs about 20 mm long, also
occur.
Skull (fig. 30). This has already been sufficiently described in the introduction to
the genus.
Habits. There is nothing to add to what has been given above under the generic
heading.
POIANA LEIGHTONI Pocock Leighton's Linsang
Poiaiia ricliardsciii libcrkiisis Pocock, 190S, Proc. zool. Soc. Loud, (for 1907): 1043. This name is rejected as
a /ii/'ji(5 calami in favour of that which next follows, Icightoui Pocock, as explained thereunder.
Poiana richardsoni Uifililoni Pocock, 1908, Proc. zool. Soc. Loud, (for 1907): 1043-1045, pi. 54, f 3. North-
east Liberia, 24 to 32 kilometres west of the Putu Mountains, situated west of the Duboc and Cavally
Rivers. Type in the British Museum, No. S.8.23.2, sex?; a headless flat skin, lacking also the forefeet,
but otherwise in fair condition; no skull. This name is accepted in spite of the apparent page priority
oi liheriaisis, which was an obvious lapsus calami. It has generally been assumed that the lapsus must
be in respect of the name Icightoui used in the key two pages later than lihcriciisis. But that Pocock's
real intention had been to give the animal the former of these names is clearly shown both by a printed
amendment slip m the Proc. zool. Soc. and the fact that tor his own private separates of his paper he
had the name amended on page 1043 of the text to leightoni before the reprinting. Pocock's pen,
therefore, obviously slipped in the inadvertent use of lihcriciisis, not vice versa. This name was given
in compliment to Mr. Leonard Leighton of the Liberian Rubber Company whose small collection
of mammals contained the r\pe.
Distribution. Although Pocock named this as merely a local race of richardsoni it
would seem, even from the very incomplete material m the British Museum that it is
probably a discrete species, geographically widely separated from the Bight of Biafra
animal. Pocock (1908b; 1038) mentions that Leighton obtained 6 skins; but only 2
ever came to the British Museum, the type in 1908, a second from the Zoological Society
of London, via Pocock, in 1939. There are no skulls. Kuhn (1965) gives the following
additional places in Liberia from which he personally has obtained specimens; Biple,
Bongle, Deaple, Duotown, Igua, Siamoiirovia, Tappita. on the road between Bia and
Zwedru. All these are in north-eastern Liberia, rather further north than the type
localiry, and west of the upper River Cess. The vegetation is rain-forest. F. de Beaufort
(1965) says the species also occurs in the Ivory Coast at Gagnoa. Like richardsoni, there-
fore, h-ightoni would appear to be a rare and very localized animal.
Description. Leighton's linsang is appreciably paler than Richardson's, the dorsal
groiuid-colour being a sort of golden-buff. As in the latter species the niaculation is
black, though appreciably more intense; and the spots are of similarly small size but
often of more angular outline. There is a definite, more or less continuous, narrow
black spinal stripe. The dorsal pelage is very short, vers' soft and springy, the abundant
imderfur measuring about 10 mm, the bristle-hairs about 12 mm but exceptionally
reaching 15 mm. The head and neck being missing from both British Museum speci-
228 Till, ,>
thioiighout till.- diMj.il pchigc. liic black, occmsikiuHv dci.'p-bro\vn, spots torniiiig
the dorsal pattern arc ot fairly small size and rather irregularly disposed, not more or
less clearly arranged in longitudinal series as they are in most Giitctiii. Broadly speaking
they are roughly siibcirciilar, about to mm in diameter, or less, mostly independent of
one another, though in a few cases tending to coalesce transversely. But in some speci-
mens they arc much smaller. There is no black spinal stripe or crest, although occasion-
ally in some skins there is a combination of spots that gives some impression of a
medial stripe.
The flanks are almost completely unspotted; the belly is yellowish and clearly,
though not sharply, divided from die flanks. The two yellow or creamy spots on the
shoulders are somewhat oval and at their maximum are about 25 mm long and vers-
plain to see; but they vary and are often indistinct or sometimes virtually lacking.
Forward ot these lie the markings of the neck. These are considerably variable but
basically consist of a medial black line reaching to the crown of the head between the
ears. This is flanked on either side by another, more or less parallel, black line reaching
to the base of the ear. There may be a row of dots, sometimes very faint, between die
medial and outer band; or tlris last may itself by broken up into spots: or some or all
t these elements may be entirely lacking. However, in most, though not all, West
African specimens the three main black lines are clearly present. The face is greyish-
brown, without distinct markings. The ears are veiw rounded, low 111 height but broad
at the base; and a bursa is always present, its semicircular posterior flap arising, bodi
top and bottom, behind the piima.
The feet are 5-toed, each digit armed with a very sharp, well-curved claw like a
cat's in shape and similarly retractile. They are slightly webbed between the basal
parts of the toes. The soles are very characteristic, apart from the naked, v.-ell-developcd
pads almost completely densely but shortly hairy. Li the forefoot the palmar and the
carpal pads are united; a central depression boimded by the four main palmar and two
carpal sections naked but of granular appearance. The ist di2;it of this foot is joined to
Its section of the palmar pad by a naked strip; and there are four small triangular
naked areas just anterior to the other sections. In the hmdfoot the plantar and tarsal
pads are similarly joined, but the latter unite into a single, large pad, very wrinkled
or ridged posteriorly and reaching almost to the heel. There are similai small naked
areas forward of the central pad and also joining it to rile 1st digit.
The tail is somewhat longer than the head ib\ body, bushy and woolly throughout
Its length but usually distinctly broader proxiniallv than distally. It is darker above
than below through the presence of long-black-tipped hairs, which in some specimens
impart a wholK blackish appearance to the terniiiial four niches. Its covering consists,
like that of the bodv, of abundant, long, dense underfur, about 30 mm in length:
and of sparser bristle-hairs, normally measurmg about 40 mm but which here and
there attain almost 60 mm. The tail may be looselv described as ringed; but it is very
irregularly so, nothing like the genets or Imsang, the "rings" being unevenly spaced
and mostly only half-rings across the dorsal side, or sometimes little more than broad
patches. The scent glands have been fully described by Focock (1915b): those of the
male are situated anterior to the penis, diose of die female in front of the vulva.
NANDINIA 233
Skull (figs. 3 I and. 32). This is considerably larger than that oi Gciniui but otherwise
superhciallv ver)- similar. The braiiicasc is long, ovoid, and bounded anteriorly by a
marked intertemporal constriction wliich is narrower than the interorbital breadth.
The supraorbital processes arc long and sharp. In old males and very old females there
is a pronounced sagittal crest, v^^hich posteriorly jouis a broad, flange-Uke supraoccipital
crest. The zygomatic arch is strong, almost semicircularly curved, the jugal process
slight.
The most notable pecuharity of the Niindinici skull lies in the bulla, the posterior
part of which is generally held to be entirely cartilaginous and is lacking from normally
prepared specimens. A detailed description of this auditory region is given by Hough
(1948), who finds it, despite the peculiarity now under discussion, otherwise quite
rv'pically viverrine. She accepts the generally held view that the entire posterior portion
of the bulla (the entot)'mpanic) is unossificd; Van Valen (1963), however, disagrees
with this, holding tliat "the dorsomedial side of the entorv'oipanic is commonly,
perhaps always, ossified in mature and nearly mature individuals, contrary to the
usual statement, although normally there is a cartilaginous region between the tympanic
and ossihed cntorympanic where they approach each other". Be this as it may, for
practical purposes of recognition almost ever)' prepared Naudiiiiii skull has the main
bulbous portion of the tympanic bulla, so conspicuous a feature of other carnivorous
skulls, lacking. Li this it is unique amongst the Hving (but not fossil) carnivora, and
exceptional as regards the mammalia as a whole. The anterior, wholly ossified, portion
around the meatus is often missing too, but for the entirely different reason that it has
been swept away through lack of its normal mechanical support provided by the bony
posterior region. The lack of a posterior portion of the bullae is accompamed by a
second peculiarity of the Nandinia skull. In other Feloidea, in which the bullae are
fully ossified, the paroccipital processes are closely applied to their posterior faces,
over which they spread to a greater or less extent; in XiVidiniii there is no ossified
bulbous portion to which they could become attached and they stand isolated as con-
spicuous components of the posterior part of the skull, long and pointed, and distinctly
more canoid than feloid in appearance.
The dental formula is ^-ia-i ~ 4-°" '^^^ upper mcisors are set m a straight or slightly
curved, compact row, the outer ones being somewhat stouter than the iimer ones.
The upper canines arc much straighter than usual, grooved on the outer face as in
Gcnetta but more obviously so; and also on the inner face, a character that is obscure in,
or lacking from, the genets. There is nothing remarkable about the rest of the check-
teeth except that tifi is very small, peg-like; and, in fact, considering the much greater
size of the animal the whole series is small in comparison wth Gawltn.
The lower jaw is strongly built, wth deep rami. The incisors arc bifid; the canines
are more curved tlian the upper ones and are similarly grooved on both outer and inner
faces, the latter sometimes rather indistincdy. The posterior molar is relatively small,
but nevertheless much better developed than the upper one.
Habits. In spite ot its being both widespread and numerous not much is known of
the life ot the two-spotted palm civet. Something has already been said, in the opening
-34
THE CARNIVOKKS OF WnST AFRICA
paragraph dcaliiii; witli this animal, ot its conjectured attinity with the Asian I'did-
iL'.xurus, in respect ot habits as nuich as of phylogeny. Though Wvuiiniii does chnib
pahn trees, and has been shot in diem, there is at present no significant evidence that,
despite its common Enghsh name, its habits are in any exckisive or predominant way
connected widi them, and particularly in so far as thieving palm-wine is concerned.
In an area such as West Africa where wine-tapping is so abiuidantly practised such a
habit, if it existed would be a matter of common everyday knowledge. This is not so.
Fic. 31. Naudinia liinotiila: skull, B.M. No. 48.814,
I ; Literal
Further, ex-Anglo-Lidiani, relyuig on parallelism with Asiatic species, have in the past
saddled NaiiJiiiid with the character of being "a noted fowl thief", certainly a more
credible failing in a viverrid than wine-bibbing. In combination of these two reputed
habits, the present writer was warned many years ago that it was foolhardy to preserve
oil-palms standing in one's compoiuid since there might always be a palm civet lurking
therein, ready to descend at night to rob the fowl-house. Whether this danger is real
is open to some doubt; for this small carnivore is, in truth, largely vegetarian. G. L.
Bates (1905), who has recorded more of this animal in its natural surroiuidings than
anyone else, wrote: "... there is no doubt that the usual food of the Nandine is
vegetable. It never catches chickens, as do other J'ivnjiddc" . And T. S.Jones has the
impression that ripe palm fruits form a significant part of the diet. On the other hand,
Thorneycroft (195S), writing in Malawi implied that the oiJy time die palm civet was,
as a rule, encountered, was "in connection with raids on the hen house". He also sup-
posed a pair he saw in the bush to be interested in a flock of guinea-fowl. And R. W.
Hayinan has noted {in lift.) that 6 adult specimens taken by him in the Ituri forest in
1930 were all lured by baits of oifal into traps set on the ground between die buttresses
ot giant trees.
NANDINIA
235
The fruit-eating habit is well confirmed by those who have kept Xiindinia in cap-
tivity'. Ball (1955) for example, who reaixd a yoiuig specimen in Nigeria, found it to
eat banana at an early age; and this remained its favourite fruit though it would eat,
also, pawpaw and avocado. This animal was, in addition, given crickets and other
insects, fat moths being very welcome; but as it was diftkult to obtain a sufficiency of
Fig. 32. Naiiditiia binoiaia: skull, B.M. No. 48.814, sc.\ ?, x i; p,iliital & dorsal views
such items the young palm Livet was eventually provided with a small lizard each day.
Such food had to be quite fresh, for even if it was only a little stale it would be refused.
This animal also accepted scraps from the table of a varied kind such as might come
from ordinary human mc.ils, including fish and pieces of meat; and it was very fond
of sweetened condensed milk and of chocolate. It liardlv ever drank water.
236 THE CARKIVOHf.S oy WEST AIRHA
All this IS doubtk'ss a t.iir indication ot tlic sent ot dietary m die wild: a good deal
ot hiiit tempered with insects, small rodents, birds or anything of like kind offering
Itself easily. Bates, in fact, records raids by a two-spotted palm civet, repeated over a
series ot nights, on the shreds of flesh still attached to the skeleton of a chimpanzee
hung up to dr\''. He also indicates that the fruits of the umbrella tree, Mnsaiii^ii cccro-
jHoiiks R. Br. and ot the climbing, yellow-flowered cucurbit Cof^iiiatixiii podolacna
Baill. are recognised to be such favourite foods of this animal that they are used by
local hunters to bait traps set with the object of capturing Kauduiia. Sanderson (1940)
foimd the stomachs of his many Mamfe specimens "nivanably" crammed with vege-
table remains — that is to say plantains when caught near farmed land.
The two-spotted palm civet is nocturnal. Under natural conditions it wakes at about
sunset and goes to bed soon after dawn, being, presumably, active during most of the
night. It may theretore sometimes be seen 111 the half-light at either end ot the day;
or It may be picked up by the red glow of its eyes in a torch in the darkness of the
forest at night. Since this animal is both arboreal and terrestrial such glimpses may be
either on the ground or on the lower branches of trees. The sight ot a pair of glowing
eyes peering down through the blackness can be rather eerie. These nocturnal habits
are basically retained in captivity but may be partially overcome in response to the
offer of food. Climbing for this vivcrrid, with its extremely sharp cat-iike claws, is a
matter of considerable ease; indeed Bates, in connexion with the nightly visits to the
chimpanzee skeleton mentioned above, makes it clear that the animal concerned in
this must have walked some distance clinging upside-down to the underside of the
ridge-pole of a hut. Its sure-footedness is also indicated by 13all (1955) who relates
how his domesticated animal used to spring onto the curtains, scramble to the top
and walk along a curtain-rod only 13 mm in diameter. Li climbing vertically up the
bole of a tree the claws may well receive assistance from the large, rough-surfaced pad
under the hindfoot. It always climbs down heaci first, Taylor (1970) giving a series of
pictures of the successive moves.
Probably the majority of its foraging, however, is carried out on the ground; and this
is one reason why it is fairly readUy trapped. When on the ground it either prowls in
feline fishion or proceeds at a trot, though not very tast. Thorneycroft (1958) records
an interesting incident concerning the palm civet's capacity tor leaping. He observed
one high up a tree, frightened by the barking of dogs at the foot, to sail gracefully —
"almost float" — to the ground 111 an effort to escape, descending at "an angle greater
than half a right angle", legs and tail fully stretched out. The landing on the bare
ground was perfect, on all four feet, at a considerable distance from the tree. This
performance was rapidly repeated from a neighbouring tree; but an attempt to reach
a third tree was foiled by the dogs. Taylor (1970) rates Xdiuliiiia as the most efficient
African vivcrrid at lumping.
During daylight hours the two-spotted palm civet curls up tightly and sleeps. Bates
said that it did this in thick tangles of vines in the tree-tops. This is a little surprising
in that climbers do not very commonly form tangles such as would naturally provide
suitable beds for quite heavy animals; especially in the upper strata of trees, at which
level lianes have generally achieved a good deal of independence. Tree-tops, also.
NANUINIA 237
would sccni to imply a good deal of sunlight, whereas Wmdiniii, in common wirii most
daytime sleepers, prefers deep shade. Tangles certainly exist; but at ftirly low levels
or in partially destroyed forest to which simlight has been admitted. It would seem
probable that for their daytime sleep these animals would more commonly seek the
shelter of holes or well-shaded crooks where large branches join the bole. There seems
to be no record at all of an actual breeding nest. This must almost certainly be a hole;
but whether up a tree or in the ground it is impossible to say. According to Walker
(1964) the period of gestation is about 64 days, the litter size being 2 or 3. Juvemlc
specimens exist in the British Museum captured in February, March, May, June and
August, thus indicating that breeding may take place in either the wet or dry season.
Two-spotted palm civets, if obtained very young, have shown themselves to make
charming, friendly and trusting pets, though suspicious of and aggressive towards
strangers. Given the free range of a house they have been said to preserve it free of
rats and snakes. Ball (1955) found his yoimg animal to enjoy games, even fairly rough
ones, with a dog, but not with a cat. This author also records that though his palm
civet had the habit of marking out certain areas with its scent glands it never at any
time had, itself, any offensive odour. Its fur, which was rather rough and bristly,
always had a pleasant, warm, musty smell. Bates heard two palm civets calling to each
other in the evening in the forest with a kind of faint kittenish mewing; Ball's pet
when pleased uttered a sort o[ cwiick-cwuck sound mingled with a purr; but when
amioyed gave a shrill, ratthng warning note. This animal was also responsive to human
utterances, recognising the sound of its master's voice, and answering to its name and
coming to a whistle.
Taxonomy. Three races have been described from other parts of Africa. These are
dependent on variations of colour or pattern. Some degree of variation is common ;
(. A. Allen (1924) writing of nearly 75 north-east Congo specimens said that there was
amongst them "a wide range of individual variation in coloration"; and the British
Museum West African material, of about 50 skins, displays appreciable variety of both
colour and markings. It is therefore ditficult to know how far races founded on these
characters may be justified. At least one Sierra Leone skin has a more intense colour
that recalls Cabrera & Luxton's ititcnsa from the southern Congo ; but it is not quite the
same, and the pattern is certainly less marked. And though nuchal striation becomes
obscure in certain other West African specimens there is none exhibiting the com-
plete absence of any trace of dark markings in this area that is the mam characteristic
of Thomas's (lerrarJi. J. A. Allen foimd this to be the case also with liis Congo series;
and since, moreover, there are in London several skins from East Africa with this
distinguisliing feature it seems probable tiiat i^crrardi is something more than a mere
individual extreme. Allen considered the third named race, arborca Heller, "to be
surprisingly different ". It would therefore seem that these races arc justified; and, that
being so, the West African two-spotted palm civet is correctly designated Xandinia
hinotatii hinolatd (Gray). If colour alone is regarded as a valid subspecific distinction it is
always difficult to know where to stop; but it is possible that the appreciably paler
skins of the Cross River basin (south-eastern Nigeria and Cumeroim) may merit a
distinguishing name.
;;,S Till'. CARNIVORES Of WEST AFRICA
I .iWe 12: Niiniciic.tl ilatj lur .\'im.
All these matters are dealt with in some detail and with illustrations in Pocock's
paper on the external characters ot die mongooses fig 1 fie), though by no means all
West Atncaii species are covered.
Skulls and Deutitiojl. As might be expected, mongoose skulls more closeK'
resemble those ot the iiearK' related genets (Viverrmae) than those of an\ other West
Afnciii carnivores. There are. .ipart from size, certain clear differences, which can be
seen b\ a comp.irison ot tig. Z'j w itli tig. 36. In dorsal aspect the rostrum m the ITerpes-
tiiiae, with the notable exception ot Lilwrliilis, is bro.ider; .md the also very much
bro.ider frontal area combines widi the complete or almost complete orbital ring to
gi\e tile structure .1 quite difhreiit .ippe.ir.uue in this region. In the veiitr,,! .ispect the
IIEnrRSTINAF.
241
much longer post-dciual palate of the mongooses is at once obvious; and the bullae
instead of being long arc for the most part much more highly and acutely domed.
Mostly the anterior portion of the bulla is very much smaller than the inflated posterior
chamber, with the single exception o{ GalcrclLi in which the two are subequal.
The dentition in the Mungotinae is remarkable for its variery. There may be 36,
38 or 40 teeth due to diversity- in the number of premolars, which may be ^ in Munoos
and Crossiircluis; j in Gi^lcrcUa; or j ui Ichncwniii, Giilcrisais, Xeiwgalc and Liberiiais.
Both Herpcstes and Atihx are capricious in this respect, the former having either 3 or 4
lower premolars, the latter exhibiting the same variation both above and below.
Occasionally skulls occur m these two genera with differing numbers of premolars
on either side. The form of the cheekteeth is also widely variable m the different genera,
being sometimes fairly typically carnivorous with sharp-edged flcsh-cutring carnassials
(Xaiogale, Atilax), sometimes sharply-cusped insectivorous {Afiiii(;os, Crossarchus),
and in one case, Galeriscus, fraiikly crushing. In some species the dentition is relatively
reduced in size, and this is particularly so in Libcriictis.
Fig. 33. Herpestinae: illustrating the two contrasting positions of the cheekteeth:
a. p"^ well anterior to the root of the zygoma {Idmeumia albicaiida, B.M. No. 25. 5. 12. 13) ;
b. posterior comer of p"* about level with the zygoma (Xeiwgale imso, B.M. No. T0.6.1.14)
There are nvo distinct rs'pes of herpestinc skulls m accordance with the positioning
of the cheekteeth in relation to the zygoma (fig. 33 a & b). There are two reference
pomts: firstly the extreme posterior comer of the upper camassial; secondly the
point, mostly fairly obvious though not exactly dcfmed, at which the lower curve of
the zygomatic arch arises from the main body of the maxilla, just above the toothrow.
In one category of skulls {Hcrpestcs, XfiW(;(ih\ GcilcrellT, Arihix), the posterior comer of
/!■' is situated at this point and, in consequence, /n^ lies wholly posterior to it, its outer
face forming a sharp angle with the outer face of the camassial. In the other category
{khncwnia, Gakriscia, CrosSiirclnis, Miingos, Libcriictis), the comer of ;)^ and all or at
least part of m^ lie anterior to the reference point, mostly very clearly so, but in some
Mii/Jijo.'; skulls the root of the zygoma is rather obscure. In this gi^oup the outer faces
of />•* and ml form oiJy a shallow angle with each other, or even a gentle curve.
Habits. Compared with other groups the habits of the mongooses have in some
respects been relatively well recorded, though often more as regards their everyday
behaviour as captive animals than as concerns the more secretive aspects of their lives
::.\i Tin ( arm\i)ui:s or wisi aiiika
m their natural environments. This is because, by reason oi' some ot their activities
and the exceptional readiness with which many of them take to domestication, they
have excited man's sympathetic interest over many centuries. Of all wild African
mammals there can be little doubt that mongooses make the most acceptable pets,
presenting much less difficulty of control and management than monkeys and offering
in return a certain indcfmable charm together with unending interest, amusement,
often surprise and sometimes apparent affection. The subject must, of course, be taken
in hand while still ver\' young; and, with this proviso, probably nearly all species
can be tumed into quite lovable, diverting, playful companions — and even useful
additions to the household since there is no doubt that one of these creatures helps to
keep the premises free of luidesirable pests, be they rodents, reptiles or cockroaches.
However, not all West African species have yet been tried out in this respect. Of those
that have, probably the kusimanse {Civssardnis) would lead as favourite — from the
point of view of size as well as ardency of spirit. Accounts of this and other species
will be found below in the appropriate sections.
Of all the activities or reputed activities tif mongooses that for which they are most
widcl)- famous is snake-killing. There is a good deal of misapprehension and misin-
tormation about this. It is often sweepingly beheved that all mongooses make a regular
practice of dehberately seeking out and destroying snakes, the ancient Eg)'ptians going
so far as to say that Hcrpcstcs habitually entered into long preparation tor the fray by
encasmg itself in layers of hardened mud as an impenetrable protection against the
tangs of its opponent. The information relative to snake-killing is by no means so
positive as generally supposed. Possibly several species will kill a snake if they should
happen in the course of foraging to come across one and it was not too large; but it is
extremely doubtful whether any would purposefully seek out snakes in general as a
matter ot sheer innate enmit)-, as popular renown would have it. Certainly as far as
the mongooses dealt with in this present accoimt are concerned Herpi'stes ichneumon
has since ancient times had a reputation, though seemingly unconfirmed by modern
observation, for an exceptional devotion to killing snakes; Mimgos mtinf^o and Ichiiciimia
dlhkandd have both been rehably reported as doing so and eating their prey; but for the
other species evidence is either lacking or merely hearsay. Indeed, Cansdale is quoted
m Hinton & Dium (1967) as observing that his tame kusimanse was afraid of even a
ilead snake ; and recorded ( 1 946) the same thing of a marsh mongoose.
There can be no doubt, however, that where a fight between a mongoose and a
snake does take place the former is aided to victory by two tilings: its great speed and
agihty which enable it to spring upon its enemy, make an effective bite and leap clear
before its opponent can strike; and the deceptive nature of its long-bristled coat in
full erection, giving a false impression of size and misleading the reptile mto striking
at what proves to be notliing more solid than hair. It has been experimentally shown
that some mongooses are markedly more resistant to the effects of snake venom than
other animals are, though they nevertheless succumb to a sufficiently large dose.
A peculiar activit}^ that is more certainly common to part, if not all, of the sub-
family is the practice of smashing objects by throwing them against something hard —
usualK' interpreted as being basically intended for cracking open eggs, but on good
HEUPKSTINAE 243
evidence applied to other things, sonic familiar as foods, some strange, and some frankly
useless. Tliis operation is carried out in two ways. The more spectacular is hurling the
object, with the forcpaws, horizontally backwards through the hindlcgs against a rock
or wall; the other is to stand upright, hold the object in the forcpaws against the chest
and then cast it with surprising force vertically down to the ground. Both methods
are very effective ; and if they do not immediately achieve the desired end are repeated
until they do. West African species that have so far been observed to act in one or the
other of these ways are Ichiieuiniti albiaiudd, Mnn^cs inni{i;o and Atilax paliidinostis.
As for more general habits, mongooses may be diurnal or nocturnal, sohtary or
gregarious. They are, by and large, pretty omnivorous, taking flesh when they can
get it, killing and eating small mammals, birds and their eggs, snakes, lizards, probably
frogs, crabs, sometimes fish, snails and, seemingly always as an essential mainstay,
insects, particularly orthoptera, beetles and their grubs. Some, at least, greedily eat
fruit; and in captivity more exotic foods. Not a great deal has been recorded of shelters
and breeding nests; but it would seem that all species normally use convenient holes
in the ground, in fallen logs, amongst tree roots or gaps between boulders. Some live
communally in so-called warrens but consisting usually of a single chamber though
with two or three entrances. It has sometimes been asserted that mongooses are generally
fairly closely associated with water. While this is clearly so in the case of some, such as
Atilax and possibly Hcrpcstcs, it is not so obvious with other genera; and without
question these animals are sometimes encountered at some remove from the nearest
river.
Little has been recorded of the various steps and behaviour patterns in raising a
family. Courtship play and coupling have been seen, as noted later, in the case ot
Muiiqos imingo; and some rare and very interesting though brief observations on the
period of gestation and frequency' of pregnancy are given in the accoimt o£ Crossdrchns.
Litter size can, to all intents and purposes, only be guessed at: 2 to 3 would seem to
be the commonest, but some species have been credited \%dth 4 or even 5.
One other peculiarity may be mentioned in this general sketch of the subfamily.
All mongooses, as already described, are furnished with anal scent glands, one of the
chief uses of which is for the demarcation of territor)\ This activity is carried out m
two different ways: one is by squatting down and dragging the circumanal sac across
the ground; the other by the much more imusual method ot standing erect, upside
down on the forepaws — called in gymnastics making a "handstand" — and in this
position pressing the scent pouch against trees, rocks or other objects well above
ground level. The purpose of this is not altogether clear since the warning patch is well
above the nose level of any mongoose of similar species that should happen to trespass
on the territor)'. The scent, of course, diffuses and would doubtless be picked up at a
distance by a keenly attuned nose; but the advantage gained or the end served by
dehberately making it less obvious is obscure.
Mongooses have little direct economic impact upon mankind. Their sometimes
ornamental skins occasionally provide simple pouches, scabbards and so forth in \'illage
communities; and their flesh similarly helps to reduce animal protein defxcieno,- in
areas where the more conventional butchers' meat is scarce or irregular in supply.
2-11 I III ( A l!M\ OKI S Ol \VI S r AI KICA
Indirectly tlicy may be regarded as in some me.isiire benetlcial in lielpmg to keep in
check a varied list of creatures many ot which can be looked upon as undesirable;
but. on the other hand, it must not be overlooked that they also take birds such as
trancolms and guinea-fowl that are themselves of human food value; and that in
killing lizards and frogs they are disposing of potential insect destroyers. They them-
selves arc preyed upon by the wild cats and dogs, hyaenas, pythons and hawks. Certainly
they fill an important role in the complex network of nature; but the sum total of
their effect on man himself one way or another, is probably not vcrf great. Mongooses
of various kinds, mostly under very confused nomenclature, are credited (Stiles c\
Maker, 1935) with a long list of parasites, both external and internal.
Taxonomy. Diiker's (19.S7) unorthodox belief, founded on observation of living
animals rather than study material, that the Herpcstinae are in fact more nearly related
to the Mustelidae dian to the Viverridae has been referred to earlier on page 163.
The nomenclatural position in the Herpestniae is exceedingly involved. Had anyone
deliberately set out for malign reasons to devise a situation so complex as to be almost
beyond comprehension he could scarcely have bettered the labyrinthme confusion
that by chance came about 111 diis group. The binominal system of Latinised nomen-
clature was devised to overcome the imprecision of vernacular names; but m the
1 lerpestinac the absurd position arose wherein in using the name Muliiics miiiiiio it
became advisable, or even necessary, to make clear, m the vernacular, whether reference
was intended to the common mongoose of hidia or to the only distantly related
banded mongoose of Africa. This is only one of several traps and obscurities. Fortunately
1. A. Allen eventually went into the matter with extreme thoroughness (1919b and,
more fully, 1924) and by reasoned argument brought order out of chaos. In consequence
there is little need tor more than brief mention ot this past complexit)' here; and all
systematists must tor long be deeply grateful to Allen tor his painstakuig unravelment
ot so tangled a skein. Those to whom the matter is one of detailed interest must study
his accounts, where they will tuid a score of pages crannned with the history and
pitfills ot lierpestine nomenclature.
.Mien's tuidmgs have been prett)' generally accepted; but though, thanks to his
labours, the situarion has been cleared up as far as modern writmgs are concerned the
tact must always remain that the study of early literature is rendered highly misleading
.ind ditticult of comprehension without a good deal ot knowledge of this complex
history. So many scientitic names, both generic and specitie, and so many animals ot
no present interest to West Africa are involved in the muddle that no good purpose
would be served by their mention here; but the general warning may be given to the
West African student who attempts to derive intonnation from natural histories or
scientific literature published well into this century to be cautious in interpretation ot
Miiiioo<. Hcrpvslcs. Cro.Viiriliiis. iimn^o, linciatiis and iclimiiiihvi, in various combinations
with each other or with other genera such as the previously all-embracing t Vi'crn;.
hi this general note the main questions of present interest are diose ot the name
and the status of the mongooses as a taxonomic unit, hi Simpson's classification (1945)
they stand as a subfamily, the Herpestinae, of the Viverridae; but some authors, notably
I'ocock, had previousK' regarded them as constituting a fimilv in its own right.
HERPESTINAE 245
Pocock (1916c and 1919), examining the question almost purely from the standpoint of
external characters, felt strongly that this was their correct taxonomic rank; and since
at the time he believed Herpistcs to be preoccupied and Miingos comcquently the vahd
name for '"the typical mongooses" he called the family the Mungotidae. J. A. Allen
(1919b) showed this to be an error and the family, or subfamily, name to be more
logically and correctly derived from Hcrpcstcs. More important than mere nomenclature
is the question of the taxonomic rank to be assigned to the group. There is a good deal
in favour of a reassessment of the Vivcrridac and its component subfamilies. The
mongooses exhibit morphological differences, external, cranial and dental, that in
sum present a strong case for independent family recognition. But this is a matter
essentially beyond the limited scope of this present work. It involves questions far
wider than the mongooses, indeed the whole present conception of the Fissipeda. To
deal with such fundamental matters piecemeal in a regional work is probably to lead
more to confusion than clarity, and the major lines of Simpson's classification are
therefore adhered to for the purposes of the present account.
As regards the full generic use herein of Gakrclhi and Xcnogiile, the arguments will
be dealt with later under their appropriate heads; but it may be said in this general
account that, despite a broad overall resemblance of the skulls of these two genera and
that oi Hcrpcstcs, the present writer fully supports Allen's opinion that there is a sufficient
degree of difference, external and cranial, to warrant regarding these three as generically
distinct. It must be added that this view is not shared by G. Petter (1969) from the
consideration of tooth structure alone. Similarly, Oldfield Thomas's conclusion that
Gakriscus is generically separable from Bdcogalc seems, on several counts detailed later,
to be quite justified. His reservation oi GalcrcUa for cchracca alone, with the consequent
erection of Myonas to cover the remaining species in this complex is, however, not
accepted, the arguments in favour of such action being at present not fully convmcing.
Thus, in this work these 9 genera are considered to be valid for West Africa : Mimgcs,
Hcrpcstcs, Crossiirchiis, Atihx, Ichiietimin, GalcrcUa, Gcilcriscus, Xenogalc, Libcriictis.
These may be separated by the following keys :
KEYS TO THE GENERA OF HERPESTINAE
(previous key page 163)
A. Cranial characters
I. Condylobasal length of the mature skull about 92-96 mm; the rostrum long
and narrow (length of skull anterior to the postorbital processes very
nearly equal to that posterior) ; all the cheekteeth very small for the size of
skull (e.g. breadth of/)'' is less than 6 mm) . . Liberiictis {fiigc 336)
Mature skull usually either appreciably longer or markedly shorter than the
above; rostrum short, blunt and mostly broad (length anterior to the
postorbital processes nearly always appreciably less than that posterior);
cheekteeth relatively much larger ....... 2
246 THE CARNIVORES OF WEST AFRICA
2. The posterior outer corner ot p* lies very' close to the point where the outer
posterior margin ot the maxillary root ot the zygomatic arch arises, and
all or nearly all of in^ conseqently lies posterior to this point; the outer
face of »(i forms a sharp angle with that of /)'* (fig. 33b) ... 3
The posterior outer corner of/)'' lies well anterior to the point defuied above,
and all, or a great part, of »ii is consequently also forward ot it; the outer
face of /»! forms a very obtuse angle with that of p^ (tig. 33a) . . 6
3. Condylobasal length of the mature skull about 100 mm or more; the anterior
chamber of the bulla flattish and much smaller than the inflated posterior
portion ............ 4
Condylobasal length ot the mature skull under 70 mm; the anterior chamber
ot the bulla inflated and more comparable in size with the posterior
portion ......... Galerclla (pin;c 307)
4. Postdental palate about as broad as long; the interorbital breadth over 19 mm
and the cheekteeth light in build, the occlusal outline ot the narrow
upper carnassial being scalene, its anterior breadth appreciably less than
the external length ....... Xctiogalc (/niijc 329)
Postdental palate longer than broad; it the interorbital breadth is as much as
19 mm then the cheekteeth are strongly bmlt, the occlusal outline ot the
stout upper carnassial being more nearly equilateral, die anterior breadth
usually well over 80 per cent of the external length .... 5
5. Width of the rostrum at the canine alveoli nearly always less than 20 mm,
never much more; greatest breadth across the outsides of the cheekteeth
less than 33 mm; breadth ot />■* less than 7 mm; length of /;j2 less than
5-5 mm . ....... Herpcstes {pa(;c iCifi)
Width of the rostrum at the canines 22 mm or more; breadth across the
cheekteeth over 33 mm; breadth of;)'* over 7 mm; length ot ;»■.; over
5-5 mm. ....... Atilax (p,n^c 2gi)
6. Condylobasal length ot mature skull well over 100 mm; premolars ;j . .7
Condylobasal length of mature skull well under 100 mm; premolars y . 8
7. Posterior upper cheekteeth about as broad as long, subquadrate in outline;
upper canines straight, tall, above average size, laterally compressed and
with peculiar, sharp, knite-like anterior and posterior edges; postdental
palate broad (about 14 mm) ; interorbital breadth considerably greater than
the postorbital constriction .... Galcriscus (paiic ill)
Posterior upper cheekteeth generally markedly wider than long; upper canines
curved and of normal subconical form; postdental palate narrow (mostly
well under 14 mm); niterorbital breadth usually rather narrower than the
postorbital constriction, occasionally a little wider Ichneumia (/'. ;(,'(' 300)
8. Occlusal surface of upper molars subtriangular in outline, the lingual portion
clearly converging proximally to a sharp apex; skull narrow, the zygo-
matic and mastoid breadths under jo per cent and 40 per cent respectively
of the condylobasal length .... Crossarclius {paoc 279)
HERPESTINAB 247
Lingual portion of the upper molars elongate and more or less parallel-sided,
the proximal margin rounded; skull broader, with zygomatic and mastoid
breadths over 50 per cent and 40 per cent respectively of the condylobasal
length Mungos {page 248)
B. External characters
The external features oi Lihcriictis are at present quite unknown.
1. Both fore and hindfeet with only 4 digits and soles completely hairy up to
the main pad; a very large mongoose with black legs contrasting with the
paler, usually grey, dorsum .... Galeriscus [page 121)
Feet with 5 digits (occasionally Galerella, small mongooses, have the ist digit
much reduced or lacking) ......... 2
2. Digits II to V entirely unwebbed; hind soles usually quite naked, but some-
times partially hairy; a large, most often entirely blackish-browai mon-
goose ......... Atilax [page 291)
Digits II to V with small webs comiecting at least the basal joints. . . 3
3. Pelage dark blackish-brown, slightly grizzled on the head and neck; size large
(head &: body about 520 mm); the long nose projecting well beyond the
Ups; hind-sole completely hairy . . . Xenogale [page iig)
Pelage of a lighter colour ......... 4
4. Entire forelegs and hindfeet almost black, contrasting strongly with the rest
of the long, loose pelage; the long-haired tail mostly unicolorous, white or
black, but sometimes coarsely parti-coloured; hind-sole almost com-
pletely hairy ....... Ichneumia [page 300)
Not like this 5
5. Size large, head &: body 500 mm or more; tail broad at the base, much
narrower distally with a conspicuous, contrasting, long black tuft; soles
naked ........ Herpestes [page 266)
Size considerably smaller ......... 6
6. Pelage short, close-lying, of fme texture and fmely speckled; tail terminating
with a conspicuous black or reddish tuft; head & body of small size
(about 350 mm) and very slender; claws very short; sole of the hindfoot
naked for about three-quarters of its length. Galerella [page 307)
Pelage longer, coarse; animals of small size but stocky build; claws of the
forefeet very long .......... 7
7. Pelage with abundant long miderfur; one or two whorls on the back of the
neck; nose very long; sole of the hindfoot hairy on the proximal quarter
Crossarchus [page 279)
Pelage virtually without imderfur; no nuchal whorl; nose of normal length;
sole of the hindfoot naked to the heel . . . Mungos [page 2^%)
248 THE CARNIVORES OF WEST AFRICA
Genus MUNGOS E. Geoffroy & G. Cuvier, 1795
African Mungos
Mimgcs E. Geoffroy & G. Cuvicr, 1795, Mugasin EiuyclopiJi.pic, Z: 184, 187. Type species I'ii'crra mungo
Gmelin. This is a fonnalisation of an Indian vernacular (Marathi) name, maiigus.
Aricia Gray, 1864, Proc. rcci/. Sec. Lond.: 565. Type species Hcrpates tacnianotus A. Smith. Derivation of
this name is not certain but is most probably trom Shakespeare's light and airy spirit Ariel, given
with reference to the swilt and lively movement exhibited on occasion by small mongooses.
.MuMjjiU Gray, 1864, Proc. zool. Soc. Land.: 575. Type species, by virtual tautonymy, Hapeslcs mungo
Desmarest(= Mungos (asciatus Gray).
Taxonomy. The fact that a vernacular name from India is officially, and properly,
applied to a wholly African genus is but one slight indication of the muddle that has
existed in the nomenclature of the mongooses. However, this point of etymology is of
minor importance compared with the chaos in scientific naming due to misimder-
standings regarding the proper application of the names Muuiios and Hcrpcsics. It is
not necessary to recount the tuU details here; those interested should consult J. A. Allen
(1919b or 1924). Nevertheless, the following brief summary is given as a caution to
those who might otherwise be misled by the study of bygone papers or books.
For a reason which today is not altogether clear Thomas, for whom the name in
its specific form had been "so utterly barbarous" that it could be ignored in favour
of a later one (1882: 90 fn.), suddenly (1907) substituted the use ot Mhiii^os for Hcrpcstes
which he had up to a short time previously (1906) employed in connexion with the
identical species, ijriin//.?. This use of Mungos for species previously known as Hcrpcstes
was continued by Wroughton (1907). At this time the specific name tnun<;o, imder the
genus Hcrpcstes, was that widely accepted as pertaining to the common hidian mon-
goose (e.g. Blanford, iSSS); and Wroughton, expressly stating that this was the oldest
specific name for tliis species, and without comment replacing Hcrpcstes by Mtiiigos,
thus introduced the combination Mungos niungo for this Lidian animal. Four years
later J. A. Allen (1919b) showed that Mungos was "untenable as a genus name for any
Indian mongoose" and should, in fict, be apphed to the banded mongoose of Africa,
to which, further, the specific name niungo was correctly applicable. There was there-
fore a sudden and sweeping change of meaning of the term Mungos mungo from the
common Indian mongoose to the banded mongoose of Africa. This latter apphcation
is accepted today without question; but without a knowledge of this history some
references in literature are liable to misinterpretation. It is, for example, used in the
now outdated sense of the common bidian mongoose in Pocock's important papers
and keys (i9l6e and 1919).
Only one other question of taxonomy arises in connexion with this genus : that of
whether it properly embraces Crosscirchus as well, either completely synonymously
or as a subgenus. In this present work the two are regarded as quite distinct, a matter
that is more fully dealt with imder Crossarchus.
Distribution and general. The genus Mungos is wholly African and, as at present
understood, comprises two species, nningo and givnhiiuius, the latter purely West
African, the former widely distributed, in many somewhat different local forms,
MUNGOS 249
throughout the continent south of the Sahara. Neither species enters the closed forest
as such though they may occur within the nominal forest belt marked on maps where
the original vegetation has been greatly destroyed or is mere open coastal scrub.
The genus is well represented in museum collections and may be reckoned as moderately
common in the field, gamhianus in West Africa, mioi^c elsewhere in the continent.
The latter species, however, seems from the paucit)' of specimens collected, to be rare
in the territory covered by this present work, its centre of distribution lying, apparently,
far to the south-east.
Descriprion. The two species differ markedly from each other in superficial appear-
ance, the one being conspicuously cross-banded, the other an irregular mixture of
colours. Apart from this, they are very similar in general form, being both of smalhsh
size and stout, with short legs and a short tapering tail. The contour dorsal pelage
consists of annulated bristle-hairs; in mungos the various corresponding colour zones
fall together thus giving rise to a marked and very regular transverse pattern of alter-
nating light and dark bands; whereas m fiamhianus they are randomly dispersed and
consequently result in a diffuse speckling. Both species in form, and one [gamhianus)
in general colouration as well, closely recall Crossarchns, and it is for this reason that
the latter has by many authors been identified with Miingos. But, the marked pattern
of nmngo aside, there are clear differences. Externally, the most readily observed of
these arc, in Muiigos, an almost complete lack of underfur; absence of an extra-long,
overshot snout; and a hindfoot whose sole is naked to the heel. Further, more detailed,
external description will be found under the two separate species.
Skull (fig. 34). This belongs to the category of small-sized skulls having a condylo-
basal length of less than 75 mm. Two other West African genera fall in this category,
GalercUa and Crossarchns. From the former, Mungos can be immediately distinguished
by the more forward positioning in its jaw of the upper carnassial; in Mungos the
posterior outer comer of this tooth, /)'', is situated anterior to the posterior root of the
maxillarv' process, with all or a great part of in^ also lying in front of this point.
Differentiation from Crossarchns is a more difficult matter; there is little that is absolute
to go on, it being almost entirely a question of comparative lengths or proportions.
The difficulty is added to by an almost complete lack of West African imtngo skulls
in the British Museum, data in respect of this species consequently having to be taken
largely from extralimital material, which amongst itself is pretty variable. The slight
differences can best be appreciated by a comparison of fig. 34 with fig. 37. The Mungos
skull is mostly somewhat broader; of 11 adult specimens measured only one had a
zygomatic breadth shghtly under 53 per cent of the condylobasal length; whereas
Crossarchns rarely exceeds 52 per cent and is almost always less. But the most obvious
difference in the dorsal view is the long narrow rostrum of Crossarchns in contrast to
the noticeably shorter, broader and blunter structure of Mungos. This discrepancy of
length is, in skulls from which the suture has not been obliterated, almost always
reflected in the mid-line length of the nasals: 16 mm or more in Crossarchns, 14 mm or
less in Mungos~hut there arc some extralimital exceptions to the latter. Also clear is
that the postdental palate, where it becomes parallel-sided, is in M. gamhianus and the
250
THE CARNIVORES OF WEST AFRICA
Fig. 34. Mun^os gamhianus: skull, B.M. No. 36.10.30.12, ,^, X .j'
MUNGOS 251
type of M. caurinus always conspicuously shorter than broad, though in other M. mungo
there are exceptions; in Crossarchus the two measurements arc about equal.
The general appearance of the Munt^os skull varies slightly. The postorbital con-
striction may be equal to or markedly more or less than the interorbital breadth. The
postorbital processes are mostly short and sharp, never, even in the oldest skulls, joining
up with the jugal processes to form complete orbital rings. A sagittal crest may be
well-developed but in most available West African specimens it is either poorly
represented or completely lacking, including well mature or oldish skulls. A supra-
occipital crest is clearly present even in very yoimg skulls and becomes broad and
flange-like in fully adult animals. The depth of the bones forming the zygoma is,
for mongooses, relatively shallow. The anterior portion of the bulla is smaller than the
well-inflated posterior chamber and carries on its ventral aspect a marked transverse
depression and fissure covering the greater part of its breadth; and this makes the
auditory meatus a flat oval.
In Muiigos there are, apparently with complete constancy, only 3 premolars on each
side, above and below. The outer cusps of the upper camassial are low and without
any marked cutting function; its inner heel, as with those of the two molars,
terminates in an almost equally high sharp cusp, and the whole cheek dentition appears
adapted more to an insect diet than to anything else, tn^ is a fairly big tooth, generally
somewhat wider transversely than the camassial; nfi is commonly wider across than
the lingual section of m^ but in exceptional cases is much reduced or may be lacking
from one or both sides. In the lower jaw the camassial is a tooth of little or no more
importance than any of the others; in young animals it carries three small, subcqual
cusps, the two lingual ones closely juxtaposed. These cusps soon become obliterated
with wear, as do those of »i2 also. In both jaws the canines are, for the Camivora,
relatively short.
Habits. What is known of these will be found below under the two separate species.
No general generic account can be given; for while there exists a number of observa-
tions relating to the continentally-spread mungo next to nothing is recorded of the
purely West African gamhianus.
MUNGOS MUNGO (Gmelin) Banded Mongoose
Viverra mwifio Gmelin, 1788, in Linnaeus' Sysft-ma Naturae, 13th edition, I: 84. Type locality' given as
Asia, but fixed by Ogilby (1835), Prcc. zool. Soc. Land. : loi, as Gambia; though Thomas (1882) beheved
it to be more probably the eastern part of the Cape Province of South Africa, and Roberts (1929)
wrote that "By common consent the type localit)' is taken to be Natal . . .". The name is derived from
the Marathi vernacular mangiis, being simply another form of the generic name.
Herpcstcs fasciatiis Desmarest, 1823, Dicnotmaire des Sciences iiaiurellcs . . ., 29: 58. Type locality given as
India, but as this was merely a renaming o{ immgo, above, the same considerations apply. The specific
name is a Latin word meaning enveloped with bands, given in reference to the pelage pattern.
Herpeslcs gothnch Heuglin & Fitzinger, 1866, Sher. Akad. IViss. IVieii, 54 sect, i: 560. Type locality
Kordofan. The specific name is the Arabic term for this animal. Possibly usable subspecifically for
certain West African specimens.
Crossarchus iMoti Thomas & Wroughton, 1907, Ami. Mag. nat. Hist. (7) 19: 374. Type locality Bomu,
north-east Nigeria. This was named after its collector P. Amaur)' Talbot, at that time a member of
Boyd Alexander's "Niger to the Nile" expedition. Available subspecifically.
2J2 THE CARNMVORES OF WEST AFRICA
Crossarchus fasdalus mandjarum Schwarz, lyij, }b. imssai<. Vcr. Naltirb. 68: 63-^4. Type locality Bate,
near the Uham River, Cameroun. This race was called after the Mandjia tribe dwelling around the
headwaters of the Shari River. Available subspccifically.
Mimgos aiurmis Thomas, 1926, Ann, Mag. nat. Hist. (9) 17: 1S2-183. Type locality Gunnal, Portuguese
Guinea. The name is from the Latin cmmis, the north-west wmd, because of the situation of the speci-
men's source relative to the main distribution ot the genus. Available subspccifically.
Distribution. Like several other mongooses — of the genera Hcrpcstcs, Atihix,
Ichncwniit and GidtnlLj — Miin^^os nuini^o has a wide distribution over the continent
south of the Sahara ranging from Senegal to almost 30°S. It is an inhabitant of the
open grass-woodlands, never of the closed forest, and seems to be centred more on
south-eastern Africa than anywhere else. At any rate, although fairly common in
that region it is apparently rare in West Africa, whence only 4 skins exist in the British
Museum. These are from Gumial in Portuguese Guinea (2), liornu in north-eastern
Nigeria, and Fort Laniy (Chad). But the species is reputed to have a wider distribution
in West AfHca than this would imply, for specimens were brought to England from
Gambia in 1835, which caused Ogilby to regard that as the real type locahty; and
Schwarz (191. <;) described the race numdjimiin from Bate near the Uham River in the
Central African Republic (about 6''4iN, I7°02E). These extremes embrace a wide
variety of open-woodland vegetation: Guinea, Doka, Sudan and Sahel.
Description. With its prommently and uniquely cross-banded pelage this mon-
goose cannot be mistaken for any other in West Africa (Plate 6). This is one of the
smaller mongooses, the head & body length being of the order of 380 mm, the tail
about 230 mm, and the weight i to 1-5 kg. It is therefore just slightly more bulky
than Crossdrcluis. The pelage is fairly long and slightly rough in appearance in life
though possibly not quite so much so as m the kusimanse. Li prepared skins it generally
hes very flat, and this is because it lacks the usual cushioning support of dense underfur,
this category of hair being almost totally absent. The bristle-hairs are of flat section,
about 24 to 31 mm long, and are markedly ringed light and dark. There are, almost
without exception, four regions of aimulation. The long basal section may be either
pure white or this colour interrupted by one or two indefmite blackish areas. The
three distal zones are always the same: a deep, blackish-brown ring, an off-white or
yellowish ring, and a black tip, usually longish. But there is no common length for
each of these zones, and especially the basal one, even in a single specimen; for since
the hairs arise at different points of the skin such irregularity is necessary for the corres-
ponding zones to fall coincidentally from dispersed points of origin and so produce a
total pelage effect of alternating light and dark transverse bands.
This cross-banded pattern applies only to the top of the back posterior to the
shoulders: it fides out on the flanks; and the shoulders, nape and head are an ordinary
random "pepper and salt" mixture. The hairs of the underparts arc plain or very
inconspicuously aimulated, and may be very sparse, leaving much of the belly almost
naked. The hairs of the throat are directed forwards and outwards; those of the chin
are directed backwards, so there is a pronounced line where the two meet; as there is,
also, along the side of the neck, backwards from the lowest point of the ear, where
the outAvardly directed hairs of the throat oppose the downwardly pointing neck pelage.
Plate 6
Kiisim;,mc {Crosscrclnii ohsam,>): C.inibMi,
Mongoose (A/,,,,,,...,.,,,,/,,,,,,,,,); Talbot's IJandcd Monaoos.
(.\/i(;/i>().< /»(,/,(.,i ,,,//„,„
M U N G O S 253
The head is broad and the muzzle fairly blunt. The largish, rounded ears are positioned
well to the side of the head. The tail, which is in broad terms about three-quarters of
the head & body length, is long-haired but not bushy, of a nondescript speckled
colouring like the anterior portion of the body pelage, but always with a blackish
terminal zone. It is evenly tapered from base to tip, very much in the manner of
Crcssiirchus. The legs are speckled ; but the backs of the forefeet and the digital area of
the hindfeet arc generally, but not always, dark, even blackish. The digits are webbed
between the proximal joints; the claws are very long, particularly those of the forefeet,
deep basally and fairly well curved; and the sole of the hindfoot is naked to the heel.
The scent glands have been described in some detail by both Chatin (1874) and
Pocock (i9i6e), the former dealing with the male, the latter the female. The sac
surroimding the anus and the orifices of the glands is subcircular m shape and very
large compared with most species. The glandular system itself is more complex both
in external appearance and in structure than in other mongooses; for, besides the usual
simple pair of organs with single orifices cither side of the anus, there are subsidiary
glands and ducts. The floor of the deep-wallcd sac is not a plain surface but is complexly
wrinkled in the male, and in the female comprises integumental folds or depressions.
Summarising Pocock's description of these latter, there are two pairs of depressions,
an upper and a lower, above the anus, and a lateral pair situated one each side of the
rectal orifice and more distant from it. Each of these six depressions contains a small
glandular pit with several secreting pores, none of which, however, represents the true
anal glands found in other species. The orifices of these, the normal glands, are to be
found on each side of the sac close to the inner margins of the two lateral folds. These
true anal glands arc a pair of large muscular sacs containing a dark-coloured, strongly
smelling, oily fluid which escapes into the sac. The smaller, subsidiary glands are very
different, of a more rudimentary nature in the female; but in the male, according to
Chatin's description, appear to have become more highly developed and to approxi-
mate to the true anal glands both in their structure and in the nature and amount of
their secretion.
Skull. A general description of the Mungos skull has already been given above;
here is not a great deal to add to this in respect of the species. There is, in fact, in the
British Museum only one adult West African skull of tnungo from which to derive
data for comparison with five adult gtuiihianns. Recourse is therefore necessary to
extralimital material of the former, and of this there is an abundance. Hayman (1936)
first drew attention to significant differences between the skulls of the two species.
The most unusual of these concerns the carotid foramen, which in the species now
under discussion is a largish round opening in the basisphenoid, but which in gnm-
hiimus is obscured, as described imdcr that species.
The inwii;o skull appears to be slightly broader than that o[ (^dinhidiius its zygomatic
breadth averaging about 56 per cent of the condylobasal length as compared with
only some 53 or 54 per cent in the other species. The posterior projection of the ptery-
goid process is narrow and club-shaped, terminating in a small knob. In the great
majority of extrahmital imiiifio skulls measured the interorbital breadth is greater than
the postorbital constriction, whereas in the available gambianus material it is almost
254 T'"- CARNIVORES OF WI.ST AIRUA
alw.iys less. The most obvious distinction between the two species lies m the teeth,
which in ijii/ii/x'./iiii.'.' are strikingly smaller. This is ni some measure brought out in the
table ot measurements on page 265.
Habits. Because it is not imcommon and ticcurs ui large and noisy parties in relatively
open coimtry the banded mongoose has possibly more frequently attracted attention
to itself and been observed than any other African species. Moreover, it has often
been kept as a pet or zoo specimen. There are consequently a good many notes on
certain aspects of its life and behaviour — though not a single one emanating from
West Africa. The most complete and up-to-date field accounts are those given by
C. D. Simpson (1964 and 1966) and Neal (1970). These relate to the animal in the wild
in the southern and eastern part of the continent and to captive or semi-captive indivi-
duals but can with little doubt be taken as applying equally well to the forms occurring
in the territory covered by this present work. The notes ^\■hich follow herein are
indebted in large measure to the patient work of both these observers.
The banded mongoose is probably wholly diurnal in its activities. Like the kusimanse
It limits in companies which as a rule number between half-a-dozen and a score.
Occasional packs of 30 or 35 have been reliably reported; but now and again claims
to have seen very much larger congregations, even up to 100, have been made. If
these estimates were accurate they concerned very exceptional gatherings. It is, in any
case, generally no easy matter, except at the nocturnal shelter itself, to count even a
moderate-sized band or merely to estimate their numbers with close accuracy since
at any given moment a large proportion of the part}' is on the move and often in and
out of the undergrowth. The members of the unit keep fairly close together but not
m tightly packed formation unless they are rravelhng purposefully from one given
point to another, say a new nest or fresh feeding grounds; or if danger threatens, when
they bunch together or fall into line, head to tail. The younger members of the group
may from time to time lag behind the rest, delayed by play and a lack of that fervent
application to the business of food-finding commonly exhibited by their elders.
Realising their separation they hasten forward and often actually run ahead of the
main body. Whether, in point of fact, a single individual ultimately determines the
general direction taken by the parry is unknown; but since it appears that some sort
of dominance is recognised between different members of the community it seems at
least probable that there is a tacitly admitted leader whose movements are instinctively
followed.
Throughout these foraging expeditions a fnrly continuous twittering is kept up,
partly the outcome of the excitement of the hunt, and pardy as a means of keeping
together by the creation of a communal spirit. Pocock (1916c) thought that it was one
of the functions of the anal glands in the gregarious mongooses to assist in holding a
group together by scent. This may be true in the more static sense of recognising and
clinging to known friends rather than risking the society of strangers ; but while on
the move it seems certain that sight and hearing, both highly developed senses in this
species, play the greatest part in maintaining close contact between all members of the
company. This, at any rate, was the opinion of Simpson. Nevertheless, should a few
get cut off from the main body for any length of time, by being frightened into taking
MUNGOS 255
cover, Ncal observed them to rejoin the main parry by pursuing exactly the same
intricate route as that taken by it, a procedure that could only be accomphshcd by
scent. The combined voices of a party of banded mongooses on the forage do, in fact,
carry a considerable distance. When it is necessary to sound an alarm or to demand the
full concentration of the pack the twittering takes on a shriller, more urgent note.
Foraging for food in the banded mongoose is no leisurely, part-time affair as it
has the appearance of being in so many carnivores. It is a highly active, continuous
performance, cverv' fully adult member of the unit displaying not only an astonishing
voracity but a dedicated eagerness and keen inquisitivcness as well. The party fans out
slightly and ever)- root, grass tuft, fallen log, rock, hole or crevice within the ambience
of each individual is determinedly explored, surface litter raked apart and such stones
as are readily movable turned over in the search for insects, their larvae, spiders, worms,
snails, lizards, mice, eggs, fledglings and other small creatures that may help to satisfy
seemingly insatiable appetites. Travel is therefore slow. An acute sense of smell enables
insect grubs, caterpillars and chn,'salids to be sniffed out though well hidden below the
soil or in tussocks. These are then rapidly dug out with the long claws of the forefeet.
Droppings of large animals are of particular interest and eagerly broken apart since
they arc often the source of a plentiful supply of insects, especially dung-beetles and
sometimes masses of dipterous larvae. Because of their richness Neal found dung trails
to be an important factor in determining the direction of the himt; and it is possible that
the paucity of such animals as elephants, buffalos and other large game in West Africa
accounts in some measure for the relative scarcit}' there of the banded mongoose.
Mostly, each individual hunts purely on its own behalf, quickly gobbling down what it
discovers before predatory interference from others can take place. But occasionally
larger prey is the object of communal attack; Simpson records having once wimessed
a pack of about 10 of these mongooses kill a large sand snake — Pstiniiiiophis sihilmis,
a species that occurs also in West Africa. Li such an attack the mongooses fluff out their
fur, dash in for a quick bite at the snake's body and leap clear before it can strike.
Simpson's statement that prey such as mice are killed by shaking, dog fashion,
is interesting as running contrary to the habit of the in many respects similar kusimanse,
in which Ewer (1968) observed that shaking was not carried out. Larger prey that
cannot be demolished at a gulp by one individual is pounced upon by the pack in
general, each member that can get at it ripping off as big a chunk as it can and carrying
it off to consume at leisure apart from its companions. There seems no doubt, however,
that insects are the basic food, and vast niunbers must be eaten in the course of a day.
In captivity a wide range of human foods is accepted; but in sharp contrast with the
kusimanse, fruit was found to be not very acceptable to captive specimens of muu(;o
though it is said that, as in a number of other small carnivores, fallen fruits form part
of the natural diet. If water is available these animals drink, by lapping, once or twice a
day; but though the species has often been held to favour the vicinity of water, com-
panies have been observed in prett)' arid territory where bodily water requirements
must be satisfied by extraction from food. It is possible that in such localities wild
fruits play a more important role.
No one has yet succeeded in following a pack continuously throughout the day,
2<,f> Till; t AIlNIVdUF.S OF WEST AFRICA
tliough both Simpson and Ncal have observed dieni for long periods, the latter with
binoculars from vantage points. It is not certainly kirown, therefore, whether or not
hmiting is, in fact, unremitting throughout the entire daylight hours. For any animal
to be persistently and so vigorously active as banded mongooses from dawn to dusk
would be a severe tax of energ)', and it seems at least possible that they, like so many
other animals, large and small, nocturnal as well as diurnal, suspend their pursuit of
food for a period, or periods, of rest. Be this as it may, at the end of the day, at least,
the entire company seeks the shelter of a common warren. This commmial resting
place may occasionally, m suitable territory, be more or less on the surface amongst
boulders, tree roots or other convenient cover; but more often it is subterranean,
having as its basis a burrow originally dug out by some larger fossorial animal such as
an aardvark, or, more commonly, a hole beneath a deserted termite mound. This last
certainly seems to be the favoured site, particularly if it is on a slope affording a clear
view roimd, has partial shrubby cover and a not too distant water supply. Entrances of
convenient size are dug or enlarged with the long-clawed front paws which cast the
excavated earth backwards between the hindlegs much in the common hcrpestine
manner of "egg-throwing".
Once the general appearance of these warrens has been grasped it becomes, according
to Neal, fairly easy to pick them out. If one has been in occupation for any length of
time it can be readily identified by the piles of droppings on its summit and scattered
for some distance around. There are generally two or three separate entrances and
several feet of internal tiumels leading to a central chamber. In the deserted warren
excavated by Neal this chamber measured some i -o by 1-5 metres and had a mid-height
of 0-5 metre. There were also two side tunnels leading to small chambers; and it is
likely that this special accommodation is for the use of breeding mothers.
It would appear that more than one commmiity may occupy a single warren; for
Shortridge (1934) mentions "an unusually large colony of at least thirry individuals"
that daily divided into several troops which hunted independently. A warren may,
by and large, be occupied for a long period of months; though not necessarily con-
tinuously, for the pack may, possibly when food runs short in the immediate vicrmty,
temporarily desert it for other warrens within its hunting territory. Simpson observed
one pack to use three different warrens located within an area of more than a square
mile. Besides these more or less permanent nocturnal shelters there may be within a
territory other temporary "bolt holes" to which a foraging company can retreat in the
face of danger; and it is at least possible that such established refuges, above or below
ground, may be used for a recuperative phase of inactivity during the heat of the day.
Two general points concerning warrens may be mentioned here. The first is that a
good deal of noisy activity, clearly audible from above, takes place below ground;
and the second, that Simpson found the entrances to abound in fleas.
The extent of a hiuitmg range has not been closely investigated and there is nothing
to indicate the sort of area covered beyond Simpson's incidental observation just
quoted. Nor is there much information concerning scent-marking of boundaries.
Neal makes no reference to this at all; but Simpson recorded that there is little activity
of this kind during food foraging but that it was common at drinking sites. The methods
MUNGOS 257
of registering scent signals follow the usual pattern : pressing the anal pouch against
convenient trees, logs or rocks, sometimes at ground level, sometimes by raising the
hindquarters well into the air; and, besides this dabbing, marking by dragging the anal
region across the ground.
According to Snnpson (1966) the birth and nursnig of the young takes place "in a
grass-hned chamber deep in the warren". Whether this statement is derived from actual
excavation of the site or is deduced from the fact that a tame female offered several
alternative possibilities for a natal chamber always chose a dark box accessible through
a tumiel in a mound of soil is not clear. Sexual maturity occurs certainly in females
at the age of 9 or 10 months, possibly a httle later in males, hi heat, a good deal of the
more active, dehberate courtship seems to be carried out by the female, lying on her
back and wrestling with the male, or flattening herself against his back and rubbing
her vulva through the fur. Both sexes, however, become playful, jumping around,
pouncing on each other, or nudging with the head; and during this the male also
"scents" the female, though seemingly less designedly than in her case. His anal glands
become enlarged and exude quantities of a white secretion with which the female
unavoidably becomes covered during this precoitional romping. Finally, the two
animals generally chase each other roiuid and round in ever narrowing circles, or the
male may circle the female, his tail held high. Neal observed mounting to be repeated
three or four times, with a short chase between each; and on one occasion a second
male, which had been watching throughout, mounted soon after the first male had
separated.
The period of gestation is in the nature of 2 months, its precise length being as yet
undetermined. It is not possible to derive information regarding a preferred breeding
season in West Africa from the limited material available ; but it seems probable (C. D.
Simpson, 1966) that, like the kusimanse, these mongooses are capable of breeding more
or less conrinuously. From 3 to 5 young seems to be the general number at a birth;
but a litter of 6 has been recorded. They are born blind, with a dark skin and a fine
transparent pelage which, however, faintly reveals black cross-banding on shoulders
and rump. The eyes open about the loth day. At between 2 and 3 weeks the pelage
begins to become more apparent; and it starts to assume adult appearance after 6 weeks,
though it does not attain its mature coloration imtil the age of 3 months. The young
appear to be held in common; for in one community Neal observed that when the
pack left on the daily forage one female remained behind to suckle and look after the
eight young. And on return of the parry to the warren in the cvcnuig all the females
niu-sed any of the young without distinction.
The weight at birth is about 20 g but this increases very rapidly, though the figures
published by Simpson (1964) exhibit an extremely wide variation between different
individuals. One young male achieved what may be regarded as a fully adult weight
of 1-75 kg as early as about 5 months; an adult male, on the other hand, is given as
only 155 kg; but, an adult female reached the exceptionally high weight of 225 kg.
As regards longevity, one banded mongoose is known to have lived in captivity to the
age of II years; and another of 8| years.
Several miscellaneous matters of behaviour must be briefly glanced at. In connexion
258
THE CARNIVORES OF WEST AFRICA
With feeding it is interesting to note that the backward egg-hurhng procedure common
to several members of the siibtamily is well confirmed in inmijio. It has been observed
and illustrated by C. D. Simpson (1964 and 1966), Davis (1966) and Kiiiloch (1964).
The force with which this is carried out is surprisingly powerful; nevertheless, an egg
docs not always crack sufficiently at the first attempt, in which case the action is deter-
minedly repeated until success is achieved. This procedure is, however, not reserved
solely for eggs but is employed with other desirable but encased foodstuffs, some of them
necessarily strange to the mongoose; and, moreover, as KinJoch has shown, is applied
to a variety of quite unprofitable articles, including even a bimch of keys. One obser-
vation involving this behaviour, made by Davis in the Bronx Zoo, is of especial
interest in as much as it concerns large millipedes so commonly occurring in Africa.
A banded mongoose, discovering after repeated attempts that its teeth were imablc to
pierce a cc-iiled up specimen of one of these, fmally hurled it dirough its hiudlegs against
the wall twice, thereby cracking the hard chitmous coat sufficiently for its teeth to
effect penetration. This achieved, the millipede was then readily consumed. Neal,
in fact, found these myriapods to constitute a very important item in the dietary in
Uganda, imini^o droppings invariably showing clear and abundant remains of the
chitmous shell.
Lrke others of its kind this mongoose is almost wholly terrestrial and probably
normally has little urge to climb for any distance up vertical tree trunks. Its long, almost
straight claws are scarcely adapted to this; and more especially to getting down again.
Kinloch describes a few rather inept attempts at tree climbing made by his pet /;i/»njp,
in which, it is true, the upper branches of acacias were attained but descent thence
found too nerve-racking to be imdertaken. Simpson, too, in his opening paragraph
(1964) makes some mention of tree chmbing; but this may refer to a pack of banded
mongooses that, escaping from hunting dogs, scrambled to the top branches of a tree
that had been pushed over by elephants. The sloping trunk and interlacing branches
of this would, of course, considerably lessen the difficulty, bodi of getting up and of
getting down. Wliere there arc positive footholds such as offered by the wire netting
of a cage it is known that iiningo can reach a good height from the ground. Apart from
an ability to scramble over low obstructions it can leap onto them provided they are
not more than 500 mm or so high. Though banded mongooses often live near water
Simpson says that they never seem to take to it of their own accord and are, in any
case, poor swimmers; on the other hand, Dalton (1961a) records how a pet of this species
leapt into a flooded river in order to rejoin her on the other bank and successfully
swam across.
Play is an important factor 111 the lives ot all young carnivores, and tins fact is con-
stantly illustrated by juvenile and subadult banded mongooses, both in the vicinity of
the burrow and while actually on the move durmg forays. Play follows the common
pattern of mock-fighting, chest to chest wrestling, tail-chasLng and scampering after
each odier. Simpson found, however, that play does not occupy much of the adults'
time. These, basking in the early morning sun or resting around the wan^en after the
day's hunt, go in for a good deal of grooming of dieir own or other adidts' fur. At
diese times, too, but especially in the morning after rising, they like to stretch their
MUNGOS 259
limbs out quite straight tore and aft, lying with their bellies pressed to the soil. Play
and other intercourse within the group, however, whatever it may appear on the
surface is not necessarily on a socially equal basis; for domination of one animal over
another is established, in the observation of Simpson, by putting the foreleg across the
shoulders of the inferior animal and seizing this latter by the back of the head lightly
in the jaws. Such ascendency by no means always belongs to the male. Simpson noted
female dominance in two different groups, in one of which a female established her
position over two males of the same litter at the age of about three months.
All who have studied banded mongooses in the wild or had dealings with them at
closer quarters have been impressed by the obvious acuteness of the senses of sight,
hearing and smell. These animals, though recklessly courageous in the face of close
danger, are at all other times highly suspicious of possible attack and timorous to the
point of fleeing for cover at the least imagined threat. This caution and an obsessive
curiosity that is often vital to the discovery of food hidden 111 soil or vegetation lead to a
constant exercise of these highly developed senses. Both at the warren and frequently
during the course of the daily excursion one, or sometimes nearly all, of the group will
sit upright on the haunches, or if necessary stretch up fully erect on the hindfeet, and peer
round, intently watching and listening. High sounds seem to make more impression
than low; and there has been no suggestion that crackling is a terrifying noise as it is
to the kusimanse. Simpson noted that pet animals could pick him out visually at some
distance, and that lizards cUmbing trees or insects flying away were followed by sight.
Neal found that this constant vigilance and acuteness of perception rendered close
observation of a pack a matter of difficulty since a stationary Land Rover or other
strange object at about 40 metres aroused nervous suspicion, and the shghtest movement
was detected at 20 to 25 metres, at which distance, also, the click of a camera attracted
attention.
A display of wariness can be seen on the occasion of leaving or return to the home.
The banded mongoose is not so early a riser as other diurnal animals, mostly making
its first appearance an hour or so after sunrise rather than in the grey light of dawn.
Before actual emergence from the warren the immediate vicinity is carefully scrutmised
from the mouth of an ennance hole, and if all looks clear one or two adults come out
and, if it is an old termitarium, moimt to the top of the nest and stand or sit erect
scanning the surrounding territory. Then, if no danger threatens, there is a general
exit followed, as a rule, by a period of play, siui-basking, yawning, stretching, grooming
and defaecation before moving off^on the day's forage. The return home is commonly
effected a little before night closes in; though should the afternoon be heavily overcast
the pack, possibly deceived by the poor light, may arrive at the home territory con-
siderably earlier. The returning party never enters the warren direct but pauses to rest
and relax at a little distance from it. Thence, approach to and actual entry into the
nest is carried out in gradual and cautious stages, puncuated by the usual play and
grooming activities, irntil the final bolt down an entrance tminel as night falls.
Banded mongooses have a range of different calls and notes. That most commonly
heard is the rather bird-like twittering of the whole pack when on the hunt. This
may change when danger is suspected to a strident note of alarm; but if a single in-
260 THE CARNIVORES OE WEST AFRICA
dividual is scared or threatened it spits radier like a cat and growls, and if this proves
ineffective it alters to a staccato chatter ot rage — or, rather, ot blind fury, for at these
moments one of these mongooses will hurl itself without reserve in attack at whatsoever
has aroused its anger. At the other end of the scale, in moments of pleasure the note
uttered is something of a low whine or even a kind of soft purr.
The species appears to have few natural enemies apart from birds of prey, which it
quite obviously holds in considerable fear; for a hawk may swoop swiftly from the
sky, seize an mdividual and be safely away without the possibihty of counter-attack;
whereas the combined tooth-power and lightning agihty of 20 or 30 enraged mongooses
must be a considerable deterrent to almost any ground predator. Nevertheless it is
said that the larger carmvores such as lions and leopards will pursue and attack a mun^o
hunting party. As some sort of defence against assault from the sky, of which their
keen eyes give them early warning, these mongooses instinctively bimch together m
tight formation; and this may possibly have the beneficial effect of deceivuig hawks
and eagles, making them more hesitant of attacking an apparently large body than
they would be of attempting the comparatively simple task of picking up isolated
individuals.
The majority of mongooses, obtained sufficiently young, make excellent and
fascinating pets, and Mungos mungo is no exception. These animals, however, cannot
be kept confmed to small cages but must be given more or less free range. This being
the case it shoidd be fully realised before taking on the responsibihry for such a pet
that, while the effort is usually very rewarding, a great deal of time and trouble are
unavoidably involved m feeding, play and general watchfulness, together with good-
humoured patience in the face of inevitable accidents to possessions due to an insatiable
curiosity, and restraint sometimes in response to slight personal injury, the outcome
of an incalculable uncertainty of animal temper. The banded mongoose is at heart a
very friendly creature, with its owti kind, with humans, and often with dogs; m
domestication it becomes trusting and, at least apparently, affectionate. It is by nature a
cleanly animal, easily house-trained; and its bodily smell is, as a general rule, shght
and moffcnsive. Its feeding is a matter ot simplicity since it avidly accepts a very wide
range of foods. All this has been vividly brought out by Kinloch (1964)-
Taxonomy. A species such as imin{;o with a pelage composed predominantly of
light and dark ringed hairs must inevitably display a range of colour variation, within
a smgle population as well as, iir the wider aspect, in response to differing ecological
conditions. No less than 16 subspecies are, mdeed, hstcd in G. M. Allen's African
Checklist (1939) ; and although, in the type descriptions of these, general body and tooth
sizes receive occasional reference and rarer emphasis, the great majority of the forms
are, m fact, described almost purely on the basis of colour, and in almost every case
from one or two specimens only.
Without question, clear differences of colouring exist between specimens; but even
now, after many years of collecting, the amomit of material available for study is for
the most part quite insufficient to judge reliably the constancy of colour in any given
locality, and hence the real validity of presumed races. Thomas, erecting CMrinus on
the basis of two differing specimens, himself thought that "probably it will be foimd to
MUNGOS 261
grade hereafter into others of the banded forms . . .". Certainly as far as West Africa
is concerned it is futile to pretend to the accuracy of deterniuiation implied by a tliird
name. Four specimens alone exist in the British Museum from the region. Two of
these, from a single locality and both ascribed by Thomas to caurinus, one being the
type, differ very distiiictly from each other in coloration. The other two, from well
separated locahties but both ascribed to talhoti, one the type, also differ from one
another, possibly due to age, possibly not. Until a great deal more material has been
gathered together for study it seems pointless to go into fmer division than the species
and imply a degree of understanding that does not exist.
Moreover, for practical purposes in a work such as this present it is not possible to
go any further. The plain fact is that with only a couple of sharply different forms to
differentiate or ^vith comparative specimens actually in the hand such expressions as
"paler" or "darker", "pinker" or "buffer" may well be full of meaning; but with a
range of several dehcate nuances it is an almost impossible task to convey absolute
colour to a student possessed of a single skin. Very few users of this present work
have access to standard colour charts; and even with these at hand the task is still fraught
with difficulty since the selection of colours in them is too limited to find an exact
match for the slight nauances involved. Li any case the colours themselves vary quite
appreciably in different parts of the pelage.
However, since certain names have been connected with West Africa the foUowing
notes are given for what they are worth. At the outset it must be pointed out that
there is uncertainty regarding the precise appearance of even the nominate race since
three different African localities have been assumed to be the type locahty. If Ogilby
was right in thinking tliis to be Gambia — and there is no positive reason for supposing
his guess to be less accurate than those of Thomas or Roberts — then caurinus might
fall into the category of a synonym.
Mungos mungo gothneh (Heughn & Fitzinger) Kordofan Banded Mongoose
This was described from Kordofan (Sudan) in the Sahel zone of vegetation but has
sometimes, with possible justification, been assumed to range westwards through this
same belt at least to the eastern side of Lake Chad. If it does this there is no reason why
it should not range yet further. The only clue to its appearance given m the type des-
cription, and that almost useless, is that the white colour of the imdcrside is less well
developed than in zebra, an Ethiopian species. Measurements of a few Sudanese speci-
mens assumed, from their locahty of origin, to be gothneh are given in the table on
page 265.
Mungos mungo talboti (Thomas & Wroughton) Talbot's Banded Mongoose
The brief distinguisliing features of this race (Plate 6) from Bornu (north-east Nigeria)
were given as medium size and very pale coloration — though Schwarz later wrote
that it was not always so pale as Thomas & Wroughton had said. What specimens
Schwarz had as a foundation for tliis statement, and on what grounds those specimens,
differing as they must have done from the t^'pc, could be held to be tallwii is not clear.
A second specimen in the British Museum from Fort Lamy, and determined two years
later as this race, is, in truth, somewhat darker; but it is a juvenile and not strictly
s
THE CARNlVOliliS OV WEST AFUICA
comparable, liirthcr, the two may, thougli not certainly, have conic h'om dirterent
vcL!;ctation zones; Fort Laniy lies hi the Sahel, Aduia raddiana, type of vegetation,
but Bornii is a district covermg a large area which incluties both Sudan and Sahel
woodland. Comparison was made in the type description with sonudiais, the general
appearance being characterised as even paler than in that race; and udhoti was said also
to have a black tip to its tail, a feature that certainly does not show very clearly today
in the type, the end of the tail being missing.
Mungos imingo niandjariini (Schwarz) Schwarz's Banded Mongoose
Bate, whence the type specimen of this race came, is in the Central African Republic
(6'4iN, 17"02E) and in a much moister vegetation zone, Doka woodland, than is
usual for this species. Possibly for this reason its coloration is duller: Schwarz described
the form as ver}' like i.tlhoil but at once distinguishable by its darker, more yellow or
browiiish ground colour, especially on tlie head and neck, and feet that arc quite
bkick. He also ascribed to this race two odier specimens trom i .So km further south,
from the source of the Pama River, which flows into the Ubangi and is hence just
extralimital to this work and in die (niiiiea woodland zone.
Muiigos mtingo caurinus Thomas North-west Banded Mongoose
Thomas described this race as "rather small"; the skull of the t)pe, an old animal,
m fact exlubits little diflerence of measurement from Sudan specimens presumed to be
gotlinch. The general colour was said by him to be "nearly as dark as in typical M. iimngi.\
quite unhkethat of the pale M. lalboti of Bornu". hi the r\'pc the colour is certainly
relatively dark; but Thomas omitted to say that the specimen was in moult and that
in consequence tlie bands are obscure or entirely lacking from the posterior halt of the
back. A second specimen, yoiuig, taken at the same place (CTiinnal, Portuguese Guinea)
six weeks carher is very different; and if it is compared with the type q{ taihoti it will
be seen that though the flanks oi taihoti are paler the backs match exactly. The material
available is quite inadequate to form the basis of any expression of opinion on the validity
of this proposed race, the type locahty of which lies in the Guinea woodland.
MUNGOS GAMBIANUS (Ogilby) Gambian Mongoose
Hcrpistcs ^anihiamis Ogilby, 1835, Pioc. zool. Sot: Loud.: 102. Gambia. Type m tlic liritisli Museum
No. 55. 12. 24.22ft, 9; skin in fair condition but probably much faded since it was once mounted and
on exhibition, and with h.ilfthe tail missing; skull much damaged.
Distribution and general. Although there has been some confusion regarding the
cenus of this mongoose, it having tor a long time been held to be Crosfardius — and,
mdeed, so classifie'd in G. M. Allen's Checklist (1939)— there seems never to have
been the slightest question respecting the species. Quite difterent m general appearance
from its strikingly cross-banded near relative imiu\i,\ it does, in fact, bear considerable
external resemblance to the more distant C.rossarchus ohsciinn, with which it has in
consequence quite often been confused both in the field and in the museum (Plate 6).
The essentia! distinctions between the two are brought out m the section devoted to
this latter species and need not be further entered into here.
MUNGOS 263
This is a small-sized mongoose apparently entirely of West Africa, its known range
being from Gambia to western Nigeria. It occurs almost exclusively in the Guinea
woodland zone just inland of the high forest but may penetrate into the rather similar
Doka belt. However, it has also been obtained from Bonthe island on the coast of
Sierra Leone, the vegetation of which, apart from fringes of mangroves, consists of
sand ridges and sparse grass. How the species got there is an interesting speculation;
but its presence there implies that it may also be discovered in other stretches of coastal
scrub such as the Accra plains, Ghana. G. S. Child (private commiuiication) records
that packs are not i:ifrequcntly to be seen in the dry-season throughout the Borgu
Game Reserve area in western Nigeria.
The actual places from wliich the 1 1 British Museum specimens came are : Gambia,
unspecified (2); Sierra Leone, Bonthe and Dumbaia; Ghana, Ejura (4) and Bole;
Togo, Kete Krachi; Nigeria, near Shcpeteri, Ogun Forest Reserve. OiJy 6 of these
are adult.
Description. The overall colour o( qiwihianus varies somewhat with the area from
which the specimen comes ; but the broad general appearance is that of a smalhsh,
dark brown, heavily speckled mongoose, with a head & body length of some 350 mm
and a tapering tail measuring about three-fifths of this. The face, though not exactly
blunt, lacks the conspicuously long protruding snout so characterstic of the rather
similar-looking kusimanse.
The pelage is fairly long and fairly harsh. On inspection it will be found, like the
other species of this genus, iniingo, to lack any obvious sign of underfur below the
contour coat of bristle-hairs. These latter are flattish, but not quite so much as in other
genera. They vary considerably in length as regards extreme examples, from 30 to
52 mm; but the more representative range is from about 35 to 45 mm. The annulation
of these bristles is distinctive and serves as one of the points of difference between
this mongoose and the kusimanse, as explained in the description of that species, hi the
proximal half there may be four zones of coloration, but these are not always all
present. Exceptionally there is a basal white portion measuring 3 to 5 mm; the next
distal zone, present in most specimens, is black and averages 5 to 7 mm; nearly every
specimen exhibits the next zone, which is white and averages 5 mm ; and this is very
often, but not always, followed by about i mm of deep yellow, dividing the proximal
region from the three terminal, and constant, rings. First of these is a black zone the
length of which varies between different specimens, in a Ghana example averaging
II mm, in a Nigerian one 18 mm; there is then a golden-yellow subterminal zone
averaging from 5 to 7 mm ; followed by a fmc black tip which averages 5 mm, is rarely
less than 4 mm, and may reach 10 mm. Because of the much longer subterminal yellow
zone the overall pelage colour in (^ambiaims is almost always rather lighter than in
C. obscurus, the ticking having a considerably coarser appearance than the fme puctula-
tion of the coat in this latter. Unlike the nearly related timngo there is not the least
suggestion of the corresponding colour zones falhng together in the pelage and there-
fore no hmt of any regular transverse pattern.
The fur on the belly is very scanty and, for the most part, imicolorous red. On the
throat it is imicolorous white, or yellowish, and directed outwards and upwards
264
THE CARNIVORES OF WEST AFRICA
from a longitudinal nuxli.il parting. Where its direction conies into opposition with
the shorter, posteriorly directed hair of the neck it curves backwards and forms a more
or less distinct white line. The short fur with which it comes iitto contact is blackish;
so a characteristic giiiiihiiiniis pattern is formed along the side of the neck, from the ear
to the foreleg, of two longitudinal black and white lines, but much clearer in some
specimens than in others. The rounded ears, set well down on the sides of the head,
arc clad with short, speckled hair, both inside and on their backs. Since the nose is not
elongated the depth of the upper lip from the rhinarium to the mouth opening is short,
quite different from the kusimanse. The arms and legs are ticked with yellow in the
manner of the back, but the backs of the forefeet and at least the digital area of the
hindfeet are black. The toes are slightly webbed; the sole ot the hindfoot naked to the
heel. The evenly tapering tail is mostly speckled but has a longer or shorter black
termmal region. No one appears to have investigated the form of the anal scent glands
and pouch.
Fig. 35. MiiwiiL^s i^anthiaitjts: huWa, ■ 2
Skull (fig. 34). This follows the general MtDii^os pattern as described above under
the generic head. There are, however, certain points of difference from nmngo. The
most interesting of these, to which Hayman (1936) first drew attention, is the existence
in i^ainhiivnis of a projection of bone from the antero-intemal angle of the bulla which
at least partially covers and obscures the relatively small carotid foramen in the basi-
sphenoid, with which bone it eventually becomes fused around the latero-anterior
rim of the foramen (fig. 35). This structure appeared to Hayman to be unique in the
carnivores; but a similar bony projection is, m fact, to be found from time to rime in
other mongoose skulls, especially in }hriHsiis ichneumon, as for example m B.M. No.
95.9.4.6 and B.M. No. 33.3.3.5. It can be seen also, though less well developed, in the
type o[ Crossarchus iinsor\;ci, B.M. No. 10.4.8.7. Judging from the slender material at
present available, five adult specimens, this is a somewhat narrower skull than that ot
niun^o, the 2r\-gomatic breadth averaging about 53 or 54 per cent of the condylobasal
length. The mterorbital breadth is less than the postorbital constriction ui four of the
five adult skulls, in the fifth the two measurements being equal. Whereas in numfio
the postdcntal palate is generally slightly broader than long, in '^amhianu'. the difference
IS more marked, the length being only about halt the breadth. The posterior part of the
MUNGOS
265
The most obvious diffl^rencc bcm^m the two species, however, lies m the teeth
sTm ",f:r k;;i'rTr/""f ^'^^ smalJ very considerably less in bulk for .n Zt
imilar sized skull. The visual sharpness of this distinction ,s only partially conveyed bv
the measurements given 111 the Table below. ^ tonveyea by
noI^tS;^" 'VTu' '""^ ft[P°'" "°''^^g '' ^''''''' of tl^^^^^ not a single field
record a in^Tf ''v?' 7^ 'f'''°' '''^ '^^ independent observation havmg been
XtneaTthe ti"ot T t''^'' "" "^Z" '^ J°""'^ ^P^"'"™^ ^o- Bonthe wa^
young near the end of June. Li a personal communication he states that he collected
another young specimen at the end of September; and young or very yot^^ ones
t^rw^TheTh'Ih " ^.^-t'^-"gJ--n' and Februa'ry. if is ther^o^e nS even
known whether this species shares m any way the communal habit of ,,,,.,^0 and the
Table 13 : Numerical data for species o£ Mimgos
Vegetation
Number in mean
Condylobasal length
Basilar length
Palatilar length
Zygomatic breadth
Upper cheekteeth breadth
Nasals, length
Interorbiul breadth
Postorbital constriction
Braincase breadth
Toothrow (c — 111-)
p* length
("1 breadth
m^ breadth
"11 length
"12 length
Head & body
Tail
Hindfoot
Ear
mimgo imwgo
?gotlmch caiirimis
(ex Sudan) (Type)
gambiamis
RATIOS (per cent)
Tail/head & body
Zygom. br./condylob.
Braincase/condylob. I.
Braincase/zygom. br.
Palatilar l./condylob. I.
Interorb./postorb.
Sahel
Guinea
Guinea and
Coastal
scrub
4-
I
5
67-0
66-2
69-9
63-3
61 -2
65-0
33-7
32-6
35-1
37-3
39-5
37-4
23-1
22-8
2I"7
i8-3
—
i8-8
14'0
14-0
13-4
I2-I
13-8
14-2
26-9
27-4
28-3
22-8
23-1
22-9
5-3
4-4
4-3
6-5
S-6
5-2
45
—
4-0
4-7
4-1
4-2
4-4
—
3-6
—
38s
350
•^
23s
210
—
72
69
' —
26
23
—
61
60
57c< iiliih-tiiiioii: skull, li.M. Nn. 9.11.2.10, sex ?, x i
HERPESTES 27I
Skull (fig. 36). The skull is long and rather narrow, terminating anteriorly in a very
short, bliint but narrow rostrum. The ovoid braiiicase carries, with few exceptions,
a well-developed sagittal crest in males and females alike, posteriorly joining the
broad, flange-like supraoccipital crest in a T. The frontal region is fairly broad and,
for the most part, iirflated to a slightly higher level than the cranium, quite smooth
and falling away in an even curve to the rostrum, and also laterally to the orbits.
The ratio of the intcrorbital breadth to the postorbital constriction is pretty widely
variable being sometimes lO per cent less, sometimes lO per cent more. The post-
orbital processes nearly always meet and fuse with the jugal processes to form com-
plete circumorbital rings. The nasal sutures also fuse and disappear fairly early. The
zygomata are strong and broadened through most of their length. The postdcntal
palate is clearly longer than it is wide; the anterior part of the bulla small, the posterior
chamber very inflated, high.
The incisors, top and bottom, are in a compact straight row, the outer ones larger
than the inner. The upper canines are slightly curved, the lower ones rather more so.
There arc always 4 upper premolars on each side; and there are mostly 4 in the lower
jaw; but of 11 skulls examined 3 mandibles have only 3 premolars on each side, and
one has 4 and 3. The anterior premolar is always very much smaller than the rest,
a simple peg. The posterior outer corner of the upper camassial lies near the outer
posterior root of the maxillary process, m^ is of moderate size, short but wide, its bulk
about that of the buccal section of/)"*, its outer face forming a sharp angle with that of
this latter tooth. The outer and anterior cusps of the lower camassial (;»i) are markedly
larger than the iimer posterior one. i)fi is small, its transverse width about that of the
lingual portion of /h^, its bulk clearly less.
Habits. In spite of the fact that the alleged habits of this animal have been written of
spoken of, and handed down almost as folklore for many htmdreds of years the truth
IS that very little that is up-to-date and reliably factual has ever been recorded. The
ichneumon has a long-established and almost unassailable reputation as the prime
enemy of both crocodiles and poisonous snakes, having, besides courage, such dedica-
tion to the destruction of the latter that it was reputed in ancient times habitually and
deliberately to prepare itself for the fray by covering itself in successive layers of mud,
drying out each in turn, until it was enclosed in a thick fang-proof armour of hardened
clay. With this protection it was able to seize its opponent in its jaws; when, instead
of biting it to death, it fliuig it into the Nile to drown. As for its other foe, not only
did it, according to the ancients, destroy crocodile eggs but it also regularly and courage-
ously leapt do^vn the open throats of these animals, which commonly sleep with their
mouths agape, and ate its way through the internal organs, making an exit through a
hole gnawed in the belly wall. The crocodile was presumably supposed to remain
placidly on dry land imtil this intestinal timnelling had been brought to a conclusion
and a safe emergence achieved. Even modern accounts of this mongoose are invested
with some air of fantasy, as when Brehna (i88o) describes a family on the himt, the
male in the lead, closely followed by his mate, and she again by the young ones all in
line close behind one another "as diough the whole chain of animals were only a
single being, somewhat rescmbhng a remarkable long snake".
27- llll ( ARNINUKES Ol WKST AlKICA
Olio may suppose that such ancient and persistent stones, together with tlie Greek
name "tracker", must have some foiuidation in fict however slender. Also that there
should be some significant basis to religious veneration, although this last would seem
to be discounted by the sacred ibis, which has little to it beyond its rather striking
colciration yet was embalmed in countless numbers. The crocodile stoiy may well
be a simple enhancement of egg destruction as an important factor in. the control of
these greatly feared reptiles. Nevertheless, it has been suggested that even this last
reputed practice may be pure invention since nest-robbing would be difficult if not
impossible with the female crocodile constantly on the watch in nearby undergrowth.
But that Nile monitors (I 'araiius niloiiais) acting in consort can rapidly create havoc in a
cache of crocodile's eggs is well-established, and the marsh mongoose has been said
to be a great destroyer of these too (page 297); so there seems no good reason why
I Icrpcstcs should not behave similarly. However, no record has been traced of such
depredation having been reliably observed.
In regard to snakc-killing one must distinguish berween habit and ahiliry. While
ichneumons may kill and consume snakes met with in the ordinary course of foragmg,
it is very unlikely that plain inborn enmity would lead them purposeful!)' and regularly
to seek out these reptiles. The ancient Egyptians knew, and doubtless honoured, the
ichneumon's capacity' to attack and overcome so dangerous and feared an opponent
as the asp or cobra; and possibly the spectacle of such combats was from time to time
staged in or aroimd temples, where these mongooses were habitally kept as sacred
animals. Yet there seems to be no record of anyone ever having witnessed a fight
between ichneumon and snake in the wild; or, for that matter, any recent account of a
staged one. Hinton & Dunn (1967) in writing a general work on mongooses and all
that was kno\\ai of them obviously combed the literature pretty closely; but they
make no reference to any fight, even a put-up one, between a snake and an Egyptian
mongoose. Even such verbal accoimts from travellers as may from time to time come
one's way may be open to some question since there has in the past been a good deal of
traffic between India and Egypt; those who claim to have seen such set fights arc rarely
competent to tell one kind of mongoose from another, and the Indian mongooses
arc not unlike the iclincumon m appearance. Flower (1932), indeed, recorded that
Itinerant conjurers were often to be seen in Egypt with the small hidian mongooses
as part ot their stock in trade.
It is, in fact. Flower who gives the most positive, though very brief account oi the
habits oi Hcrpcsics, at least in so far as Egypt is concerned. He says that it is diumal and
crepuscular, never to be seen in the early morning but on the move equally in brilliant
noon-day sun and m die dusk ot evening. It must be noted that other observers state
It to be positively nocturnal or possibly nocturnal as well as diurnal; and T. S. Jones
(personal commmiication) says that in Sierra Leone it appears to be largely nocturnal.
Flower characterises it as hunting aroimd in leisurely fashion; and it seems to be little
disturbed by the presence of people or vehicular traffic since it has been known to cross
Cairo streets, force motor-cars to slow down, and to have been run over by a tram.
It is as diough the ages have passed it by and it still expects the protection and respect
that were its heritage in ancient Egypt.
IIERPESTES 273
Hcrpfsics is, indeed, very unsuspicious and easily trapped. It takes readily to water
and swims well, being sometimes caught by accident in fish-traps when it is attempting
to rob thcni (Pitman, 1954). It is, in fact, commonly, but not exclusively, riparian,
dwelling and hunting amongst reeds and other dense imdergrowth; at the same time
it is by no means averse to woodlands, open countr)' at a distance from water, and
even to montane grassland (Ansell, 1965). Wlien living on river banks it is said to
feed on fish and possibly crabs and frogs; but away from such a locality it takes small
mammals, birds, insects and reptiles. It has a doubtful reputation as a fowl thief. T. S.
[ones states that in Sierra Leone it will certainly attack chickens when it gets any chance;
and it has been said to play havoc in a fowl house and to be detested for this reason
throughout modern Egypt. Flower (1932), on the other hand definitely states that the
peasants assured him that the iclmeumon never steals chickens — though a fairly well
authenticated record of the destruction of seven geese was once sent to him by a medical
officer. Flower's assertion is the more interesting in running directly counter to views
expressed by Zeuner (1963) that Hcrpesles had declined from the position of a sacred
and much respected animal in ancient Egypt to that of a widely hated pest in later
times due to the introduction of the domestic fowl from Lidia and its eventual spread
amongst all classes, with consequent bitter resentment of depredations amongst both
birds and eggs which had come to occupy a highly important place in the Egyptian
domestic economy.
Little, then, that is rcbable or not controversial has been recorded of the daily and
yearly hfe of the Egyptian mongoose, and it is to be regretted that no systematic field
study of it has so far been undertaken, as it has for some of the larger carnivores. The
fact is that, while such predators as lions, cheetahs, hyaenas, jackals or himting-dogs
are fairly readily observable and capable of being followed, the ichneumon, though
widespread over the continent and not uncommon, besides being of less immediate
economic moment presents considerably more difficulty. It spends much of its time
in rather concealed conditions of dense ground vegetation and can, as a rule, be caught
sight of only for brief moments as it hurries from one cover to another. Hoogstraal
(1964), however, states that sometimes these mongooses sit, play or move slowly in
easy sight of humans. The gait is normally a quick, purposeful trot.
For the rest, it is known that Hcrpcstcs shelters, and doubtless breeds, in burrows,
which may be of its own making or taken over from some other excavator; but gaps
in rocks, holes under tree roots and similar convenient cavities arc also made use of.
So far as limited observations indicate there seems to be no fixed breeding season;
there are no West African juveniles in the British Museum from which deductions
might be drawn, but one Sierra Leone specimen taken on 2nd November contained
two foetuses. Beyond this, defmite figures for the number of young 111 a litter are
almost non-existent; but though these animals probably lead much of their hves in
solitary fashion they have been said when raising young to go about in family parties
of about half-a-dozen. Like other vivcrrids, the ichneumon can emit an offensive
odour when alarmed or otherwise excited; and like so many other mongooses, when
caught young, it takes readily and kindly to domestication and makes an admirable
pet. In view of the fact that it has been known in this capacirs' for five thousand years
274 T'"- < ARMVORtb Ol WEST AFRICA
It IS remarkable that more has not been recorded of its behaviour, its nature, hkes,
dislikes, eating and sleephig habits, postures, breedmg, period of gestation, litter size,
parental care, voice and so forth. However, there is one detailed account (Diicker,
i960) of mating-play, copulation and parturition as exhibited by a young pair, of a
single litter boni in captivity. Play of sexual significance started between these two
when they were only about 15 months old; but real sexual readiness, evinced by a
swelling and redness of the vulva, did not come about for another 6 months. Con-
siderable wooing play took place thenceforward together with repeatedly imsuccessful
attempts at coition. The male frequently uttered an 00-00-00 sound, which was answered
by the female. At moments of great agitation this cry became something like hc-hc-
hc-hc, rising in speed and intensity to a pitch of excitement. When copulation was
actually achieved the act took some 4 to 5 minutes, during which the male repeatedly
nudged the female in the region of the throat with wide open muzzle. Two young
were eventually born at a half-hour interval ; but owing to the long-dra\vn-out nature
of the mating period it was not possible to tell the actual period of gestation. From the
dates indicated it might he an)n,vhere between, say, 7 and 11 weeks.
After the birth the parents were both very excited, ruiming and climbing about the
cage: but it was some time before they took any notice of the cries of their young.
At lengdi, after about a couple of hours, the mother carried the babies, separately and
held by the middle of the body, into the sleeping box, where they simply lay on the
floor while the parents sat snuggled up closely against one another taking no interest
in their oftspring. The only occasion on which they seemed to be aware of a parental
dur\' was when the cage was approached and they growled and erected their fur.
Next morning there was no sign of the babies, which must have been eaten by their
parents — a common event in nature, and more so under the unnatural stresses of
captivit)-. Ten days after parturition the female was again m season and the pair strongly
sexually interested in one another. This would seem to confirm that breeding is not
merely an annual event related to a particular time of the year. Also, it might be
inferred that two is a common litter size — there is the London record of two foetuses,
these two parents were themselves twins, and they had two offspring. No further
records were possible with this pair since the female suddenly died. It may here be
noted that a specimen has been known to have lived in captivity to the age of about
124' years [Int. Zoo Yr. Bh., 2).
Taxonomy. The broadest question is whether there is, in fact, more than one species
in the genus; that is whether the separation of the southern African animals at this level,
as differ, from the northern ones, once the practice, is valid. It seems fairly unanimously
held today that the species ichneumon satisfactorily and correctly covers all mongooses
of diis genus from Spain to the Cape; and this view is held in this present work.
With so wide a distribution it is obvious that local races must almost certainly exist;
but the problem here is not so easy of solution. With a pelage of the composition of
Hcrpcstcs, consisting of a top coat of long amiulated bristle-hairs in which black and
white play a dominant role there is, as in other similar genera, particular scope for an
almost infinite variet)' of effect due to slight differences of ring width or nuances in
the tone of the two main colours or the transitional zones between them. Some of
HERPESTES 275
these may be simply idios^ticratic; others may, in truth, be pecuHar to all the inhabi-
tants of a locality or set of conditions, though the plain fact is, as so commonly the
case, that existing study material is often too meagre to cstabhsh on a firm basis whether
such divergencies are sufficiently constant over a given area to justify nomenclatural
recognition.
Ten or eleven races oi ichncuimin are, in fact, currently recognised, only one of them,
occidentaUs Monard from Portuguese Guinea, dcfmitely West African, though it has
been suggested that parvidens Lonnberg from the lower Congo possibly reaches Lake
Chad. One incontrovertible fact exists, however: there are m West Africa two very
distinct colour extremes, the one pale grey, the other dark red-browTi. The former is
not ver)- dissimilar from specimens from the type area, Eg)'pt — which, not unusually,
differ somewhat amongst themselves. But in the West African examples the hairs of
the dorsal pelage have rather wider and whiter rings; the miderfur is orange, whereas
in Eygptian material it may sometimes be redder, sometimes drabber; and the belly
is better clothed, with longer, paler ("oft-white") hairs. Menard's description oi ccci-
daitiilis, though in respect of mdividual components detailed, fails to convey the overall
appearance of the coat. His type has not been seen; but by the courtesy of Dr. Villy
Acllen and Dr. J. L. Perret of Geneva it has been possible to make a direct comparison
of a small but adequate sample of the type pelage. From this and Monard's characterisa-
tion of the underfur as orange it seems sufficiently clear that the three pale West African
Herpesics in the British Museum are indeed occidciittilis. These are all from the Guinea
or Doka woodland.
The five dark form specimens, of very marked contrast to those just dealt with, are,
with one possible exception gone into later, animals of the closed forest zone between
western Nigeria and Sierra Leone. There is nothing m the large London collection
like these dark reddish-brown skms except an aberrant one from Spain, and they
would thus certainly seem to merit a descriptive name of their own. It might be deduced
from tins that Hcrpcstes ichneumon is pale grey in arid country becoming deeper and
redder as the atmospheric humidity increases; but there is an intermediate form which
comes not from an intermediate ecological zone but, so far as can be told from the verv
limited material available of each of these forms, from somewhat further inland than
the pale form, that is the drier Doka and Sudan zones. These skins partake rather
more of the reddish forest colour than they do of the grey; but the pale rings are broader
and whiter. Since they do not fit into either category they, too, though less obviously,
seem to justif,- a distinctive name.
The subspeciation described below hinges entirely upon coat colour; there are
two other specimens of West African ichneumon in the British Museum, one from
Gambia (no localit)') the other from Oda (Ghana), which since they consist of skulls
only cannot be particularised. No satisfactor)' evidence for the existence of Lonnberg's
parvidens in West Africa has been foimd in this present investigation.
Herpestes ichneumon occidentaUs Monard Western Ichneumon
Distriburion. Monard had two specimens before him when he named tliis race.
1-6 THE C:AIINIV0RES op west AFRICA
.111 adult tciiKilc h'oiii Mansoa and .1 male frmn Cachcu, both 111 Portuguese Guinea and
both, accorduig to the author, in dense tropical forest. The following are the particulars
and description ot the three London specimens believed to represent this race. They
come from Thics (Senegal), Cape St. Mary (Gambia), and Kita Sudan (Upper Guinea) —
which seems most likely to be Kita in the former French Sudan, now Mali. The first
two places are situated in Guinea woodland, the last in Doka.
Description. The overall impression ot the dorsal pelage of these is pale grey flecked
with chocolate (Plate 7). The furly dense, fme, not very long underfur, on the average
about I s mm though individual hairs may be longer, is, as Monard described, a beautiful
orange colour. This is almost completely concealed, in repose, by the very long (at the
lower back 60 to 65 mm) bristle-hairs. These arc ringed black and white, the basal
and two other zones white, separated by two major dark zones and ending with a
rather short black tip; that is to say 6 alternating rings. The "black" is seen on close
examination to be really a very intense red-brown; and there are some rings in which
this true nature of the pigment is more apparent than in others; and there is always a
brief transitional zone to the white where it becomes progressively less dense and hence
pale red. The coat, therefore, always displays a fnr amoimt of reddish colour, mostly
deep, which, as the corresponding zones of the hairs come to lie somewhat together,
tend to form short, very irregular, transverse lines or chevrons.
On the belly the imdcrfur becomes very pale buff, and the still very long bristles
cream, with one, or sometimes two, rather pale, inconspicuous dark rings. The hairs
of die head and fice have the same sort of colouring as the back but are quite short,
with correspondingly short annulation. The same can be said of the legs, where,
however, the dark clement is more prominent; so that, while the dorsal pelage may
be said to be grey speckled with dark, the legs arc dark stippled with white. The tail
is similar to the back m the broad basal half beyond which it becomes much more
yellowdsh-brown up to the long pure black terminal tuft.
Skull (fig. y>). It is not possible to draw useful conclusions from the single skull
available for study. There are in tliis only two marked differences from the remaining
West African material, and these may well prove merely idiosyncratic : the zygomatic
breadth is appreciably greater than in the others, amoimting to 55 per cent of the
condylobasal length as compared with rather less dian 50 per cent; and the breadth of
nfi is less than in any other specimen.
Herpestes ichneumon aithos mhip. iiov. Forest Ichneumon
Distribution and general. The contrast between this and the last is very marked
(Plate 7). Five specimens exist in the British Museum; these come from Newton (Sierra
Leone); Wulade, Kissi country (northern Liberia), 2 specimens; Idumiye Ono, 4 miles
north of Iseluku, west of Asaba on the River Niger, Benin Province, Nigeria (these
are spelt Idumuje Uno and Issale Uku on modern maps); and Senegal, no specified
localitv'. The first four of these lie within the closed high-forest belt, though Newton
has become degraded by fire and cultivation into grass with patches of secondaiy bush
near water. The last would seem to be possibly an exception, though, in the absence of
Plate 7
Forest Icliiicuiiicn {Hcrpcftcs idwaimon aiilios): Western Iclineunioii (HiT/Jcsfex Ulwaumv! oaichiialis):
M^irsli Mongoose {Alikx yw/i/i/iHusiis)
HERPESTES 277
an exact locality and the fact that in 1863, when the specimen was registered, the
limits implied by "Senegal" were not precise, one cannot be at all sure of the vegetation.
There is another ground for not blindly accepting at its face value this accrediting of
an apparently high forest mongoose to predominantly dry open woodland country
in the vicinage of the Senegal River. The collector was the late 19th century con-
troversial author Winwood Reade. It is a fact that on this visit to the then very httle
known and not very well comprehended western side of Africa he went to Gaboon,
Angola, Casamance, Gambia and Senegal, in that order. He might easily have said,
speaking generally, when visiting the British Museum that he had just returned from
Senegal and had brought some specimens for identification, thus conveying a wrong
impression; whereas the specimens may qiute wcU have been obtained at various pouits
of his tour; this one, for example, in the forests of Gaboon, whither he had gone
deliberately to follow the footsteps of the then famous collector Du Chaillu.
The Newton (Sierra Leone) specimen, collector T. S. Jones, B.M. No. 53.130,
a mediumly-agcd $, gravid with 2 foetuses, is nominated as type of this new race
because, although the skull is badly broken, the skin is complete; whereas the only
perfect skull in the material available has the skin incomplete.
Description. The overall impression of the dorsal pelage is a deep reddish-brown
abimdantly ticked with yellow. The underfur is of an appreciably deeper rufous shade
than the orange o( occidcnhilis; it is about 15 to 18 mm long, and though in a large
measure hidden by the bristle-hairs it does to some extent show through and help to
account for the reddish impression of the pelage. The bristle-hairs are not much more
than 50 mm long, and their annulation is quite distinct from that oi occidentalis; for
whereas in this latter race there are, including the black tip, about 6 alternating rmgs
in which the light and dark are not very different in length, in this newly proposed
form there are about 9 or 10 rmgs, the dark ones very much predominant, the pale
ones being mostly reduced to widths of 3 or 4 mm. The dark annidations are to all
intents and purposes black; but the two narrow distal pale ones, the only ones which
show beyond the underfur and constitute the ticking element in the pelage, are yellow
or off-white. Because of the vastly different widths of the aimulation in the two races
the actual physical appearance of the pale tickmg irrespective of questions of colour,
is very distinct in occicienUilis and aithcs, being in the former long and longitudinally
almost continuous, but in the latter very short, very much interrupted and numerically
very abimdant. There is a fuller note on annulation at the end of this generic section.
The belly fur, though in all rather paler, is very similar to that of the back, being
annulated yellow and blackish-brown. It is paler because the pale rings are far wider
than they are dorsally. The neck and top of the head are fairly similar to the back but
ticked more fmely; but whereas in ocddait,ilis the ticking continues forward almost
to the rhinarium, in this new race it stops short about the level of the eyes, the upper
part of the nose being clad with plain, d.irk blackish-browar bristle-hairs. The ticking
on the chin, throat and underside of the neck is whiter than that of the back. The
upper parts of the legs are ticked hke the back but the feet are deep red-bro\vn. The tail
resembles the back throughout nearly all its length but is for a short distance before the
black tuft very slightly redder.
278 THE CARNIVOliES OF WEST AFRICA
Skull. The measurements given ni the table on page 279 sliow that the zygomatic
breadth is less than in occidciinilis, amounting to 49 per cent of the condylobasal length.
The interorbital breadth is less than the postorbital constriction. Li the type skull
there are only 3 premolars on each side ot the lower jaw; in one other skull there are
4 on one side, 3 on the other; the third skull has tlie normal 4 on each side.
Paratypes. Other specimens exannned are No. 7.12.17.1 (skull broken), and No.
7.12. 17.2, both Liberia; No. 63.5.9.7 (no skull), Senegal; and No. 13.11. 6.1 (skull
perfect), western Nigeria. Although in these the shade of yellow in die ticking, and the
width ot the pale annulations do not absolutely correspond with diose given above
for the tvpe, the difterences are too slight to be ot aiiv account.
Herpcstes ichneumon niesos .<;i(/i.';^). noi\ Nigerian Iclineumon
Distribution and general. Three specimens in the British Museum collection are
assignable to this: from Aiiara Forest Reserve, near Kaduna, northern Nigeria; from
Kode area, 24 kilometres north-east of Lau (on the Benue), northern Nigeria; and from
the Northern Territories, C.hana. The first of these places lies 111 the l^oka zone, the
second in the Sudan zone; the third specimen, having no clue to die precise locality,
might have come from any of the Guinea, Doka or Sudan zones.
The Anara specimen, collector D. R. Rosevear, B.M. No. 50.320, a mediuni-aged
to old S> IS taken as the rj'pe of this new race. The skin is poorly stutted, the hmdteet
and part ot the tail missing; the skull in itself is complete but has several teeth missing.
Description. There is no doubt of the intermediate nature of this proposed new
race; m overall appearance it is appreciably redder than oaidcntiilis but a markedly
lighter-coloured animal than aitlios. Like other features, the underfur in this proposed
race is intermediate between the bright orange o( ociiiiciitalis and the rich red o( aithos.
It is about 13 mm long. As in occiiliiitdlis, the bristle-hairs have about 6 annulations.
Their overall length is somewhat shorter, about 50 to 60 mm; the pale rings, besides
being shorter, are buti and the dark ones are a less intense red-brown. A comparison
of this ammlation with that of the other races is made below at the end ot the account
of this genus.
The imdertur on the belly has lost most ot the red of the upper side, becoming pale
brown; the long bristles are butf and their dark rings relatively pale. The colouring of
the head and face is very much that ot the back but the hairs and their annulations very
short; the specklmg continues forward dorsally very nearly to the rhinanuni, there
being only a few millimetres of luispeckled hairs in this region, markedly less than in
aitlios. The forelegs are speckled, of a slightly darker tone than the back, the feet being
a deep brown; the upper part of the hmdlegs in the r\'pe is pale buff\'-browii with very
little obvious annulation; the proximal part of the feet is darkish with short speckled
hairs similar to the head; their end part is missing from die type, but in another specimen
is plain dark browni. The very long-haired tail is similar in appearance to the back;
its terminal portion is lacking from the type and one other specimen (No. 56.250);
but in the third specimen (No. 22. 12. 2.1) it becomes very rufous in the narrow part;
and the terminal tuft is not the intense black of the other races but a little reddish
throughout and more particularly so distally.
HERPESTES
279
The two paratypcs exhibit minor, but not significant, differences from the type as
described above.
Skull. This has no very distinctive features discernible in the two sets of measure-
ments available. It is of average width, the zygomatic breadth amounts to precisely
50 per cent of the condylobasal length. The braincasc is a little narrower than in the
other two races; the interorbital breadth is in one case (the t)-pe) larger, in the other
Table 14: Hair annulation in Herpcstes khneunmt subspecies
occiJentalis (7)
Basal region
Dark Pale Dark
Middle region
Pale Dark Pale Dark
17 II 7 13
Terminal region
Pale Red Black
7-5
= 60 mm
mesos (12)
aithos (5)
2 3 6
13 10 5 14
5 10 3 14
5 - 10
3 2 5
= 57 mm
= 53 mm
Table 15: Numerical data for Herpestes ichneumon
occidenlalis
mesos.
mesos.
aithos.
aithos.
Type
means
Type
means
Vegetation
Guinea and
Doka
Doka
Doka and
Sudan
Forest
Forest
Number in mean
I
I
2
I
3
Condylobasal length
99-4
96-4
98-4
100-4
I0I-3
Basilar length
92-6
88-7
95-4
—
94-1
Palatilar length
56-0
52-5
54-2
—
55-2
Zygomatic breadth
54-8
48-1
49-2
50-6
49-8
Upper cheekteeth breadth
32-6
29-2
29-7
3I-I
31-4
Nasals, length
24-2
23-5
Interorbital constriction
18-4
17-0
17-4
(l8-2)
i8-i
Postorbital breadth
17-9
i6-l
17-3
20-5
20-0
Braincase breadth
36-1
32-8
32-9
35-0
35-3
Toothrow [c — m^)
36-3
34-6
35-4
35-2
36-7
p* length
9-7
9-5
9-5
9-5
9-5
ml breadth
9-3
8-6
8-6
8-8
91
m- breadth
4-4
—
5-3
4-7
5-3
mi length
8-7
8-4
8-5
8-1
8-6
m2 length
5-0
4-3
4-2
4-6
4-7
Head & body
—
600
600
545
545
Tail
—
300
300
492
492
Hindfoot
—
—
95
95
Ear
—
26
26
36
36
RATIOS (per cent)
Tail/head & body
—
50
50
90
90
Zygom. br./condylob. 1.
55
50
50
49
49
Braincasc/condylob. 1.
36
33
33
34
35
Braincase/zygom. br.
66
68
67
69
71
Palatilar l./condylob. 1.
56
55
55
55
Interorb./postorb.
103
105
100
89
90
p*lc—m~
26-7
37-4
26'9
27-0
25-9
280 THE CAKNIVOUKS OF WEST AFRICA
less than the postorbital constriction. The type specimen h.is 4, tlie otlier skull 3,
premolars on each side ot the lower jaw. The breadth ot /»' and the leiiuith ot 1112
both seem to be less than in the other subspecies.
General comments, h must be repeated that no duect comparison ot the specimen
(No. 9.1 1. 2. 10) here assumed to be McMiard's caiJiinulis with his r\'pe has been possible:
if it should be shown to differ materially it would require a new subspecific name.
The table given on p. 279 shows a comparison of the lengths ot the various annulations
of the bristle-hairs in the three races, as illustrated by the mean values derived from a
number of diflerent hairs from each of a number ot ditlerent specimens. The ring-
margins are never clear-cut, the dark zone fading throughout a longer or shorter
distance (mosdy of the order of 0-5 to i mm) into the light zone, tliis transitional region
exliibiting a pale red-brown colour. Precise measurement is therefore never possible;
hut, with this proviso and the tact that there are always occasional exceptions, the
ring-widths arc pretty constant m all the specimens examined and for the most part
do not vary from the mean lengths shown below by more than 2 mm, usually less.
Three groups of zones may be recognised. There is a terminal region measuring
10 to 15 mm consisting usually ot two zones, the tmely tapering black tip and a pro-
ximal pale ring; but in dithos the transitional area between these two is of sutticient
width and importance to merit separate recognition. Proximal to this terminal region
is a second group of 4 annulations of the utmost importance in determining the appear-
ance of the coat; for whereas it will be seen, in the table, that the width ot the dark
rings remains almost constant throughout all three races that of the pale rings diminishes
very greatly, mi til in iilihos it is a half, or evcMi a third, ot what it measures in occidcniiilis.
It will be appreciated diat, ignoring colour, this results in a marked ditterence of
appearance, hi addition to the two sets of rings just detailed, iiithos has a further, long
basal region of 3 (and in one hair specimen 4) alternating dark and pale bands. The
length of this extra region is more than compensated for by the extreme shortening
of the pale zones of the adjacent region, to which attention has just been directed,
so that the overall length of the hair is, in fact, less than those with only 6 alternating
light and dark bands.
Genus CROSSARCHUS F. Cuvier, 1825
Lesser Long-nosed Mongooses
Miiiii^os E. GcofFroy & G. Ciivicr, 1795, A/iyiisiH Ei)cycl^j\iiiqii,\ 2: 1S4, 1X7. Type species I 'ivcrra iiiiiiif;o
Gnielin; in part.
Cwssarihus F. Cuvier, 1.H25, in E. GeortVoy & F. Cuvier, Hiiloirc Niiltiifllc i1t:< Mitiiiiniji-m . . ., 3, pt. 47,
Le M.inguc: 3. Type species Crossarclun obsainis F. Cuvier. West Atrici. This n.uiic was coined from
the Greek words crossolps fringed, and arclios anus, with reference to the appearance ot the circumanal
glandular sac, the wrinkled folds of which bear a fancied rescnibl.ince to the old-fashioned pleated
collar known as a ruff.
Distribution. The genus Croscs is available and to which it
should be restricted. Gray showed good judgment in separating the two groups
gcnerically. Attention has recently been called to the generic distinctness of these
groups by Pocock, he adopting for die banded mongooses Gray's unavailable name
.4nVln( pcrsica) near the shores of Lake Chad and the adjacent dunes; and it was
often seen at night on the outskirts of lakeside villages; but was never observed at any
great remove from the lake itself. Angus Buchanan obtained three specimens at Faniiso
(Kano, Sudan woodland) but no others occur in his comprehensive collections from
further inland in the more arid Sahcl and Subdescrt zones. Altogether, 21 skins and
15 skulls exist in the British Museum from Sierra Leone, Ghana and Nigeria. The
white-tailed mongoose may therefore be regarded as being, in the right localiries,
fairly plentiful.
Description. Icliiiciiiiiiii ranks approximately in size with Hcrpestes, AViiinjdlc,
.d//7.;.v and Giikrisciis, that is to say it has a head & body length of at least 500 mm and
a weight of some 4 to 5 kg; but it is not likely to be confused in the field with any of
these, Hcrpestes having a very sharply tapering tail with a contrastmg black tuft,
Xenogiile and .4fi7(j.v being mostly very dark animals, and Galeriais, despite its super-
30a llli; CARNlVOUliS Ol whj.t aikica
ticially very similar bl.ick legs and whitf tail, is contiiic'tl to the dense tdiests of the
Cross River. Ichncunm is, in tact, a very easily recognisable mongoose, with a loose-
furred buft or grcyish-biift coat more or less heavily splashed with black, wholly black
legs, and an abinidantly long-haired tail which in its most typical form is brilliantly
white (Plate S). 1 lowever, in West Africa it is at least as commonly black, there being
in the British Museum collection 7 white to 12 black tails, and 2 half and half; but
whichever it is, it and the black legs and teet bodi contrast sharply with the basically
buft body.
The lengthy dorsal pelage is, in texture, loose, fairly sott and springy. It is composed
chiefly of dense, relatively long, slightly wav)', tine underfur and extremely long,
oval-sectioned bristle-hairs. The underfur measures 20 to 25 mm and is buftish, pale-
brown or grey-brown and plays a considerable part 111 determining the overall colour
impression. The bristle-hairs are always vei"y long, reaching their ma.ximuni on the
lower back; the actual length varies a good deal from specimen to specimen, in some
measurnig about 50 mm or even a little less, in others attaining as much as 65 mm.
They also var)' considerably in their colournig. There is usually a white base, which
may be from 3 to 9 mm long, and then the remainder ot the hair may be either all
black or may comprise one or two white zones alternatnig with pale or very pale
brow-n. The white is never very conspicuous as such in the general impression of the
coat; but the pale areas of the bristle-hairs, together with the tuiderfur, make up the
basic pelage colour, die length and number of dark terminal zones determining the
degree of superimposed black. These two elements, wlule producing coats that are
recogmsably similar, account tor pale animals or tiark animals. On the underside the
fur is sparser and paler with many tewer bristle-hairs and, generally, much less black.
The fur of the throat is directed forwards and meets that of the chin at the angle of the
jaw m a transverse wave. In the very young the coat is always pale and contains very
fev,- black-ended bnstle-hairs, which increase in number with maturity.
The head is ot moderate breaddi, the muzzle furly pointed; both crown and face
are paler dian the rest of the upper pelage, speckled; and there is a bare area round the
eve. The ears are low in height, roimded and very wide; the outer portion situated
very much on the side of the head but the inner margins reaching much further up the
crown than in most other African mongooses. Their forward aspect is rather obscured
by a screen of long hairs rising m front ot them. The median groove ot the rhinarium
is conrinued downwards as a narrow bare band parting the upper lip.
The hindfeet and all of the forelegs and feet are practically black, but differ from
the similar ones of Gakriscus in having 5 toes in place of 4. These are webbed to the
subdigital pads. The hindfeet arc appreciably longer than those of other mongooses
of similar size and Icliiiciiiiiid tends to stand higher oft the ground; the soles are com-
pletely, or almost completely, hairy up to the pads. The very long-haired, tapering
and ver}' shaggy tail is somewhat shorter than the head &: body; but different collectors'
figures give widely differing degrees of this, var)'ing from 69 to 99 per cent. This is
probably partlv due to some measurements having been taken to the last joint, some
to die tip of the fur. But irrespective of diis there does seem to be a fiirly wide variation
of ratio troin specimen to specimen; and this has something to do widi the degree of
Whitc-tailcd Mongoose {Ichnvtitnut alhkauda)'. Black-tooted Mongoose {Giileri^ais nii^ripes)
ICHNEUMIA 303
maturity; Dalton (1961) recording that tail length increased enormously with advancmg
age. The tail, as already noted above, may be either white or black or half-and-half,
black prcdoniinatnig in West Africa, though relatively rare elsewhere. Maclnnes
(1952) records being mformed that both black and wliite tails may occur in a single
litter; but no positive evidence is given. There is the usual herpestine anal sac surround-
ing and, when closed, concealing the anus and the two laterally sited orifices of the
scent glands.
Skull (tig. 39). The West African material in the British Museum is very inadequate.
Although there are 15 skulls only 8 are reasonably mature, and of these 3 are broken.
If the skulls of the other large mongooses, Herpestes, Xenot^ale, Atilax and Galeriscus are
compared with that oi Ichncwnia it is seen that, although the distance from the glenoid
tossa to the condyles is much the same in every case, the length of skull anterior to this
IS, with the exception of Galeriscus, appreciably greater in Iclinciiiiu
26-5
2I-9-29-3
21-2
I8-6-23-7
22-1
20-3-24-0
3S-3
3 1 •7-37-3
40-2
3 8 •4-44-0
8-1
7-7-8-9
9-0
8-I-I0-0
7-6
6-5-8-2
8-4
7-7-9-0
7-4
6-5-X-2
482
423-545
361
330-395
107
99-121
40
33-50
75
SI
33
6S
58
96
20T
Genus GALERELLA Gray, 1865
Slender Mongooses
Galerella Gray, 1865, Proc. zool. Soc. Lond. for 1864: 564. Type species Hcrpestes ochracciis Gray from
Ethiopia. This name was obviously intended as a diminutive of the Greek gale weasel, a basic com-
ponent of a variety of combinations applied to a number of vivcrrids and mustelids, and in this present
case especially apt for the small, slender, weasel-like mongoose for which Gray devised this particular
variant.
Myonax Thomas, 1928, Ann. Mag. nat. Hist. (10) 2: 408. Type species Herpesles gracilis Riippell from
Eritrea. The name is a combination of the Greek words mys, iiiyos mouse, and ana.x king, implying.
lOS IHE CAKNIVOKLS Or Wl-Sr AFIUCA
111 Tlioiii.ii\ oun cxpLm.ition, "the King or I'yiaiit ot the R.its and Mice", leterriiig tci the reputed
ahilitv of thi<^ mongoose as a domestic pet to rid a house of rats and other vermin.
Taxonomy. Soiiictlimg has .ilrcady been said (pages 24') and 2(^(>) ot the question ot
wliether Giihrclla is synonynious with Ihrpcstcs or merits independent status: but the
matter must now be entered into rather more fully, before proceeding further, in order
to make the position adopted clear and to etiect detinition of the genus as nnderstood
in this present work.
Giihrclld was erected by Gray tor an Ethiopian species, ochiiiu\i, which he had named
some years earlier and at that time assigned to Hcrpcstcs. The proposed genus, however,
never came into use during the next 60 years luitil it was revived by J. A. Allen (1924);
and no new form has ever been directly ascribed to it as a genus except when 111 ii;35
Schwarz, following Allen's lead, used it in connexion with two subspecies of Siim^uincci.
There had, indeed, come mto existence only one new species that might have been
assigned to it, Heugliu's riificauJa, 1S77, between its creation by Gray and its virtual
killing by Thomas in his 1S82 review of the mongooses. Li this youthful paper Thomas
made a sweeping condemnation of Gray's proposals in which, he wrote, "such a large
number of untenable genera are formed, and so many bad species are made . . ."'.
He retransferred all of Gray's 1865 genera back to Hcrpcstcs and set a nomenclatural
pattern that lasted through several decades. In his more mature years, however, he came
(1929), as a result of J. A. Allen's 1924 paper, to see that, so far as the animals here at
present under discussion were concerned, he had made a mistake and that Allen was
"imquestionably right in maintaining that the small African Mongooses should be
considered as genericallv distinct from die larger Egyptian species, the t)'pe of the
genus Hcrpcstcs . . .' .
There was, indeed, no tmge ot doubt or ambigiuty about Alien's views through the
use ot minced words: "hi general features Hcrpcstcs and GalcrcUa are about as diverse
as two genera can well be and be referable to the same subfamily . . . ". He supported
this exceptionally luicomproinising opinion by contrasting the two genera in respect
of overall size and form, of the tail, the pelage, limbs, digits, claws and soles, as well
of cranial characters and dental formula. To sweep this aside by such a comment as
". . . it should be borne in mind that although the small African mongooses appear
very distinct from H. iciiiiciiiitoii (the type of die genus) there are many more small
species of Hcrpcstcs in Tropical Asia" (Ellcrman, Morrison-Scott & Haymaii, 1953:
119 fn.) seems unfiirly to diminish Allen's argument, in which size was only one,
the least important, character adduced.
Apart from the vast disparity of size and the altogether more slender structure of
Giilcrclla the points cited by Allen as divorcnig this genus from Hcrpcstcs are that it
has a pelage of a very diti'crent character; a narrow distichous tail; short limbs with
relatively small and weak feet in which the pollex and hallux are reduced: and soles
furred for mucli of the proximal half Li the skull, GalcrcUa ditlers in its uniquely
inflated anterior chamber to the bulla; ui the shape and siting of the postorbital con-
striction; and in the constant possession of only 3 lower premolars on each side, whereas
Hcrpcstcs most commonly has 4. The present author finds that tliese difterential charac-
GALERELLA 309
ters are valid and in sum lift the matter beyond the compromise course now often
adopted {e.g. Simpson, 1945; Ellerman, Morrison-Scott &: Hayman, 1953) of making
Gahrella a subgenus of Hcrpcstcs.
Li spite of Thomas's eventual broad agreement with Allen regarding the generic
independence from Hcrpcstcs of the small African mongooses, however, he at once
introduced into the controversy a second, cross argument in that he held that GalcrcUa
was applicable only to its type species, ochracca. This last, he maintained, differed in
its skull and in the nature of its feet from the remaining numerous forms, for which
he erected a new genus, Myonax. The skull in ochracca was said to be relatively shortened,
and Its bullae enlarged. The first is to some extent true; the second not so obvious.
But Thomas placed most reliance on the feet, wliich in Galcrclla, m his restricted
sense, he foimd "quite peculiar, very slender, narrowed throughout, with the naked
area extended backwards in a prominent line to the heel, the corresponding region
in the ordmary mongooses bemg hairy . . .". hi addition, the hallux is very much
reduced or entirely absent. Schwarz (1935) disagreed that anything more than a specific
difference was involved; and Ellerman, Morrison-Scott & Hayman (1953) adopted
the same attitude. Nevertheless, Thomas did, in fact, have something more of a case
than these contrary views might indicate. The skull differences he cited exist, though
sometimes in no very marked form; and close examination shows that without doubt
the hallux is either entirely lacking in ochracca or at least far more reduced than in the
other species ; and the nakedness of the sole does extend, in most cases, further towards
the heel. He could also have drawn attention to a difference in pelage character. But
whether these points add up to a generic or even subgencric distinction is open to
doubt; and in this present work Myonax has been rejected as requiring more con-
clusive data than at present available.
Turning now from generic to specific level, one of the basic problems as far so
West Africa is concerned is to determine whether the red-tail-tufted, pale-pelagcd
sanguinca and the black-tail-tufted gracilis are conspecific or not. This is but part of the
far wider problem embracing the whole vast range of described forms from other areas
of the continent, the whole matter boiling douii to whether all these small-sized
speckled mongooses belong to a single species complex spread over the whole of
tropical and southern Africa or whether vahd rpecific differences exist within a super-
ficially similar group.
hi so far as this present work is concerned two apparently distinct groups exist
difi"erentiated most obviously by the colour of the terminal pencil of the tail. These are
either identical with or related to saiigiiinca andgracilis, two species described by Riippell,
the former with a red tail from Kordofan, the latter black-tuftcd from Eritrea.
Wroughton (1907: 115-116) examined the question of specific distinction bervveen
these two and came to the conclusion that there was none; and since that time it has
been customary to sink the latter in the former, which had page priorit)' in Riippell's
work. Thomas (1917) was inclined to disagree; and in 1929 quite dcfmitely did so
when he deliberately named (^r.jc//!.!;, not saiigiiinca, as the type species of his new genus
Myonax. Wroughton had based much of his conclusion on a specimen, B.M. No.
6.10.2.9 from Erkowit near Suakin, which he regarded as intermediate between
v
310 THi; CARNIVOHI'.S OF WEST AHilCA
i^raiilis and Siim^iiiiifa, the tail tiitt being "lialt choct)latc-browii and half black". The
colour contrast is by no means so crystal clear as this description nnght lead one to
visualise, and the tuft, in flict. has little or nothing of the intense jet-black that charac-
terises the '"(jriifife" forms. Thomas thought it to be nothing more than "merely one
of the ordinary i;ni('i7i.'.' t)'pe with a more or less bleached tail-tip". Wroughton formed
his opinion on the basis, to all intents and purposes, of a single presumed aberrant,
intermediate specimen. Thomas had never seen "examples of the true Kordofan
SiiiUJiiiiuiis", that is to say topotypical material; or indeed, from the known history of the
collections, any red-tutted specimen other than the one he then had before hmi and
was in the process of naming phocnkiinis. The opinions expressed by either author
regarding the validit)' or otherwise of the two species can therefore scarcely be taken
as cariying any greater weight than tentative guesses.
The truth is that the material available is still inadequate to form the basis ot a really
sound conclusion. There are no topotypical examples ot (;ni(i7/V trom Massawa; but
there are three skins, with only two skulls, from near Suakm, some 400 km to the north
and, fide Keay ct al. (1959), in precisely the same vegetational belt. Coastal Subdesert.
It is of interest to note that these differ quite appreciably from each other in their
general pelage coloration, both above and below; and this is not merely a matter of
tone but of rechiess or greyness, demonstrating the degree of variation that may occur
even within a restricted locality'. None more than approximately resembles Riippell's
rv'pc illustration. One is that already reterred to as possessing a parti-coloured tail;
the other two are also of interest in diis connexion. Although by the standards ot
early 19th century mammalogical description all three tail tips could well be classed as
"black" none is, in flict, of quite the same pure jet-black that characterises so many
of tills genus. All have in some degree something of an exceedingly dark chestnut-red,
in one case so intense as to pass for black except in critical comparison widi the real
thing. The skulls have almost precisely the same mean measurements as saw^iiiucd
from Darfur and are otherwise indistinguishable except for a rather narrower post-
dental palate.
The present study material is far too meagre to support any tirm decision regarding
the taxonomic significance of red tails and black tails; but since there is no morpho-
logical distinction between specimens possessing the one or the other Wroughton's
view that qnuilis is synonymous with Siiii'i^iiiiicd is adopted in this work. To what
degree the vegetational background plays a part is not clear. Red tail tips are unknown
from moister forest habitats and in West Africa occur from the Subdesert to the
Doka zone. On the other hand, in West Africa the black-tipped forms are foiuid in the
forest belt except for Ciuui which came from Cape Verde (Guinea woodland). Extra-
limitally, black tailed Gahrclhi are commonly inhabitants of drier vegetational types.
The position of Gii/i/rl/if, therefore, adopted in this present work is:
Gdlcrclla santiuinca saiiquima (Riippell), iS}S- Extralimital.
,, ,, );it7(!/;/(rii (Martin), 1836. Forest.
,, ,, caiui (Wroughton), 1907. Guinea woodland.
„ ,, pliociiicunis (Thomas), 1912. Doka woodland.
,, ,, sahai'dc (Thomas), 1925. Subdesert.
GALERELLA 311
There is no present evidence that G. s. iiiiistcla Schwarz, 1935, from lower Cameroun
occurs within the Hniits chosen for this work; but it might eventually turn up from the
Cross River area.
Distribution and general. The genus is spread over nearly the whole of Africa
south of the Sahara except for the extreme south-west. Li the territory dealt with in
this present work specimens are known from the extreme west at Cape Verde, Sierra
Leone, Liberia, Ivory Coast, Ghana, western and northern Nigeria, and Air. There
does not appear to be any preserved specimen or published record of the species from
eastern Nigeria and that portion of Cameroun that lies north of the Sanaga River;
but, in so fir as the latter is concerned, Gerald Durrcll (personal commmiication)
obtained specimens at Bafut and Wum, roughly 20 and 50 km north of Bamenda.
Over much of its range the genus can be said to be common; and this is true of the
West African high forest form, mclanura, but not, apparently, of the other races within
the region. These are all distinctive Httle creatures that could scarcely be confused with
any other mongoose or small mammal occurring in West Africa, except possibly,
at a distance, a ground-squirrel.
Description. The GalcreUn mongooses (Plate 9) are all small, very slenderly built
animals, weighing from 0-5 to 0-75 kg, with fmely speckled pelage, sharp faces and
longish tails. Head & body length is of the order of 300 mm, more or less. Field
measurements are notoriously undependable, and in one relevant case, the type of
Siilumic, those for the head & body and the tail appear from the dried skin to have been
reversed by a usually extremely reliable collector (Buchanan). Considerable dis-
crepancy arises in long-haired genera from whether the tail measurement has been
made to the last joint or to the tips of the hairs; but so far as can be gathered from
label data the tail in GiilcrclLi is for the most part somewhat shorter than the head &
body, reaching some 85 to 95 per cent; though in a few cases it seems to be genuinely
longer.
The pelage consists of long, very fme, dense or fairly dense underfur and abmidant
annulated, flat-sectioned bristle-hairs that do not so markedly exceed this in length
as they do in many other species. The whole texture of the coat is fme, silky and sleek;
and in this it may be said to differ from all other West Africa mongooses. The luiderfur
is of variable length in all specimens, being composed of longer and shorter hairs
ranging from about 6 to 12 mm. It is curiously variable in colour from specimen to
specimen of the same race. The annulation of the bristle-hairs at first seems confusingly
different in the different forms; but on analysis a fairly consistent generic pattern
emerges. There are specific differences of ring-width and colour, and in a few cases a
complication is introduced by the division of the two pale zones by a subsidiary darkish
band; nevertheless, fundamentally the bristle-hair pattern is made up in the following
way. There is a pale basal region whose range of mean lengths is from 8 to ii| mm;
distally of this lies a narrow blackish ring mostly 2 to 5 mm wide; and this is followed
by a pale (white, yellow, orange) zone of ver)' variable width ranging from about
I J to 9J mm; fmally there is a browai or black tip of at least 2 mm but when imbroken
or unworn reaching 5 mm, and very fmely drawn out. Very occasionally this tip
engulfs the subtenninal pale zone, joining the next dark ring to form a long, wholly
312 THE CARNIVOKES OF WEST AIRICA
blackish distal halt to the hair. The total bristlc-hair length averages from 17 to 24 mm.
The overall effect is of a fuiely speckled coat. In some specimens the different colour
bands lie randomly; in others they fall to some extent m groups, giving rise to a some-
what irregular, rather narrow cross-banding, in no way comparable to the bold pattern
o( Miiii'^os imiii^io. This is by no means an invariable feature ot every specimen of the
same form; where it occurs, the speckling takes on a rather coarser appearance than the
fme pimctulation produced by colour bands that are randomly disposed.
Li many forms the imderside is imicolorous and fiirly sharply divided from the
flanks; but this is not always so, especially as regards the common West African race
tiuliiinir,!. Near each axilla there is a whorl of hair, and from here over the anterior
part of the chest the hair is directed forwards; but on the throat, though the lie ot the
hair is abnormal in not being evenly directed rearwards, exact detail of arrangement
appears to vary from form to form or specimen to specimen. Generally, here, the fur
is directed outwards from a longitudinal medial parting, but further detail in this
region is rather confused, there being sometimes a distinct gular whorl, and sometimes
part of the hair is directed forwards and meets the opposing lur of the chin in a transverse
ridge.
The head in ddcnlLi is small, the tace pointed, and the upper lip beneath the naked
rhinarium very narrow. The rounded ears are very broad but low and set well on the
sides of the head. The legs are short and speckled 111 the manner of the back; the narrow
feet either similar or pale and more or less unicolorous. Both pollex and hallux are
ver\' much reduced, the remaining digits slightly webbed basally, the claws short and
curved. The sole ot the hindtoot is hair)' in the posterior portion near the heel. The
tail though clad with long bristle-hairs is not bushy but the hairs are arranged somewhat
distichously, though this is less evident in some specimens, or forms, than others,
hi general it can be characterised as narrow, tapering from a moderate base to a tuie
tip, the terminal two or three inches being unicolorous, either jet-black or rufous.
The main portion of the tail is ot the same colour and speckling as the back on top but
sometimes more or less imicolorous, paler, below or with a rutous medial longitudinal
stripe. There is the usual herpestine subcircular pouch surrounding the anus and external
orifices of the scent glands; but no one appears to have studied these latter in detail.
The females carry two abdominal pairs of mammae.
Skull (fig. 40). Galcrcllii skulls are the smallest of the mongooses occurring m West
Africa the condylobasal length being always well under 70 mm. The long ovoid
braincase terminates posteriorly in a broad flange-like supraoccipital crest; but in the
vast majority of cases there is nothing more than a rudimentary or very slight sagittal
crest. In only one of some fifry skulls examined, a mediumly-aged male, could this
crest be characterised as well-developed into an erect flange of appreciable depth.
The small extreme posterior section adjacent to the supraoccipital crest is nearly
always present except in very young specimens. The relationship ot the measurements
of the interorbital breadth and the postorbital constriction is variable; the former is
roughly constant at about 11 mm, more or less; but the latter ranges from less than
10 nnn to nearly 14 mm even in the same group of approximately comparable ages,
and the ratio of the former to the latter trom 77 to 120 per cent. 13ut whatever its
GALERELLA
313
Fig. 40. Calerella saiiguitica mclamira: skull, B.M. No. 35-I-30.53. 3, x I
314 THE CARNIVORES OF WEST ATRICA
relative size one of tlie cliiet features ot the Giilcnlln skull distinguishing it from
Hcrpcstcs and Xciio<^iilc is the taet that this postorbital constriction lies immediately
behind or even a little forward of the back margin of the circumorbital ring, not
several millimetres posterior to it as in the others (tigs. 40, 36 and 44). In nearly every
adult skull the postorbital processes unite, or almost unite, with the jugal processes
to form ossified complete circumorbital rings. The frontal region is broad and rounded;
the rostrum short and blunt. Fusion and complete disappearance of the nasal sutures
takes place early. The zygomata are fairly strong, the zygomatic breadth about average,
that is to say rouglily half the condylobasal length. The postdcntal palate is ratlicr
narrower than it is long, sometimes more so than others, and occasionally markedly
Fig. 41. GalcrcUa saii^^ititwa iiwhiiinyti: bulla, -' 2
vvaisted. The bullae in this genus arc unique amongst West African mongooses in
that the anterior chamber is much more inflated than m the other genera and begins to
be comparable in size to the posterior one, though less globular in shape (fig. 41).
The premolars are very constantly y Only three skulls of many examined had the
upper cheekteeth with the first premolar lacking from one side alone. The anterior
upper premolar is always an extremely small tooth; all the remaining cheekteeth arc,
when unworn, very sharply cusped, the dentition having the appearance of being
largely insectivorous. Nevertheless, the carnassials are relatively amongst the largest
in the subfamily, the upper ones occupying 27 per cent of the length of the toothrow,
and the lower one having a large sharp outer blade, the posterior outer cusp being
the largest and the inner posterior the smallest of the three on the anterior portion of the
tooth, in^ is a fairly large tooth, but iii- is much reduced and of less bulk than the
lingual portion of »|1. The tightly packed incisors are in an almost straight transverse
row; the canines are of moderate size.
Habits. Although mongooses of this genus are widespread and pretty common
over a good deal of die continent south of the Sahara not much has been written of the
details of their way of life. There may be some variation of habit between different
forms; but because of the confusion which has long existed in respect of siibgeneric
classification and nomenclature this is a matter which must necessarily for the present
remain obscure. The only course is to assemble on a generic basis the little that has
GALERELLA 315
been recorded, irrespective of the specific or racial names with which the observations
were originally connected. As usual, there is almost nothing in the way of field notes
from West Africa itself. Most recent accounts, in fact, all stem from a single source,
the notes assembled by Shortridge (1934) concerning the genus in southern Africa;
but Taylor (1970) has included the genus in his observations on locomotion made in
East Africa. Except for a few odd scraps of information no other accounts of habits
have been traced.
The shortage is due in part to the fact that few people appear to have kept Galcrclk
as a pet; and seemingly no one who has done so has felt the urge to describe in writing
the animal's nature and habits as revealed imdcr these conditions. Opinions, indeed,
diifcr regarding the possibility or desirability of keeping the slender mongoose as a
house companion. Some have held that like others of its kind it is readily tamed (Astley
Maberly) and makes a splendid pet keeping the home free of vermin and cockroaches;
others have found it to take much less readily to captivity than most and to be largely
untameable. Cansdale (1946), indeed, writing of the West African form mclanura,
could not understand why anyone should ever wish to keep it about the house since
it was foul-smelling and in point of fact so objectionable that it was one of the few
animals of the forest that nobody would eat. Illustrating the saying that one man's
meat is another man's poison there is a note in Hinton & Diuin (1967: 10), under the
name "Dwarf Mongoose", that according to Dr. S. Toye the hunters of Northern
Nigeria eat it despite its unpleasant odour.
GalerclLi is almost entirely dmrnal, hunting singly or in pairs at almost any hour of
the day, though possibly more in the early morning than during the heat of the after-
noon. It has been said to come out sometimes on moonlight nights. As it runs, not
trots, (Taylor, 1970) along in search of prey it holds its tail out behind with the terminal
half curved upwards; and since its size and sometimes its coloration are not dissimilar
it may, especially when scurrying across a road, be mistaken for a ground squirrel,
the nature and carriage of the tail, however, helping to distinguish it. It is widely
agreed that this mongoose differs sharply from others in the relative facility with
which it can climb trees, and even descend head-foremost, though it lacks the expert
ability of the cats and squirrels. Its short, curved claws and light weight help it in this
activity. Taylor (1970), however, while fmding that this species was an able climber in
as much as it would rush at wire or other rough surfaces and clamber up, considered
that it was not really a controlled cHmbcr m that it generally fell down. There seems
little doubt that the slender mongoose climbs not only to escape pursuit but also to
seek part of its food in the shape of nesting birds and arboreal mammals. Taylor also
observed that this mongoose frequently rose to its hindfeet; and from this erect standing
position could, if it wished to reach something above it, jump vertically as much as
50 cm.
Like other mongooses it consumes a large number of msects. Ansell (1965) records
taking a specimen near a grass fire where it was busily eating fleemg grasshoppers;
but when the stomach contents were examined they were found to contain, besides
insects, the remains of a squirrel. Apart from this one example the question of food
materials has not been very carefully investigated; mongooses of tliis genus are credited
3l6 THE CARNIVORES OF WEST AFRICA
with the usual range of prey — insects and their larvae, birds, eggs, rodents, lizards and
snakes. As regards the last, Astley-Maberly (1959) wrote ot a southern African represen-
tative of the genus that it will kill even large cobras, and Gerald Diirrcll (personal
coinimmication) found that they would readily eat snakes so long as these were not
too large. Slender mongooses are also said to be poultry thieves if they get the chance;
and Cansdale (1946) relates, in relation to the forest form iitcldiiiiiii, an interesting Ghana
version of the folk legend usually associated in other parts of Africa with Atihix.
This is that GiilcrcUa lies on the gromid displaying its open anal scent gland pouch,
thus arousing the curiosity of some nearby fowl, which, supposing this pinkish structure
to be something edible, pecks at it, chokes and gets caught and killed. As with other
carnivores, wild fruits are said to constitute part ot the dietary; and Gdhrclla has been
accused of digging up groiuidnuts in fn'ms.
There is little uiformation relating to breeding habits. The shelter, for raising the
yoimg as much as for regular nightly use, is said to be made in hollow trees, in holes
m logs or in the groimd amongst tree roots, or in crevices between boulders in rocky
coiuitn'. The number born in a litter has been recorded as 2 or 3, occasionally 4.
Regarding development of tlie young or parental care there is nothing on record.
One specimen of Gii/crc//.; is known to have lived in a zoo to the age of nearly 6 years
{Int. Zoo Yr. Bh., 2).
GALERELLA SANGUINEA (Riippcll) Slender Mongoose
Hcrpcslcs sangiiiiu'its Riippcll, 1836, Nciic il'irbclihiac :ii il-r Fmiim von Abyssiiiicii gt'licr{^, etc., pt 7, Sauge-
thiere: 27, pi. 8 tig. i, pi. 10 fig. 3. Type locality Kordotaii, Sudan. The Latin adjective used as the
specific name means bloody and was given in respect of tlie reddish tail tuft.
Hcrpcstcs gracilis Riippell, 1836, Nckc IVirbclthicrc zii dcr Fitiaia von Ahyssinicti gcliorii;, clc, pt. 7, Sauge-
thiere: 29, pi. 8 fig. 2, pi. 10 fig. 2. Type locality the valleys west ot Massawa, Eritrea. The specific
name is the Latin word for slender, given with rcterencc to the build of the body.
Cyniclis nielanmus Martin, 1S36, Proc. zoo]. Soc. Lond.: 56. Type locality Sierra Leone. Type in the iiritish
Museum, No. 55.12.24.229, sex unknown; skin poor and with halt ot the tail missing; skull merely
the anterior fragment of each jaw. This name is composed ot' the Greek words molas, mclanos black,
.and oura tail, referring to the colour of the tip. Valid as a r.ice.
Mungos mclanurm canus Wroughton, 1907, Aim. Mag. nat. Hist. (7) 20: 114-115. Type locality Cape
Verde, Senegal. Type in the 13ritish Museum, No. 72. 12. 12. 5, ?; skin in poorish condition, skull
with the right side and back of the hraincase missing. The Latin word conns means grey or hoary, and
refers to the overall pelage colour. Valid as a race.
Mnngos phocnicnins Thomas, 1912, Ann. Mag. nat. Hist. (8) 10: 2S0-281. Type locality Panyam, Bauchi,
Northern Nigeria. Type in the Biitish Museum, No. 12.7.9.2, .^; skin and skull both in good con-
dition except for the loss of both inner upper incisors. The specitic name was coined from the Creek
words phoinix purplish-red, and onra tail, and refers to the tail tip. Valid as a race.
Hcrpostcs phocnicnrns saharae Thomas, 1925, Aim. Mag. nat. Hist. (9) 16: 1S9. Type locality Aoudcras,
Asben, Niger, 835 metres. Type in the British Museum, No. 25. 5. 12. 12, o; skin in good condition
but a little of the end of the tail missing, skull good except for one imier upper incisor missing.
Distribution, hi view of the continued argument regarding the validity or other-
wise of riputed species it is impossible to give any clear indication of die tull distribution
of Sdiioiiiiiiit. As the species is understood in this present work, and explained in the
GALERELLA 3I7
taxonomic section on page 309, it is known in West Africa from Cape Verde to Nigeria,
and is said to occur also in at least the Bamcnda area of upper Camcroun; but, as regards
this last, in the absence of specimens it is impossible to say which form is concerned.
Racial identification is impossible also in respect of sightings in the Borgu Game Reserve
of western Nigeria reported by G. S. Child (private communication). Extralimitally,
siiuguinca must, by definition, range to the Red Sea; and Schwarz's mustchi from lower
Camerouii is certainly part of the same complex; but how far south in the continent
the species reaches is, at the moment, entirely a matter of opinion.
Description. The species conforms in all essentials to the broad general description
already given for the genus; but the races into which it is subdivided are, within this
framework, so diverse that any more detailed description must be reserved for each
of these.
Habits. The habits so far as known have been dealt with above, and there is nothing
that can be specifically added for sniii^uinca.
Skull (fig. 40). There arc no differences from the general generic characters given
above.
Taxonomy. In the broad review of the taxonomy of the genus the question of
species was necessarily gone into and there is not much on this account that can be
usefully added here. Whether there are different species in South and West Africa or
whether they are all merely local forms of one species it is at present quite impossible
to determine with any approach to finality; and it would seem that not only must
more material be forthcoming but that, also, cytological and serological studies must
be made before any fully satisfactory solution to the problem can be achieved. For
South Africa, for example, Roberts in 195 1 recognised 7 independent species, none of
which was 5cH;(j»/;/e length
7-7
Head & body
640
Tail
36s
Hindfoot
99
Ear
34
RATIOS (per cent)
Tail/head & body
57
Zygoni. br./condvlob. 1.
51
Braincase/condylob. 1.
31
Braincase/zygoni. br.
61
Paiatilar l./condylob. 1.
56
hiterorb./postorb.
141
p'^jc llfl
iS-4
XENOGALE 329
captivity for 40 niontiis, but such a tigurc, of course, bears little relationship to longevity
in the wild.
Taxonomy. The present author feels that jacksoni is specifically distinct; that eastern
Congo (i/ijn/ii.s-, on the scores of size, some aspects of coloration, and pelage length,
merits racial differentiation; and that West African specimens should therefore correctly
be Gdlcriscus nii^ripcs iiii^ripcs (Puchcran).
Genus XENOGALE J. A. Allen, 1919
Greater Long-nosed Mongooses
Xenogale }. A. Allen, 1919, J/ Maniiii. i: 26-27. Type species Xenogalc microdoii ]. A. Allen, Akenge,
Congo. This name was formed from the Greek xaios strange or foreign, and gale weasel, for a reason
undefined but which may be guessed as referring to a set of characters regarded by Allen as unusual.
This genus, as at present known, embraces a single species occurring from the Cross
River (south-eastern Nigeria) to north-eastern Congo, probably ranging throughout
all of the central forest block north of the Congo River. In view of its monospecific
nature more detail regarding distribution and all other matters relevant to the genus
will be found in the account of this species which follows later. It is, however, first
ncccssar)' to examine questions of taxonomy.
Taxonomy. The mongoose included here has been the subject of some confusion,
even amongst expert taxonomists, and is still today a matter of dispute regarding its
proper generic assignment. Xcncqiilc was erected by J. A. Allen to accommodate a
mongoose taken in eastern Congo which he considered not only to be a liitherto
unknown species, named by him microdon, but also to present such "a singular combina-
tion of characters" as to call for a new genus. In taking this step Allen overlooked or
disregarded a very similar animal from the Camcroun River which de Winton had
some years earher named Hcrpcstes tinso. There is little doubt that, as Hayman (in
Sanderson, 1940) has pointed out, the two species are synonymous. The question
therefore at once arises as to the correct generic assignment. This in itself is bound up
with the conception of the breadth o( Hcrpcstes; whether this genus can satisfactorily
cover not only the iclineumon of Egypt but a number of Asiatic species, large and
small, the widespread slender African mongooses, and this bulky animal as well.
Opinion is sharply divided. Hayman, in the place cited, thought that it was unnecessar)'
to add a new genus to literature on account of characters that were no more diverse
from typical Hcrpcsics than those of certain oriental species generally today included
therein. However, Simpson (1945), Grasse (1955) and Dekeyser (1955) all accord
X(7i0(jii/(' recognition as an independent genus.
The question is certainly a very open one. Proposing XoicH'ii/e, J. A. Allen a little
obscured the matter now under immediate discussion by drawing comparisons not
only with Hcrpcsics but with Aiihix and khnciimia as well. The differential characters
he cited in support of his new genus in contrast with Hcrpcsics alone amoiuited, exter-
nally, to a short thick tail compared with the attenuate tail of the latter species; and to
33° TUn CARNIVORES OF WEST AFRICA
more completely hairv palms .iiul soles. These, bv themselves, do not .idd up to ,iiiv-
thiiig very signiticnnt, and any valid distinction between the two gener.i must rest
chieriy upon cranial and dental characters.
As regards tiiese, Allen, admitting the relative size and general structure of the
teeth in Xiiiot^dlc to be as in Ihrpcsns [scnsii stricio), found the "braincase . . . very
dirterent in torni from the braincase o{ Ihrpcsni' — though he failed to support this
view with any very precise account of the relevant distinctions. These are, in point
of fact, largely differences of proportion; and, because it is in broadly the same size
class, they are most readily seen in comparison with H. iciinciiiiioii though they hold,
by and large, for Asiatic species too. The mideile portion of the XtiiOi^iilv skull is
relatively very short; comparing it for example with //. iiliiiciiiiioii. the zygoma,
measured from the infraorbital foramen to its posterior root, is almost exactly the same
in the two skulls though the condylobasal length of the one is some lo to 20 per cent
longer than the other. On the other hand, the front portion of the skull, from the
anterior rim of the orbit to the gnatliion, is much longer, being in XciiO{;iilc almost
one-third of the skull length but in Hcqh'ilcs only a quarter or less. The rostrum
besides being longer is broader and more inflated as well; as a rough measure of this
it will be found that its breadth just posterior to the canine bulge is in Hcrpcstcs not
much more than the external horizontal diameter of the orbital ring, whereas in
XiiiO{^(ilc it is at least one-third as much again. The dentition is heavier, too, in this latter
genus, the molars more strongly built; and the upper canines are proportionately
bigger and somewhat of the compressed, flat-sided, sharp-edged Giihrisais type,
unlike the more usual, smoothly rounded teeth o{ Hcrpcsti s.
There are several other distinctions between XiiiOi;iilc and H. iclunniiioii but since
they do not, or not so clearly, hold for Asiatic forms are not taken into account in
this discussion. But several ot these oriental forms have their own additional peculiarities
of structure, such as an enlarged anterior chamber of the bulla. It seems to the present
author tliat, taken all in all, the differences enumerated above justify Allen's separation
of A'i;!('(jii/e from Hcrpcstcs.
A second taxonomic question concerns the species. There is no doubt that Xcfio{;alc
skulls from the Cross River basin are appreciably smaller than those from the lower
Camcroun and Spanish Guinea. Thomas observed this in connexion with a specimen
from south-east Nigeria, to which he therefore gave the subspecific name iii^^criiiniis
to distingiush it from the more southerly specimens, which he looked upon as typical
iidso. However, Haymaii later (in Sanderson, 1940) drew attention to the fict that the
t}'pe skull of /;ii.s-(i did in most respects, in fict, more closely resemble the Cross River
material than it did the specimens Thomas had taken as representing typical uaso;
and, further, that the locality from wliieh it had come, Camenuin River, belonged
essentially to the West African {iii\^ciidiiiis) region rather than to the central block
south and east of the River Sanaga, whence these latter came. He therefore concluded
that iiiocridiiiis and naso were, m truth, one and the same thing.
If this were so, the more heavily biult, southerly, specimens that Thomas had mis-
takenly treated as typical uijw were left without a name; and Hayman went on to adduce
reasons for supposing that these, in fact, corresponded to the mongoose from Spanish
a
XENOGALE 331
Guinea that had been described by Cabrera first as Hcrpcstcs nliiwdoi\iri and later as
Hcrpcstcs alhiauida var. abiwdovari. He concluded, moreover, that Cabrera was in
error in this latter specific attribution and that, in the absence of proof to the contrary,
It could reasonably be taken that the animal in question was not a variety of the white-
tailed mongoose but identical with the southern form oiiiciso, to which the subspecific
name ciliiiodoi'ari became, thus, appropriate. With this assessment the present author
sees no reason to disas;rce.
Further confusion regarding this genus has been brought about by the strong external
resemblance of Xliw(;o1c tniso to Atihix p: i
XENOGALE 335
about as broad as it is long. The anterior chamber of the bulla is relatively small com-
pared with the posterior well-inflated portion. The lower margin of the auditory
meatus is a deeply incised V. The coronoid process of the mandible is tall and fairly
narrow.
The posterior outer corner of/i'* is situated near the posterior root of the maxillary
process (fig. 33b). Normally there are 4 upper and 4 lower premolars on each side,
the anterior ones being relatively very small; and this is the case in all the 5 icnown
(/. nasc skulls except for the female that lacks a sagittal crest, which has only 3 lower
premolars on one side. One extralimital, ahnodovari, male has had an extra posterior
upper molar [ni^] on each side, now missing, but to judge from the alveoli they must
have measured about 5 mm across; and another very old male from the same area
has lost p^ on each side, one alveolus having completely disappeared, the other almost,
oji is always a prett)' large tooth, about three-quarters of the bulk of the upper carnas-
sial; m"^ is at least as large as the lingual section of /»i. In the front section of ;)/i the
anterior and buccal cusps are appreciably larger than the postcro-internal one. The
upper canmes are only slightly curved; they are laterally somewhat compressed and
nearly always display some sign of a narrow cutting edge on the posterior face, less
frequently on the anterior one.
Habits. Almost nothing is known of the life and habits of the greater long-nosed
mongoose, de Winton recorded that the original type specimen lived in the London
Zoo for a year and "was at all times perfectly silent and somewhat shy, but soon
became friendly with those whom it recognised . . .". It appreciated the gift of a
sparrow or other small bird, which it quickly ate. The label on the type of »/(^'cn'(i»ii?,
on the other hand, indicates that it feeds on snails and young maize, and is reputed
to be a scavenger. Nothing is known of mating or breeding except that J. A. Allen
(1924) records 3 young in one litter; and one juvenile West African specimen was taken
at the end of May. It seems likely that this mongoose would be a largely nocturnal
species. R. W. Hayman has kindly supphcd the following information regarding
the species in north-east Congo. One specimen was foiuid in a hollow log; and a yoimg
one still with milk dentition, and which, incidentally, swarmed with many more
ticks than any other carnivore collected, was taken in a night trap near a small sandy
stream. Indeed, tracks which appeared to be those of this species were frequently seen
alongside other clear gravelly streams in the forest.
Taxonomy. Enough has already been said to make it clear that all the animals
occurring within the limits covered by this work are in all probabUity naso uaso;
but it nuist be emphasized that this is based on the assumption that the type skull of
naso is not, in fact, typical. The point camiot be proved until further collecting from the
Cameroun River, as opposed to the Cross River basin, has shown this supposition to
be well foimded; and that no long and narrow rostrummed form intermediate between
Thomas's niocrianui on the one hand and the heavy-skulled Spanish Guinea race
believed to be ahnodovtiri on the other, m truth exists. It seems as though the eastern
Congo animals correspond pretty closely to West African n. tuiso; and if this is so it
gives the race a curiously distant east-west distribution contrasting sharply with its
very hmited north-south range, bounded as it is by the River Sanaga. But, as Hayman
336
THE CAKNIV()IU;S Ol- WEST AFRICA
li.is pointed out. should tlicsc eastern .nnnials eventually be shown to differ appreciably
from the nonnnate race Allen's name tiiiiroiloii (which seems to have little etymological
justification) is available for racial use.
Tabic 21 : NuiiK-neal cLita tor Xfiioj;alf iinso
Vegetation
Number iii niean
Condvlob.is.il length
B.i!.ilar length
Palatilar length
Zygomatic breadth
Upper cheekteeth breadth
Nasals, length
Interorbit,il brcidth
Postorbit.rl constriction
Braincasc breadth
Toothrow (f — !»')
;)■* length
ml breadth
m- breadth
nil length
iiio length
Head & body
Tail
Hindfoot
Ear
RATIOS (per cent)
Tail/head & body
Zygom. br./eondylob. 1.
Braincase/condylob. I.
Braincasc/zygom. br.
Palatilar l./condylob. 1.
Interorb./postorb.
iiaio
uaso.
tiii^i'iicinus.
liilM' lltlSO.
ahnodoi'ari,
T\pe
Type
Means
means
Forest
Forest
Forest
Forest
I
I
5
5
109-7
100-S
103-7
1 12-9
101-6
92-4
95-3
104-7
59-3
.S3-3
54-7
60-6
5.V4
53-3
53-1
62-0
33-0
3::-4
32-6
35-6
27-0
27-0
26-2
26-X
21-2
20-5
20-5
24-1
16-.S
17-0
16-X
16-7
3S-I
36-6
36-1
37-4
40- (I
36-y
3S-3
40-9
9-5
9-S
9-7
10-2
9-3
9-2
9-S
lo-o
fi-2
yx
6-0
6-7
9-2
9-0
9-1
9-1
S-6
5-6
5-S<
6-t
S2S
.s34
520
5S7
3X1
361
376
3X3
103
9S
95
107
30
33
34
3»
72
fiS
72
65
49
53
51
55
35
36
35
33
71
69
6S
60
.S4
53
53
54
12(1
121
122
144
23-4
26-5
25-3
25-0
Genus LIBERIICTIS Hayiiian. 19SS
Libenan Mongoose
Lihcriiah Haynian. 19JS, Ann. .\%i. imt. His/. (13) i : 44S. Type species Libcriiclis Lulmi Haynian, Liberia.
This name is composed of the country of origin, Liberia, together with the Greek Uiis, strictly regarded^
as the name for a weasel, but in modem zoological nomenclature otten associated with a number ot
other small carnivores.
Since this is a monospecific genus about which nothing is at present known bev'ond
LIBERIICTIS 337
the existence of eight skulls all relevant generic information can be gathered from the
specific account which follows.
LIBERIICTIS KUHNI Hayman Kubi's Mongoose
Libcriiclis kuluii 1 layman, 195S, Aim. Mai;, not. Hist. (13) I: 449-452. Kpcaplay, north-east Liberia. Type
in the British Museum, No. 5S.507, probably o, adult; no skin, skull in faiily good condition apart
from being badly smoked and the loss of one upper and one lower incisor (P and I'l) and both posterior
upper molars [iifi). There were 7 paratypcs. The species is named after its discoverer. Dr. Hans-Jurg
Kuhn of Heidelberg University.
Distribntion and general. This mongoose offers several points of interest. It is,
hrstly, somewhat astonishing that despite a good deal of systematic collecting in
Liberia by a fair number of different workers from 1875 onwards it failed to come to
hght luitil 1958. Next, at the present time nobody outside a few hunters in a rather
remote area of Africa has any idea of its external appearance beyond the fact that it
must have a longer and very much narrower and sharper muzzle than any other known
mongoose.* Finally, its teeth are so very much smaller and less carnivore-hke than those
of other herpestincs that Hayman was amply justified in erecting for it a new genus.
The only known specimens, 8 skulls, all come from the closed high-forest of the
upper Cess River, north-east Liberia, at Kpcaplay and Gaplay, roughly 6°36 —
7°o8 N., 8°28 — 8° 30 W. Apart from the type m London, the other specimens are
lodged in Boim and Saarbriicken. Hayman considered that LiIk rlictis was obviously
related to Crossarchus; but there arc 4. premolars above and below instead of 3. If
Hayman's assessment of relationship is correct it is interesting to hazard a guess that,
when discovered, Libcriictis will prove to be a mongoose twice the size of the kusimanse,
with an even longer muzzle and nose, a dark, speckled, rather loose coat, and a rela-
tively short tapering tail. The teeth, which in spite of the very i:.uch larger skull are
only the same size as those of Crossarchus obsciirus and not at all sectorial, would seem
to indicate a soft, easily masticated diet, probably insects, millipedes, worms and the
like, and possibly fish.
Skull (fig. 45). Comparisons of this figure with those of other West African mon-
gooses shows at once the marked difference of form that clearly distinguishes this skull
from the rest. Even at its widest point it is rather narrower than average, the zygomatic
breadth being only some 48 per cent of the condylobasal length; and this narrowness
is emphasized by the unusually long and exceptionally slim tapering rostrum, the
whole structure having a very canine appearance. The length of the skull lying forward
of the anterior rim of the orbit is about 60 per cent of that to the rear, whereas in other
genera it is at most about 50 per cent and often much less.
Only one skull has been available for this present study, the type, its sex unknown.
This has a low sagittal crest and the usual broad supraoccipital crest. In this specimen
* Since this went to press Dr. Douane Schlitter of the Smithsonian Institute has kindly shown the
author photographs of two specimens which he recently obtained in Liberia. These appear to
accord well with the projected description given in this and the following paragraph.
33S
TIIU CARNIVORES OF WEST AFRICA
Fig. 45. Lihaiiciis kiilini: skull. Type, li.M. No. 58.507, sex ?, ■ I
the intcrorbital breadth is precisely equal to the postorbital constriction; but Hayman
(1958) c-xainined the other seven known skulls and recorded that in all of these the
postorbital constriction was very slightly the greater, the mean excess over the inter-
orbital breadth being, however, only ot the order of one niillinietre. This latter brcaddi
is appreciably greater than die width of the rostrum across the canines, a character
LIBERIICTIS
339
true, amongst the other genera, only of Crossiirclius. Hayman, as already mentioned,
in his type description foimd that Crossarclius was the only other genus presenting a
degree of similarity to Libcriictis. The West African species of that genus, ohscHnis,
is of course very much smaller; but the extraliniital C. alcxandri (Congo) does more
nearly approach the present genus in size, and m the dorsal aspect bears a fairly close
general resemblance, apart from its appreciably shorter rostrum. The ventral aspect,
however, at once reveals sharp differences in the number, size, form and disposition
of the cheekteeth, and in the shape of the palate, both interdental and postdental.
The postorbital processes, though well developed, remain well separated from the
jugal processes; the zygomatic arch is relatively narrow throughout its length. The
palate besides being narrow is longer than usual, the postdental portion being very
shghtly longer than it is broad. The anterior chamber of the bulla is considerably
smaller than the highly inflated, domcshaped posterior chamber, and has a slight
medial depression.
The cheekteeth, j^, are remarkably small for the size of the skuU. Hayman equated
them to those of Crossarclius alcxandri which, however, to the present writer seem
appreciably larger. The upper carnassial, all of the first and half of the second molars
Table 22: Numerical data for Liberiiclis kiilii.
Means
(from
Hayman,
Type
1958)
Range
Vegetation
Forest
Forest
Forest
Number in mean
I
8
8
Condylobasal length
94-2
93-6
9I-8-95-7
Basilar length
85-6
—
Palatilar length
53-8
—
—
Zygomatic breadth
45-8
45-0
42-4-47-0
Upper cheekteeth breadth
26-0-
—
Nasals, length
29-0
—
—
Interorbital breadth
17-4
17-4
I6-6-I8-0
Postorbital constriction
17-4
18-5
I7-2-I9-6
Braincase breadth
32-9
Toothrow (r — 111^^
32-5
32-7
3I-7-34-5
p* length
5-4
5-2
4-8-5-7
//|l breadth
5-8
5-7
5-0-6-0
mi length
s-s
—
—
1112 length
$-0
—
—
RATIOS (per cent)
Zygom. br./condylob. 1.
49
48
—
Braincase/condylob. 1.
35
—
—
Braincase/zygom. br.
72
—
—
Palatilar l./condylob. 1.
S7
—
—
Interorb./postorb.
100
94
—
p*lc~nfi
l6-6
15-9
—
340 THE CARNIVORFS OF WEST AFRICA
.iFc situ.iu-d torw.ird ot tlic posterior root ot the ni.ixill.irv process. The cusps arc low,
tlic c.irn.issials li.ivc little cutting ability, and the whole dentition is crushing and
probably insectivorous rather than frankly carnivorous as coninionly understood.
The premolars and molars are both short and narrow, the upper carnassial occupies
no more than about 17 per cent of the toothrow, c — ;)(-; and in terms of absolute size
is no bigger than in the verv much smaller animal Crossiucliiis chsiiiriis. On the other
hand, 111- would seem from us now vacant alveoli to be almost as laige as 111^; and ini
and 1112 are certainly subec]ual. The antero-mternal cusp is the largest of the three on
the anterior part of ;)/|, very slightly exceeding the buccal cusp. The canines are rela-
tivclv small; the upper incisors, diough ot iimre normal size, have the outer, slightly
larger, inies (/■') clearly separated from the rest and situated laterally rather tjian facing
the front. In comparison with lower jaws of almost equal length in other genera, such as
Hcrpistis. Xiiii-n^tilc or AtiLix, the rami oi Lihaiktis arc much more slenderly built and
incapable of dealing satisfactorily with any very pciwerful prey.
HYAENIDAE 341
Family HYAENIDAE Gray, 1869
Hyaenas
Distribution. This family is the smallest of the Carnivora and one of the smallest
of all the Mammaha. Hyaenas are both Asiatic and African in distribution. As far as
the former is concerned they range across Arabia, Syria, Iraq, Iran, southern Russia
and Afghanistan to the northern and a good deal of the southern part of the Lidian
peninsula. In Africa they occur from the more inland regions of the Mediterranean
countries to South Africa and South-West Africa to the latitude of about 28^8., though
at both extremes they are becoming more and more scarce and have in fact withdrawn
from a wider range which not so long ago extended from the north to the south
coasts. Vegetationally they are essentially open-woodland animals, in West Africa
from the Subdescrt to the Guinea, the different species, however, having preferences
for different zones. Normally they avoid the closed forest except as rare and transient
visitors.
Taxonomy. The living hyaenids have long been recognised as forming an indepen-
dent family, and this position is little questioned today. But the fossil record shows
them to be derived from the Vivcrridae, furnishing remains of animals that could be
regarded as representative of one family or the other.
Extinct forms apart, the animals included here are clearly separable into two divi-
sions; cither (Ellerman, Morrison-Scott & Hayman, 1953) as full famihes or (Simpson,
1945) as subfanulies. The former is probably justified but the latter 's classification is
nevertheless retained here pending a major classificatory revision of the African
mammalia, the point being of little immediate moment m West Africa since only
the chief of the two divisions occurs there. Simpson's subfamilies are the Protelinae
and the Hyacninac, both of Mivart, 1882. The former consists solely of the so-called
aardwolf (i.e. Earth-wolf), Protclcs I. Geoffroy, 1824, of southern and eastern Africa,
which though it outwardly very closely resembles a small striped hyaena has denti-
tional and other characters that relate it to the Viverridae or, m the opinion of some,
the mongooses in particular. The cheekteeth of this small, shy and weak-jawed carnivore
are j-j but are often partly missing besides being remarkably reduced in size and effec-
tiveness. Moreover, it subsists largely on an insectivorous diet. With this animal this
present work is not further concerned since only the Hyaeninae occur ui West Africa.
Nevertheless, it is mteresting to note that support for its near affinity with Hyaena
ratlier than the more distant one suggested by removal into a separate family is pro-
vided by the discovery of very closely related lice (Mallophaga) of the genus Filiccla
on both the aardwolf {Proklcs cristatus) and the brown hyaena {Hyciaia hrunnca)
(Ledger, 1968).
Subfamily HYAENINAE Mivart, 1882
Hyaenas
General description. Hyaenas proper are all animals of large or fairly large size
X
342 THE CARNIVORES OF WEST AFRICA
which, although they arc anatomically more closely allied to the Felidae, seem, in fact,
in many ways far more dog-like than cat-like in general appearance. Possibly the
most immediately noticeable peculiarity of these carnivores is the sloping hne of the
back due to a greater length of torelimbs than hmdlimbs. Indeed, the whole front
portion of the body is of appreciably heavier build than the rear, the shoulders, the
thick neck, the massive head and large ears seeming overdeveloped in comparison
with the hindquarters. Hyaenas arc m consequence of clumsy rather than of elegant
appearance. The deep jaw armed with huge teeth is extraordinarily powerful and cap-
able of dealing easily with the thickest of bones — or even with tougher materials in
need. Li accordance with predominantly nocturnal activity the eyes have a very vifcU
developed tapclum luddum wliich enables hyaenas to make full use of every glimmer of
light. The ears, which may be pointed or rounded, stand up conspicuously above the
head and have no bursa. The hmbs are canine-like, terminating in thickly padded,
broad feet which have 4 webbed digits and short, stout, blunt, non-retractilc claws.
These are in part concealed by very dense, stiff and wiry, curved bristles that cover
the whole upper surfice of the foot. The gait is higlily digitigrade, the subdigital pads
bemg both exceptionally deep and flattened on their anterior fices in accordance with
this extreme verticaHty of the digits. The tail is short and bushily long-haired, either
throughout its length or at least terminally.
The coat is fairly harsh, short-haired to very long-haired, with a crest that may be
more or less confmed to the back of the neck or extremely lengthy and run from head
to tail. The pattern is of spots or transverse stripes — m an extralimital species the latter
confmed to the legs. The pelage, when not in moult, is composed of fairly abundant
underfur that is sometimes very long and always wiry, not far removed m thickness
and texture from the bristle-hairs ot other families. Amongst this occur widely scattered,
very long, terete or slightly flattened bristle-hairs, plaving a relatively small part in
the overall texture of the coat.
The scent glands are situated either side of the rectum, their external orifices lying
deep in a hairless sac situated between the root of the tail and the anus. This pocket is
surrounded by a soft, tliickened, dilatable, naked run, the upper and lower lips of
which are parted when the tail is raised but come together when it is lowered, closing
the pouch in an arch-like slit above the anus. The latter is thus, unlike m the mongooses,
external to die sac; but the naked skin surrounding it is continuous with the naked
lips of the pouch, the whole complex forming when the tail is raised and the pouch
fully distended, a practically hairless subcaudal disc. The two external orifices of the
mam scent glands are sited on the floor of the sac a little distance to each side of its
longitudinal axis; but betv/een them he, also, the pores of a number of lesser inter-
mediate sebaceous glands.
Skull. This is of outstandingly powerful build. It is cliiefly remarkable for its high
sagittal crest that arises gradually, not abruptly, from the disproportionately narrow
cranium and is projected far backwards beyond the usual limits of a skull in the form
of a subpyramidal "helmet". The supraoccipital crest, that plays so prominent and
constant a part in most skulls of the Viverridae is in this family relatively insignificant.
HYAENINAE 343
The zygoma is very strong, but the orbital ring is widely incomplete. The auditory
meatus is bony and tubular.
The bullae arc well developed: Pocock (1916c and 1928) drew attention to the
fact that they are two-chambered despite the assertions of Flower (1869), Mivart
(1882 a and b) and subsequent authors. The error arose from the more horizontal
position of the septum, which divides an exceptionally large anterior chamber from
a small posterior one, and which had hence been mistakenly assumed to be the actual
roof of an undivided bulla. There is no external indication of the existence of two
chambers, in the form of a depression in the wall of the bulla, as there is in the Viverri-
dae. The paroccipital process is highly developed, completely enfolding the posterior
aspect of the bulla, extending well below it in the form of a blunt point, and reaching
laterally to the edge of the auditory meatus. Pocock also pointed out (i9i6f) that the
same two authors were wrong in citing the absence of an alisphenoid canal as charac-
teristic of the Hyaemdae since though it was usually absent it was sometimes present,
at all events in Crocuta. Its apparent absence is due to the obHterarion of its posterior
orifice, the anterior orifice being mistaken for the foramen rotundum, which is, in fact,
hidden witliin the alisphenoid canal, as observable by cutting away the outer wall of
the latter. The mandible is massive.
The cheek dentition is j-j, the upper molar being very much reduced by com-
parison with the huge camassials and remarkably powerfial remainder of the teeth;
this is especially so in Crocuta, where m^ is minute and sometimes lacking.
Differences of detail in the structure and proportions of the skulls of the two genera
Hyaena and Crocuta have been very fuUy described by Buckland- Wright (1969).
Habits. Hyaenas have long had an unsavoury reputation as nothing better than
cowardly scavengers, feeding on stinking carrion or, at best, slinking furtively in to
finish oft what remains of a carcass when the hons have killed and eaten their fiU.
It now appears that this picture is quite misleading, at least as far as the spotted hyaena
is concerned, as described more fully below in the accomit of that species wliich recent
observation shows to hunt and kill on its own accotmt. However, there is no doubt
that hyaenas do also scavenge, coming in to villages at night to take what offal they
can fuid. If opportunity' offers on these visits they steal sheep, goats or calves as well;
and they have been known to take children too. As for their cowardice, it is true that
they make Uttle attempt to defend themselves but both kinds have been observed to
drive off" a leopard or a young lion from a kill. Their usual careful approach may
be as much the outcome of excess of caution as of lack of courage.
These are chiefly nocturnal animals but are occasionally active during the day and
especially in the early morning. As a rule during dayhght hours they shelter and sleep in
any cover, in aardvark holes, under boulders, or in dense vegetation; but they sometimes
choose more open sites such as the shade of a tree. Though by nature not truly gregarious
animals they do often live in family groups and frequently gather together in some
numbers either at a kill or for a kill, as will be detailed later in the specific accounts.
Since they are mainly night feeders they are not commonly seen except when the moon
is full; but anyone who has been in the correct territory' is very well acquainted with
344
THE CARNIVORES OF WEST AFRICA
thcni through the hideous noise that they sometimes make throughout much of the
hours of darkness. Both striped and spotted hyaenas do this; but the latter is somewhat
louder and its risnig whoops and cackle have made it famous to all as the "laughnig
hyaena".
Little is certainly biown of their breeding habits. The maniacal laughter is thought
to be made luider the stinudus of sexual excitement. The females are capable of breeding
at all seasons of the year. The period of gestation has been estimated as between 3 and
}k montlis; there arc mostly 2 to 4 in a litter. The young of the two genera arc quite
different: those of Hyiiciui resembling their parents in their markings; those o( Crociita
displaying no pattern at all, merely a blackish-brown colour.
Taxonomy. No argiuncnt attaches to the separate recognition of this subfimily and
that of the Aardwolf (Protelinac) apart from whether they are each of family or sub-
family status. Within the subfimily itself, it was at one time customary to relate crocuta
to hyihiui as species within a single genus HyMua; but there can be no doubt of their
true generic independence, the nature of the teeth, the pelage and the sexual organs
being very distinct.
KEY TO THE GENERA OF HYAENINAE
(previous key page 160)
Coat pattern of transverse stripes; a lengthy crest from head to tail; ears pointed;
upper molar much more than 5 mm in transverse length . Hyaena [page 344)
Coat pattern of spots; if any crest present, relatively short and confined to the neck ;
ears rounded; upper molar very small, only about 5 or 6 mm across
Crocuta [page 353)
Genus HYAENA Brisson, 1762
Striped and Brown Hyaenas
Hyaena Brisson, 1762, Rft;iiuin Animalc in classes IX dislrihulnm. 2nd. edit.: 13 and 168. Type species
Canis liyaena Linnaeus from Iran. This name is the Latin torni of tlie Greek hyaina, itself reputedly
derived from hys a liog, and -aina a feminine termination, and said to have been given in reference
to the long spinal crest similar to that occurring, less exaggeratedly, in the European wild boar. The
nomcnciatural validity of Brisson's work has been questioned, the matter having been put before the
International Commission in i()io (Bull, zool Nom. 4: 314) hut no decision liaving yet been reached.
Should this work be rejected the name Hyaena is still available from Briinnich, 1771, see below.
Hyaena Briinnich, 1771, ZiU'/oijifli' Fundamenia: 34, 42 .and 43. Type species Cain's hyaena Linnaeus.
Distribution. This is the more widespread of the two hyaenid genera, being found
over a good deal of two continents. It has been known for very many centuries from
its chief species hyaena. This today occurs throughout much of the western half ot the
Lidian peninsula, nordiwards as far as southern Russia, and thence across Iran, Iraq,
Syria and the whole of the Arabian peninsula into Africa. In Africa the distribution is
in two disjoined parts according to the two extant species. The striped hyaena, hyaaia,
which is also the Asiatic species, is to be found all around the Sahara, north as well as
south, and as fir east as the Red Sea and soniewh.it south of the Equator; the brown
HYAENA 345
hyaena, hruunca, is confuicd, broadly speaking, to South-West Africa and a few scattered
locahties further cast to Mozambique. In former eras the genus occurred in Europe.
General description. Hyaenas of this genus arc externally characterized by much
longer pelage than that found in Croaiia; and by a pattern of transverse blackish stripes,
either on the flanks or at least, extralimitally, on the legs. The ears are pointed and pear-
shaped, markedly different from those of the other genus; and the general bodily size
is less. Further detailed description is reserved for the sole West African species, hyaena,
below.
Taxonomy. There is little to add under the generic head to what has already been
said. Clearly, in spite of former practice, it is correct to separate the spotted hyaena,
Crocuta, genetically from those now under consideration. It is equally clear that the
genus covers two distinct, extant, species : hyaena of Asia and the northern part of
Africa; and hrunnca, with a different pelage pattern and a slightly larger skull and teeth,
of the south, a specific distinction that has never been brought into question.
HYAENA HYAENA (Linnaeus) Striped Hyaena
Canis hyaena Linnaeus, 1758, Systana Naturae, loth edit. 1 : 40. Type locality Benna Mountains, Laristan,
southern Iran(/i(/t' Thomas, 1911, and Pocock, 1934: 809). Linnaeus' type is thought to have been
the description, pp. 411-412, and plate (fig. 4) facing p. 407 in Kaempfer (1712) Anwenilalwii Exoti-
cartim . . . Fasc. V. . . . Rerum Persiaim et Ultcrioris Asiae. The derivation of the name has been given
above under the genus.
Hyaena striata Zimmermann, 1777, Specimen zoologiac geographicae etc.: 366. A renaming of hyaetia
Linnaeus. Zimmerm,inn's work was ruled unavailable in 1950 {Bull. zool. Nom. 4: 547). The name
striata is a Latin word implying marked with narrow bands.
Hyaena diibbah Meyer, 1793, Systcmatisch-sunnnarische Uchersicht ier nenesteii zoohgisclien Entdcckungen
in Neu Holland und Afrika: 94. Atbara, Sudan. This was entirely founded on the description by Bruce, J.,
1790, Travels to discover the source of the Nile ... 5: 107-120, pi. Dubbah is the Arab name. Sometimes
held to be of racial application in West Africa.
Hyaena orientalis Tiedemann, 1808, Zoologie I: 350. A renaming of hyaena Linnaeus. The name is the
Latin word for pertaining to the east.
Hyaena Jasciata Thunberg, 1820, K. svenska VetenskAkad. Haiidl. I: 59. A renaming of hyaena Lirmaeus,
This name is the Latin adjective meaning enveloped with bands.
Hyaena antiijiiorum Temminck, 1820, Annals, gen. Sci. phys. Briix. 3: 51. A renaming of hyaena Linnaeus,
this being the Latin word meaning of the ancients, since this was the only species known to the early
classical writers.
There is a further fairly extensive list of synonymous names either exclusively applied to the species
outside Afiica or those of extralimital races.
Distribution. The striped hyaena has been known and been the subject of a good deal
of far-fetched misinformation throughout south-western Asia and the Mediterranean
region since ancient times. Lr Africa it occiurs today all around the edges of the Sahara,
and thence it ranges as far east as the Red Sea and, on that side of the continent, south-
wards to about 4^5. Within the desert itself it has to all intents and purposes disappeared,
and it has become fairly rare aroiuid its northern borders, that is to say in the interior of
Morocco and Algeria, though it is still not imcommon in southern Libya. It is essentially
an animal of the arid zones, the Subdesert and the Sahel woodland, further south than
346 THE CARNIVORES OF WEST AFRICA
which it occurs oiilv sporadically as an occasional visitor. It can be expected anywhere
within these two zones especially the latter far-spread belt o{ AcMia radiiiana woodland,
which extends in West Africa from Senegal to Lake Chad and thence across the con-
tinent eastwards to Somalia, turning south into parts of Kenya and Tanzania. Reputed
occurrences outside tliis normal range in moister zones of vegetation are mentioned in
the following paragraph.
bi the right areas the striped hyaena is moderately common but is nowhere so
abimdant as the spotted hyaena may be in its optimum habitat. It is impossible to judge
its numbers in West Africa from the British Museum material since this comprises only
three specimens: one a skin complete with skull, but juvenile, from Manakaoki in Air;
and two skins without skulls, one of them incomplete, in full moult, and almost
certainly purchased front local hiuitcrs, the other from the London Zoo, where the
animal had hved for 19 months, and so is probably not altogether typical. Both these
latter skins come from Nigeria, the Zoo specimen without a locality, the incomplete
one from Gwoza (ii°05 N, I3"42 E), possibly in the Sudan zone but very near a tongue
of Sahel. Two reasons probably contribute to this poverty of material: firstly, that
Hyaena being almost purely a night animal is therefore rarely shot; and secondly, the
arid zones of West Africa being far less stocked with game and with large game-killmg
carnivores such as the hon and the cheetah than other parts of the continent it is likely
that the hyaenas arc in consequence correspondingly few. Nor are there many reliable
up-to-date records of observed occurrence. The species reputedly occurs as a rarity m
the extreme north of Sierra Leone according to Stanley (in Goddard, 1925) ; and T. S.
Jones (personal commiuiicarion) heard of one shot by a Veterinary Guard near the
northern border in 1954 which from a description must have been hyaena. A.J. Hopson,
in a private commimication, says the striped hyaena is to be seen occasionally on flats m
the Yobe valley between Yo and Lake Chad; and another trustworthy observer reported
a few years ago seemingly the most unusual of all occurrences when an African hiuiter
shot a specimen near Nabardo, about 75 km from Jos on the Bauchi road (Nigeria),
that is to say in Doka woodland some 400 km south of the normal range. Zimara
(1935) recorded a specimen front Fort Archambault in the Sudan woodland.
Description. The striped hyaena (fig. 46) is the smaller of the two West African
species, standing about 720 mm at the shoulder, or somewhat less, and weighing 35 to
45 kg. The sloping back, from high shoulders to low hindquarters, is very noticeable.
The overall impression is that of a shaggy animal due to the generally long hair of the
pelage accentuated by the extremely long and abimdant crest running from head to tail.
Tltis crest, and indeed the whole pelage, is erectile. The colour of the coat varies between
individuals and from region to region; and in the face of die pauciry of specimens it is
impossible to be very precise regarding its appearance in West Africa. In general the
background colour ranges between pale grey and yellowish-buft, and on this, along
each flank ber^veen the belly and the crest, are imposed a number (c 6 to 8) of deep
brown or blackish transverse stripes of very irregular length, breaddi and contimiiry.
The legs, too, especially the upper portions, bear similar dark transverse bands. For the
most part the body stripes are some 10 to 15 mm broad at their widest, but usually taper
at each end. They are formed by bristle-hairs that instead of being unicolorous as in the
347
Fig. 46. Striped Hyaena [Hyaena hyaaia)
Spotted Hyaena {Croaita croaila)
348 THE CARNIVORES OF WEST AFRICA
rest of the coat arc blackened tliroughout all or a great part of their length. The spinal
crest is composed of dense bristle-hau's that may reach well over 200 mm ni length and
are also blackened, though so much ot the long pale basal portions shows as well that
the general effect is parti-coloured black and buff. The throat, and sometimes a little of
the chest, is black, this colour extending here and there to the sides of the neck.
The composition of the pelage is normally of moderately long underfiir and much
longer bristle-hairs. The lengths vary a good deal from specimen to specimen but an
average is 25 to 30 mm for the underfur and 50 to 60 mm, sometimes more, for the
bristle-hairs. In some cases, through moult, die underfur entirely disappears leaving a
thin coat of bristle-hairs inadequate to conceal the skin. The tail is short, that is to say
about a quarter or a little more of the head & body length, and very bushy with
extremely long, coarse bristle-hairs similar to those of the crest, with which it is in a
manner of speaking continuous. The long ears are pointed, and hence m sharp contrast
to those of Crea/fd. Another marked difference between the genera lies in the external
sexual organs of the female, which in Hyiuihi are normal and do not bear that superficial
resemblance to those of a male that has given rise in the other genus to stories of
hermaphroditism. The supra-anal scent pouch has already been described above in the
general account of the Hyaenidae. The female has three pairs of abdominal mammae.
Skiill (tigs. 47 and 48). This has been broadly described in the introductory accoimt of
the 1 lyaenidae, above. It differs from that ofCnu'tila partly in its appreciably smaller size
and generally slightly less massive build. It is none the less a very powerful structure well
adapted to the anchorage of an exceptionally strong muscular system. The posterior
projection of the sagittal crest is much more roimded than in Crocutd. The most positive
distinction between Hyaena and Crocnta, however, lies in the dentition. This, though m
general smaller, differs in having a far larger upper molar which is by no means the
smallest tooth in the head, its measurement transversely across the palate being of the
order of 15 mm.
Habits. Because of its distribution, in the African Mediterranean countries, through
Arabia to India, the very axis of the ancient world, the stnped hyaena has been known,
talked and written about for countless centuries, the source of horrific stories, wild
beliefs and misleading information. Topsell (i6sS) in his indefatigable fashion compiled
five or six closely printed pages crammed with tabuloirs beliefs and outrageous medicinal
recipes gathered from Aristotle, Pliny, Galen, Albertus and a dozen odier ancient
authorities, information once a matter of most earnest acceptance though today almost
wholly risible. Nor are these beliefs yet dead; for in Libya (Zammarano, 1930: 21) the
local inhabitants still speak of this hyaena as of a legendary animal and attribute to it
magical powers and fabulous medicinal properties. Harrison (196S) mentions a some-
what similar situation 111 Arabia. Despite a lengthy history of close association with
man the plain fict is that by modern standards our knowledge of the way of life of this
nocturnal and secretive creature remains extremely sketchy, still often amounting to
little more than matters of hears,ay since no set field study of the bionomics oi Hyaaia
has yet been luidertaken as it has for Crocnta.
Little that is up-to-date and supported by good evidence has come out of Africa
itself. Few competent observers have written of the inhospitably arid zones in which
HYAENA 349
this hyaena hvcs; and those that have mostly cither failed to mention the animal or
limited themselves to remarks regarding repulsive feeding habits and nocturnal howling.
The behaviour oHiydi lui has always been tacitly presumed to be in every way precisely
that of the rather better known crccuta; but while this may be so as regards the gross
pattern it is in so far as the finer details of existence arc concerned httle more than an
unconfirmed assumption. Even if we turn to hrdia for information the results arc still
disappointingly meagre.
The striped hyaena is a more strictly nocturnal animal than the spotted hyaena, rarely
emerging from its diurnal shelter before nightfall and returning to rest before daylight
or soon after dawn. It is thus hardly ever seen except by moonlight; and this in part
accounts for ignorance of the fmer points of its habits and conduct. Further, unlike
Croaita, this hyaena does not as a rule join up with others into hunting or feeding
parties but remains solitary or in pairs. Like the spotted species it is said to travel long
distances in search of food. This last consists largely of carrion or, at least, the remains of
carcases after other predators have fmished with them. Almost nothing comes amiss,
though Hilzheimer (1915) records that the flesh of vultures is scorned even by hungry
hyaenas. Bones of all kinds and sizes are eagerly eaten, even those already denuded of
all flesh by vultures. There is no difficulty regarding the breaking up of even those of
large size. Nor is there about digestion; bones or the partly digested calcareous remains
of them, in fact, form a high proportion of the droppings, which are very characteristic
since they quickly dry to hard, pure white, subglobular masses.
Striped hyaenas are known to attack farm stock. The victims are commonly of the
smaller or younger kind, sheep, goats and calves ; but successful attacks arc sometimes
made on much larger animals up to the size of a donkey or a horse. The young of
antelopes, or sick adults unable to defend themselves also fall to these hyaenas as occasion
offers. Normally these predators ravenously and gluttonously get on with stufluig them-
selves with food as soon as they can safely get near the meal, and continue luireniitringly
until they are satisfied — sometimes consuming enormous quantities. At times, however,
they carry food back to their dens, but whether this is as a further meal or for the
purpose of feeding the yoiuig is not clear. Hilzhcimcr (1915) records that not infre-
quently a pair of tame hyaenas was, through lack of supplies, forced to go without
eating for three or four days, and on one occasion eight days, without trouble or ill
effect apart from eventual tremendous hunger. Something similar to this might well be
a pattern in the wild. Flesh is without doubt their favourite meal; but in captivity they
will eat other things, including bread (Hilzheimer, 1915) and fruit, for which latter
Flower (1932) records an extreme partiality. Because they so readily scavenge anything
available they can be easily trapped or poisoned. One of their more horrific reputations
is the digging up of corpses. This has been positively asserted over and over again but
occasionally writers have denied its truth. Both opinions probably have some justifica-
tion: the story is almost certainly true in relation to hurriedly buried bodies lightly
covered with sand; but seems less likely to be so of properly deeply interred corpses.
Striped hyaenas rest during the day in holes in the ground, such as porcupine or
aardvark burrows duly enlarged when they are to form permanent breeding homes; or
in cavities amongst rocks, or sometimes fairly narrow crevices. In so far as the last are
350
THE CARNIVORES Or WEST AFRICA
conCL-riu-d Zamniarano (1930) relates a strange story of the bolder luinters of Libya
worming tlicniselves into the depths of such shelters m order to bind and drag out the
occupant on a long rope; such performance being not ahogcther so dangerous as it
might at first seem since the hyaena retreating head-first into the narrowest part of the
crevice is unable to turn round to confront and bite its aggressor. Confirmation of this
is given by Salez (1954) who states that in Algeria it is claimed diat certain Arabs, going
quite naked into a hyaena burrow, are easily able to capture the animal alive.
Fig. 47. Hyuciiii hyaciui: skull, B.M. No. 23.1. 1.78, j.
later
Breeding, also, takes place in these shelters. The period of gestation, according to
observations quoted by Hilzheimcr (1915) for the Leipzig and 13erlin zoos, is probably
90 or 91 days, not 7 months as mentioned in Pocock (1941) which would seem to he far
too long. Litter size is said to vary between 2 and 4, but observations are few and un-
reliable in spite of the fact that the species breeds readily in captivity (Crandall, 1964).
The young differ from those of Crocuta in bearing a recognizable resemblance to the
adult pelage pattern; the hair is short, silky and white but the transverse stripes on body
and legs are clearly discernible; and the eventual spinal crest is indicated by a dark band.
According to Pocock (1941) the newly horn cubs have their eyes closed and their ears
sealed down. If this is so it is in very marked contrast to Cmciilii in which the young
are born well developed and almost immediately active. Nothing is known of the course
of development or of parental care; but as fir as one aspect of the latter is concerned it
is possible that the young are fed, after weaning, by regurgitation. At the other end of
the scale, it is recorded that one of these hyaenas lived in captivity to an age of about
HYAENA
351
Fig. 48. Hyaena hyaena: skull, B.M. No. 23.1. 1.78, ,5, X i; palatal & dorsal views
24 years (S. S. Flower, 193 1), still tame and friendly.
Strange as it may seem, these hyaenas can be fully trained. The yomiger the start is
made the better; but Hilzheimer (191 5) gives a long account of the domestication of a
352 THI- CARNIVORES OF WEST AFRICA
pair, still juvenile but socniingly past their babyhood. These were difficult at first until
complete domination had been established by strong-arm methods; but they eventually
came to conduct themselves like well-trained dogs and even exhibited obvious affection.
It is interesting to note that this last was displayed in its highest form, as an excited
welcoming, by the opening and eversion of the supra-anal pouch, the tail being held
stiffly erect. However, no mention is made by this author of any objectionable odour
trom these glands, then or at other times; but a curious use of the scent for criminal
purposes is recorded by Salez (1954). This author relates that the odour of the striped
hyaena is so feared by dogs that they freeze with terror and flee without uttering any
sound; and this effect is made use of in parts of Algeria by bandits who, furnished with
a hyaena skin or themselves anointed with the scent, can thus without raising an alarm
from the watchdogs enter settlements and drive off^herds, later to be restored in return
for the payment of ransom. Whether the odour of striped hyaenas really does have such
a terrifying effect on dogs is imconfirmed, though Bruce (1790) discovered that his
normally brave greyhounds refused to hunt them. On the other side of the picture,
both Bruce and Hilzheimer relate that these hyaenas are extremely keen on the flesh of
dogs and will display considerable audacity in carrying them offl Pocock (1941), how-
ever, mentions hyaenas m India shamming dead when attacked by dogs; and also a case
of a tamed hyaena living in perfect amity with several dogs. The vocalizations of this
species of hyaena are somewhat similar to those o(Cnuiila but less noisy.
Taxonomy. A good many races of the striped hyaena have from time to time been
described, five being currently recognized. 3 in Asia, 2 in Africa. None has ever been
named for the area dealt with in this present account. Authors have depended for
subspecific distinction very largely upon colour, with or without some other attribute
of the coat, and occasionally on the size of the teeth especially the upper carnassial, p'*.
Pocock (1934) went into the question of the races o( Hyiumi in great detail, carefully
examining the evidence for all the named forms. He came to the conclusion, at least in
so fir as tropical Africa was concerned, that colour could not be regarded as a basis for
subspecific separation, many of the distinctions claimed as ot taxonomic importance
being merely seasonal or individual. Re-examination of the British Museum specimens
confirms this view. The plain fact is that the material is too meagre and of too scattered
origin to form a pre^perly sound basis of argument; but there can be very little doubt
that in any given locality there is an appreciable individual variation, of coat colour and
marking, apart from any differences attributable to age, moult or season. It must not be
overlooked, too, that these hyaenas are great wanderers and travel many miles in search
of food. Those fortuitously gathered together m any one place at a kill may, in fact, have
come from appreciably different backgrounds; and the existence of any strictly "local '
race is open to doubt, the more so as the wandering habit renders breeding contacts
very wide.
Pocock reached the conclusion that two races could be recognized in Africa: harhara
Blainville confmed to north-west Africa, and dnhhah Meyer covering all those forms
named for the eastern side of the continent. The latter was held to be characterized by a
smaller skull and teeth as indicated by />''; but the evidence for this seems pretty slender,
and Pocock himself admitted a link between the two races afforded by specimens from
CROCUTA 353
Uarfur. Since the publication of his paper it has become customary to assume that West
African hyaenas arc also assignable to diibhah. Bruce's description of this form [i.e. the
original English of the German translation used by Meyer) is of a vast animal with a
head & body length of 1750 mm, a tail of 530 mm to the end of the hairs, and weighing
some 50 kg — as Pocock remarked, an exceptional giant amongst striped hyaenas. The
pelage colour was given as yellowish-brown, the head and cars hghter; both the tail
and the spinal crest were of a reddish-browar colour "without any rings or bands of
blackness upon the points". Certainly nothing like this is so far recorded from West
Africa, the only known skin closely corresponding to this colouring coming from
Uganda. Pocock himself made no attempt to assign the only West African specimen
then to hand, an immature skin and skull from Manakaoki (Air), either to his widely
variable diihhah or to any other form. Since his detailed review further West African
material has become available but as this comprises only two skins unsupported by
skulls, the one a zoo animal reputedly from Nigeria, the other incomplete and in full
moult without underfur from Gwoza (Nigeria), it is quite insufficient as the foundation
of any reasonable judgment. The first two are densely haired and basically buff; the last
almost hairless, what little there is of the thin top-coat being white.
The present author sees no virtue in pretending to a degree of accuracy implied by the
attachment of a third name, either diibbdh or anything else, to West African specimens
on the basis of the present available material.
Genus CROCUTA Kaup, 1828
Spotted Hyaenas
Crociita Kaup, 1828, in Oken's Isis, 21: column 1145. Type species Caiiis crocnta Erxleben. This name was
really preoccupied by Crocuta Meigen given to a two-winged fly in a paper published in i Soo. Although
Meigen's pamphlet was declared available in 1944 by Opinion No. 152 of the International Com-
mission on Zoological Nomenclature it was, in 1962, suppressed by the same body under Opinion
678. The name is derived from the Greek krokolos meaning saffron-coloured, with reference to the
pelage.
Crocotta Kaup, 1829, Skizzirte Eiitu'ickclmi(;s-Geschiclili: imJ nalurliclws System dcr Europiiischen Thier-
u'clt ... I; 78. Type species Canis croaita Erxleben. A rc-spelling of the above.
This genus differs from the last in being purely African in its present distribution; but
in late Tertiary times it was an inhabitant of Asia and Europe, at least as far north as
England, where caves are known containmg abundant remains. Though, in its heavy
forequarters and sloping back, Crocuta bears a broad resemblance in bodily form to
Hydaid, the general pelage pattern and character are quite different, consisting of spots,
not stripes, and being much shorter in length, lacking any marked spinal crest. The
rounded ears are also quite distinct; and there is an important difference in the dentition.
Since the genus is tnonospecific there is no need for further generic description.
CROCUTA CROCUTA Erxleben Spotted Hyaena
Canis crocuta Erxleben, 1777, Systema Regiii Animalis . . . Classis I, Mammalia: 578. In this work the range
was given as Guinea, Ethiopia, to the Cape of Good Hope; but Cabrera, 1911, in Proc. zool. Soc.
3 54 THE CARNIVORES OF WEST AERICA
Loud. : 95 argued that the type locality was most probably Senegamhia. The derivation otthis name has
been given above under the genus.
Hyaena maculata Thunberg, 1811, Mini. Acad. Sci. St. Pcicrsh. 3: 302. Type locality South Africa. The
name is the Latin adjective meaning spotted.
Hyaena capensis Desmarest, 1817, Nouvcaii Dictionnaire d'Histoirc Nalnrclle etc., Nouvelle edition, 15:
499. Type locality Cape of Good Hope.
Hyaena rufa Desmarest, 18 17, Noiweaii Dicticnnaire d'Histoirc Namrclle etc., Nouvelle edition, 15: 499.
Type locality Cape of Good Hope. The name ra/a is the Latin tor reddish, referring to the pelage
coloration.
Hyaena croacuta A. Smith, 1826, A Descriptive Cataloi^iie of the Sotilli African Mitsetiin, Part I, of Mammalia:
12. A misspelling o( crocitta.
Hyaena eiicrila A. Smith, 1827, Trans. Linn. Soc. Lond. 15: 461. A misprint for crocuta.
Hyaena ciwieri Boitard, 1842, Le Jardin des Planles: 233. Type locality Cape of Good Hope. This was
named m compliment to the great naturalist Baron Cuvier.
Hyaena {Crocotta) thierryi Matschie, 190D, Shcr. Ces. natiirj. Freniuie Bert.: 30. Type locality Sansanne
Mangu, Togo. This was called after its collector Oberleumant Thierry.
Hyaena (Crocotta) to^oensis Matschie, 1900, Sber. Ges. naliirf. Frciinde Berl: 31. Type locality Ketc Krachi,
Togo.
Hyaena [Crocotta) tioltei Matschie, 1900, Sber. Ges. natiirf Freniuie Berl.: 215. Type locality Yoko, upper
Sanaga, Cameroun. Named after Ohcrleutiiant Noltc, who collected it.
Distribution and general. It may be broadly said that the spotted hyaena is spread
through Africa south of the Sahara with the exception of the closed forest. It is true
that occasional occurrences in this last zone are reported but these mostly refer to places
that are not far distant from the open grass-woodlands or at least from clear-felled
farmlands; and, in West Africa at any rate they are very exceptional. One such visit is
mentioned later in this account. On the western side of the continent Crocuta occurs in
Senegal but apparently not so far north as Rio de Oro. From this extremity it ranges
across the continent in the Subdesert zone (but not the desert itself) Sahel, Sudan, Doka
and Guinea woodlands to Eritrea and Somalia; thence down the eastern half to parts of
South Africa, South-west Africa and Angola. It is known to ascend to an altitude of
about 3500 metres. Its distribution, thus, m western Africa m parts overlaps that of the
striped hyaena, chiefly in the Sahel zone. Although it occurs outside them the spotted
hyaena is most at home in the Sudan and Sahel woodlands, though rarer in the latter
than the former. Its appearance further south in the Doka and Guinea zones is to a large
extent a matter of season; for m these, especially the latter, it is impossible to see and not
easy to progress through the tall dense grass that normally covers them. When this is
burnt in the dry-season, however, large stretches of the coimtry become wide open, the
newly sprouting green grass attracts antelopes, and the zones become for a few weeks a
profitable hunting groiuid. In parts of eastern and southern Africa the spotted hyaena is
exceedingly abundant; in West Africa this is not so, though in certain areas of the Sud.rn
zone it is not uncommon. An interesting comment on the relative importance of the
two hyaenas in that zone in West Africa is provided by the fict that whereas the Hausa-
speaking peoples have only one or two terms for the striped variety they have nearly a
score of names and epithets for the spotted one. In Sierra Leone it was formerly common
in the open woodlands of the north; and, in fict, these animals proved a pest in Kabala
town between 1924. and 1930, coming down at night from the rocky hills to steal cattle
CROCUTA 355
hides pegged out to dry. T. S.Jones (1966) noted that the speaes was well-known and
common in the wilder parts of Bombali and Koinadugu; he foiuid (personal commiuii-
cation) that the numbers had been greatly reduced and that it had become localized in
hilly areas away from towns. Yet further west, Monard (1940) recorded that it was then
abundant in Portuguese Guinea; and G. S. Child says (1971, private communication)
that it occurs but is not common in the Borgu Game Reserve (extreme west Nigeria).
However, as far as the whole region covered by this present work is concerned
Croaita is very poorly represented in the British Museum, study material comprising
merely 2 skins and 4 skulls, half of each being juvenile. These come from Gambia and
north-cast Sierra Leone. It is certain that in recent years the species has become less
plentiful m West Africa. This is due in part to increase in human population and
extension of agriculture, resulting in less and less game; but in some measure it is more
directly due to a pohcy of poisoning, the Bauchi (Nigeria) local government, for
example, a few years ago offering a bounty of jCzjio/- a skin, resulting in near exter-
mination (Sikes, 1964b). Poisoning has also been carried out from time to time in Sierra
Leone by the Veterinary Department in response to complaints from Fulani cattle
owners that their calves were constantly stolen. Some of T. S.Jones's specimens m the
British Museum are the outcome of such control in the Gbcria area where about 20
hyaenas died, several in their subterranean holes. It is possible that widespread destruc-
tion has similarly been carried out throughout the length and breadth of West Africa.
Nevertheless, Matthews (1939a) found that although in Tanzania Crccuta was merci-
lessly killed at every opportunity, and sometimes in large numbers, it remained super-
abundant. The difference is without much doubt attributable to the very disparate
availabihties of food.
Description. Crocuta (fig. 46) is a far larger, more heavily built animal than Hyiuiia,
rather clumsy in appearance m fact. This greater size is succinctly conveyed by its weight
which normally lies between 65 and 75 kg, and may be more, as compared with 35 to
45 kg in the other species. It can stand 750 mm at the shoulder. Like the striped hyaena
its forcquarters are far more heavily developed than the hindquarters, and the forelegs
longer than the others so that the back is in consequence similarly sloping from neck
to tail.
Colour is widely variable and these differences have been used as the basis of several
local races, as discussed later. But speaking in broad terms the pelage has a ground
coloration of pale greyish-brown or yellowish-grey on which is superimposed an
irregular dorsal and lateral pattern of dark, roundish spots. These, mostly very distinct,
may be reddish, deep brown or almost blackish, and are of variable size, in the same
skin as well as between specimens, but are commonly of the order of 20 mm diameter.
This maculation is continued, though sometimes less distinctly, on to the legs and belly
but not the throat and chest. On the back and sides of the neck the spots are often
replaced by five, paler, not sharply defmed longitudinal bands; but these markings are
sometimes so obscure as to be virtually absent. The broad medial band on the back of the
neck is nearly always lengthened mto a slightly forwardly directed crest sometimes
broad enough to constitute a small mane; and it is very often continued posteriorly as a
spinal crest, rather shorter in length and less forwardly pointing. It usually fades out
356 THE CARNIVORES OF WEST AFRICA
somewhat to the rear ot the shoulders; and it is nowhere in any way comparable in
lengdi and importance to the crest o£ Hy', 111 use closing against the posterior fice of the lower carnassial. hi the
mandible the two premolars are broadly similar to /i^, consistmg of a large triangular
median cusp with smaller anterior and posterior cusps basally. The lower carnassial.
fELINAE 377
0)1, is quite different in profile from the other subtriangular cheekteeth since it consists
of two compressed blades, its anterior and posterior margins parallel and upright, the
superior, cutting, edges inclined inwards towards each other (fig. 2).
Although it is not strictly a part of the skull one small portion of the skeleton closely
associated with it must here be mentioned since it has been given unusual taxonomic
importance in its use by Pocock (i9i6g and 1917b) and subsequent authors for the
separation of the genus Panthcra from Fclis and AcinmY-'^'. This is the hyoidean appara-
tus; but as it is of practical use only to those with the facdities and inclination to
make a careful anatomical dissection it is not employed in this present account. Briefly,
therefore, without going into unnecessary detail the hyoidean apparatus consists of a
chain of small bones, known collectively as the suspensorium, passing from the bulla,
on each side, to further small bones at the root of the tongue and embracing the top
of the windpipe, hi most of the cats this apparatus, except at its cartilaginous extremities,
is fully ossified and the larynx is thus held up firmly to the base of the skull and limited
in its movement; but in Pantlura, that is to say in West Africa the lion and the leopard,
part of the suspensorium remains unossified and clastic and is much longer, allowing
greater freedom of movement. Clear detailed illustrations are to be found in Pocock
(i9i6g, igryb and 1939) and Mivart (1881).
Habits. The Fclinac are by nature very largely nocturnal or crepuscular, a fact
which is basically true of the domestic animal as of the wild species, though the excep-
tional, atypical circumstances of life in close association with man tend to mask it
somewhat in this case. But no species is exclusively active at night, and although in the
wild much of the daylight hours may be spent in sleep, all cats can at least occasionally
be seen during the day prowling around with no apparent purpose, or feeding, or
training their offspring, or grooming their coats, or, if young enough, playing. Never-
theless It is not mitil evening draws in and darkness approaches that the important
business of life, the hunting and killing of prey and the finding and couphng \vith a
mate, commences m earnest. In these matters, as in several others, the cheetah seems to
form an exception to the majorit)' of felines since its pursuit of prey, at least, takes
place in fuller daylight. Because of their predominantly nocturnal habits details of the
behaviour of many species remain obscure and in some cases virtually unknown. The
majority of cats, also, are soHtaiy except at mating time or when a mother is followed
by her litter; but lions live together more permanently in small bands and so, to a lesser
extent, do cheetahs, which at least often congregate together for hunting.
The cats, unlike the dogs, make little use of confmed shelters either for their daily
rest or for breeding. Sometimes they \vA\ make a temporar)- home in a cave or in a
large cavity amongst boulders, but they rarely make use of holes in the ground, a cover
of dense vegetation generally serving all their needs. As the vast majority of them are
excellent climbers rest and safety" is often foiuid in the lower branches of trees, curled
up m a crook against the triuik or stretched out along a stout bough; and possibly
some of the smaller forest species may use a convenient hole in a triuik, either at ground
level or higher up, for giving birth or shelter to the yoimg during their early babyhood.
Singularly little is knowai about the sex lives of the cats. Observations have been
made on some species in captivit)' but on few under natural conditions. So far as present
378 THI; CARNIVORES OF WEST AEKICA
kiunvlcdgc goes it would seem that in most species the male and female rcmam together
for only the limited period of coupling or of raising one specific litter. Li view of the
great disparirv' of bulk occurring in the felines no general figure for gestation or for
size of fiimily can be given, though as regards the latter the cats rarely produce at a birth
the large numbers which from time to time are born to some canid mothers, two or
three being a common issue and nine probably an exceptional maximum. The young are
knowii variously as kittens in the smaller species or as cubs in the lion and leopard. They
are always born in an incomplete state of development, mostly requiring lo days or
so for the eyes to open completely and yet more for them to become fully functional
as regards focussing; further, new-born kittens need one to two weeks to become
properly mobile and steady on their legs; and considerably longer for their teeth to
erupt.
The very sharp curved claws possessed by all cats except the cheetah, aided by their
very powerful leg muscles and relatively light bodies, enable most species to climb
trees and other fiirly vertical rough surfaces with considerable case and speed. Even
in the cheetah the young, before the claws become worn and blunt, are quite good
climbers; but in the lion, possibly owing to its greater bulk and general massiveness
of bmld, climbing ability is more limited and ascents arc normally restricted to the
lower branches of smaller trees, and those often with sloping trimks that can be readily
walked up. Yet, that actual weight has little to do with limiting the ability' to climb is
shown by the extraordinary power of the leopard to ascend vertically for a considerable
distance carrying in its mouth a heavy antelope — a vivid demonstration of the immense
muscular power of the jaws, neck and legs foimd in the Felinae. The power of the legs
is shown in another way, for the typical cats have the facility for making prodigious
standing jmnps, sometimes employed for the initial stage of a vertical climb or for
getting onto and over obstacles, but more importantly for leaping suddenly onto
prey.
Various gaits are used b}' the cats according to circumstances. The commonest is a
leisurely saunter with tail outstretched or sometimes raised. Wlien prey is being
deliberately stalked a similar action is used but very much more slowly and with the
body in a crouched position. When more purposefully on the move from place to
place than implied by the usual idle stroll all the cats, but especially the bigger ones,
engage in a springy trot; but when it is necessary to move really fast they bound over
the ground both fore and hindfeet more or less coming together then stretching fully
apart as the body flies through the air, one forefoot next touching the soil a little before
the second, the hindfeet then being brought forward together again to close proximity
on the ground with the forefeet.
A diversity of vocalizations is to be heard in the subfimily from the well-known
mew of the domestic cat, through the snarl ot the leopard to the roar of the lion, the
latter's louder calls being made possible by the far greater freedom of the lar) nx due
to the form and less ossified structure of the hyoidean apparatus in Panthcra as compared
with Fclis. In the latter genus some of the larger wild species utter sounds not much
removed from a "mew"; and most of them in alarm draw back die lips to expose the
teeth and "spit", hi several members i>f the subfamily a vibrant soiuid related to the
FELINAE 379
contented "purr" ot the domestic cat is uttered, though not always expressive ofserciiity.
However, between the Hon and the domestic cat Httle has, in fact, been clearly recorded
of feline utterances, and they are probably more difficult to express verbally than
those of dogs or hyaenas without the use of specially coined onomatopoeic words such
as "mew" and "purr".
The Fclmae, whose diet is possibly more completely derived from the flesh of
mammals and birds than that of any other section of the fissipedes, fulfil an important
function in preserving the balance of nature though since they are less numerous than
other members of the suborder their total effect may in sum be shghter. However,
the lion, leopard and to a smaller extent the cheetah exercise an influence in an other-
wise imtouched quarter as they are able to kill fully adult ungulates of a size beyond
the powers of lesser carnivores with the possible exception of a pack of hyaenas. The
essential part these animals play in a balanced countryside has not infrequently been
overlooked, with disastrous results; as, for example, when the offer of a bounty for
destruction of leopards as "vermin", or their overkillmg in plam greed for their valuable
pelts has led in the course of time to a marked increase of baboons with consequent
severe depredation of agricultural produce by vastly larger troops of these notorious
and wasteful crop thieves. By what, in turn, the numbers of felines are kept in check
is less apparent. There is always, as just mentioned, man out to kill unwelcome pre-
dators to protect himself and his property; or in search of warm and often highly
ornamental skins for his own use or for sale. Man apart, without doubt the smaller
carnivores are preyed upon by larger members of their kind besides forming meals
for hawks, eagles and snakes; but when it comes to holdmg in reasonable check the
lion and the leopard the position is not so clear. However, it must be remembered
that even the largest and most powerful animals arc very vulnerable whilst juvenOe.
There are many hazards between birth and safe maturity. There is first the uncertain
temperament of both mother and father which sometimes leads to abandonment
and not infrequently to destruction and cannibahzation of their own young; there is
always the chance of theft by hyaenas; there is disease or injury resulting in inability to
survive. Finally, as regards a species as a whole, there is the vital question of the suffi-
cicnc)', or even availability, of food. No area can support a greater number of pre-
dators than there is sustenance for, surplus to the maintenance of a stable population
in the food species itself, hi any struggle to obtain meals from a marginal or inadequate
source of food the weakest and less efficient predators soon die off. This factor is of
considerable moment in much of West Africa where over vast areas, owing to expan-
sion of human popidations and of agriculture, conditions for the survival of prey
species, of all except the smallest kinds, are now much deteriorated from what they
were even 50 years ago and are annually worsening.
Taxonomy. As a group the cats have excited a great deal of interest and attention
and in consequence, as Simpson (1945) has pointed out, not only is the literature
enormous but there have as well arisen "irreconcilable differences of opinion regarding
the phylogeny, and hence the major taxonomy, of the fclids . . .". It would be pointless
to attempt to discuss here more than a very small part of what has been written. What
follows is merely a brief resume of the position in so far as the existing cats of West
380 THE CARNIVORES OF WEST AFRICA
Africa arc affccud; and it .u the s.inic time affords sonic key to the conflicts of views
and nomencLuurc that may be encountered in the hteratiire.
There is httle or no dispute regarding the identity and individuality of the various
units which make up the group under discussion but only as to the taxononuc level
to be assigned to them; and in this matter there arc two widely divergent schools of
thought. At one time all the cat-like creatures were included in the single Linnaean
genus Filis; but it is now pretty generally agreed that two major separations from the
r\'pical cats can and should be made: firstly, that the cheetah, by reason of points of
external form, notably the absence of digital sheaths into which the claws can be
withdrawn, is clearly generically distinct and assignable to Acinonyx; and secondly,
that the lion and leopard (together with certain other extralimital large species) should,
on the score of the form ot the hyoidean apparatus, noticed above luider the description
of the skull, be ascribed to the separate genus Panthcra. Common practice today is
therefore to divide the existing cats into three genera, ft 7/5, Panthcra and Acinonyx.
It is with the first of these genera that disagreement starts; for in 1858 Severtzov
gave almost every known species subgeneric rank, to the number of nearly two dozen;
and though the majority of recent authors have continued to treat these mostly mono-
specific units as nothing more than subgenera Pocock (1917b) in his classification of the
Felidac and in two subsequent major publications (1939 and 195 1) has insisted on giving
full generic status to 14 names of Severtzov and other authors, hi this he had a sub-
sequent measure of support by J. A. Allen (1919a). Largely in consequence of his
attitude towards Fclh and these others as cognate genera Pocock felt the need to
emphasize the distinction between this group and those of Panlln ra and Acinonyx,
and this he brought about by raising all three to subfamily rank as the Fclinae, Pan-
therinae and Acinonycliinac. Pocock specialized to a large extent on the Carnivora
and more especially on the cats, his last published work being an imcompleted mono-
graph on the Felidae (195 1). His experience was unusually wide and in some ways
unique since he had, over many years, close acquaintance with livuig captive animals
and the dissection of newly deceased specimens as well as with abundant museum
material. His views, therefore, cannot be dismissed lightly; yet the general consensus
of opinion today, led by Simpson (194s), and with the single exception of Ognev
(1962), is that his taxonomic ratings are at too high a level and that the numerous
genera of Severtzov, Gray and others are best relegated to the position of subgenera
of ft/is, and that Pocock's new subfimilies are unnecessary. This view of classification is
adopted in this present work.
It may be added that Hopwood (1947) troni examination ot skulls came to the
conclusion that the lion was the most primitive of the great cats and probably merited
separation from the leopard and odiers to a genus of its own. The whole situation is
complex and, in truth, not very well understood, the present fluctuating classifications
of the Felidae being based on differing individual evaluations and interpretations of
characters derived often from quite inadequate museum material and without taking
into accoimt patterns of behaviour or vital information derivable only from living
animals. Until an obviously sounder based concept of the Felidae has emerged from a
study of the group 111 depth involving not only considerably better material but a fir
FELINAE 381
wider range of valid taxonomic characters as well there seems little pomt in tinkering
piecemeal with a classification and nomenclature which is widely understood and works
practically though phylogcnctically possibly capable of improvement.
As a matter of convenience the keys which follow deal with individual forms,
treating genera, subgenera and species alike without attempt to set them forth in their
correct taxonomic grouping. External characters are rendered difficult by phasal or
developmental changes in pelage colour and pattern.
KEYS TO THE SPECIES OF FELINAE
(Previous key page 160)
A. Cranial characters
1. Total length of adult skull over 180 mm ....... 2
Total length of adult skull less than 180 mm ...... 3
2. Total length of adult skull less than 270 mm; length of ;)•• under 30 mm
P. pardiis {[hJiic 439)
Total length of adult skull over 270 mm; length of ;)■* over 30 mm
P. leo (/).!(;(■ 460)
3. Upper premolars 2 only; nasals descend ver^' steeply from the inflated frontal
(fig. 54) ....... F. caracal {pngc 402)
Upper premolars 3 ; nasals do not descend so steeply ..... 4
4. Total length of skull less than no mm ....... 5
Total length of skull over no mm ........ 6
5. Bulla length approximately equal to the width of the rostrum across the
canines ........ F. /iftjca (yni^e 384)
Bulla length markedly greater than the width of the rostrum across the
canines ........ F. margarita {page 395)
6. Nasals broad posteriorly (about 20 mm) not very tapering; total adult skull
length about 160 mm ..... A. jubatus [page ^g})
Nasals narrow posteriorly (about 6 mm) sharply tapering; total adult skull
length not more than about 145 mm ...... 7
7. Postorbital constriction about 50 per cent greater than the interorbital breadth;
p^ small; total skull length about 125 mm . F. serval [page 412)
Postorbital constriction not much greater than the interorbital breadth;
p^ extremely small, total skull length about 140 mm F. aurata [page 425)
B. External characters
I. Head & body length of adult not more than about 600 mm; tail little more
than half as long; ears without any black pattern on their backs . . 2
Head & body length of adult at least 750 mm ; ears (except aurata) with some
black on their backs ......... 3
5S2 THE CARNIVORES OF WEST AFRICA
2. Eais si.-t taiily lugh on the head, pointed and with a snudl apical tutt; tail with
two or three siibterniinal black rings and usually a black tip
F. libyca {pin;c 384)
Ears broadly rounded and set low on the side ot the head ; tail without, or only
ill-dehned. black rings . . . . F. tiiargarita {pa(;c 395)
3. Tail at least half as long as the head c\ bod}' ...... 4
Tail well under half the length ot head & body ..... 6
4. Tail tapiering, clad with very short, cleise-lying, pialc brown hair, unspotted
in tlie adult but always with a contrasting blackish terminal tuft
P. leo (/') in the southern part ot the continent.
FELIS LIBYCA Forster African Wild Cat
Iclis lybiiH G. Forster, 1780, 111 Burton's h'ciniri;vsLlnihti: dcr I Vii/iis5ii;(" Tliitrc, cii. 6: 313. Gafsa, Tunis-
The name seems to h.ive been based on a not very clear description iurnishcd by tlie explorer James
Bruce. Forster's spelling of the specific name is an obvious Inysiis since the country fiom which it was
derived has from ancient times been written as Libya (see Ellcrman & Morrison-Scott, 195 1).
Fflis lynx lyhictisis Kerr, 1792, TlicAniiiial K/iiiji/i'i/i etc.: 136. C.ips.i, Libya(= Gafsa, Tunis).
Felis crislaia Lataste, 188$, Acl. Soc. liiw. Bonhmix 39 (i.e. (4) 9): 229. Not of Falconer & C.uitley, 1S36.
The name is the Latin for crested.
Fclis haiissa Thomas & Hinton, 1921, Novit. Zool. 28; 2, 3. Zinder, Niger. Type in the British Museum,
No. 2T.2.11.16, o; skin good, skull in fair condition but with the frontal region .and bullae partly
smashed. The name is derived from that of the people inhabiting the region ot West Africa whence
the specimen came. Valid as a race.
Ixlis lowci Pocock, 1944, Proc. zool. Soc. Loiul. 114: 68. Jebel Marra, 1200 m, Darfur. Type in the British
Museum, No. 23.i.t.66, 5; skin and skull both in good condition. This was named after the collector
Willoughby Lowe.
Felis lybica lyiicsi Pocock, 1944, Free. zool. Sec. Loni. I14: 68. Collected 56 km north of El Fasher, LDarfur.
Type in the British Museum, No. 23. 1. 1.65, ^ ; skin good, skull with right upper cinine area sm.ashed
and missing, and one ramus of the mandible Licking. This was named 111 honour of the collector
Rear-Admiral Hubert Lynes.
fffa ]yhica fo.xi Pocock, t944, Proc. zool. Soc. Loiui. 114: 71. Kabwir, 820 m, Nigeria. Type m the British
Museum, No. 12.1 1.7.5, sex ?; skin fair, with paws Licking, no skull. This was named after the collector
Dr. J. C. Fox. Valid as a race.
Felis libyca sai'anicola Dekeyser, uj^o. Bull. Inst. jr. Afr. noire, 12: 704, 705. Messirah, south Senegal.
Type in the Institut fr.in(;.iis d'Afrique noire, No. 48-5-38, i; skin and skull. The name is coined
from savanna .ind the Latin colo meaning to inhabit.
Distribution and general. The African wild cat (Plate 10) has an extensive dis-
tribution. In Africa Itself It is to be found over most of the continent from the Mediter-
ranean to the Cape with the exception of the closed forest block. It occurs also in a
restricted area of Southern Europe, that is to say in Italy and on the islands of M.ajorca,
Sardinia and possibly Corsica; and beyond this it ranges east across Palestine, Arabia
and Persia to Turkestan and northern India. With such a distribution, embracing an
exceedingly wide range of vegetation, it follows that there is an equally varied degree
of external appearance. This renders any general description difficidt; but colour and
distinctness of niaculation apart there are certain characteristic markings all or at least
some of which are, with greater or less degree of clarity, always exhibited. These arc
detailed below.
Five races have been .ittributed to the region covered by this present work; but this
point is more fully discussed in the later taxonomic section. It is natural that the abund-
ance of this cat should vaiy from locality to locality throughout its wide range according
to local circumstances, food supply, predators and the degree of severity of genera!
existence; in regard to West Africa, study specimens are fiirly rare though the species
FELIS LIBYCA 3S5
IS kiiowii, by repute, to occur from the extreme west to the extreme east, and broadly
speaking is not at all uncommon in the Sudan and Sahcl zones. It is also known from
well into the Subdcscrt zone at Tchsiderak in Air; another form occurs chiefly in the
Doka and, apparently more uncommonly, in the Guinea woodland. The species is
almost certainly absent from the high-forest. The most southerly specimen in the
British Museum is from near Obubra on the Cross River; but this would seem to be
exceptional.
Recognition of lihyca is sometimes made doubtful by two facts. Firstly, that it is
not unusual in Africa for domestic cats to go feral and, by force of circumstances in a
highly competitive environment, to lose all trace of their normal placid nature and
become so uncompromisingly aggressive in behaviour as to give the impression of
being unquestionably members of the true indigenous fauna, hi the second place,
lihycii does from time to time cross with domestic cats, the offspring sometimes dis-
playing the markings of the one and the character of the other. Within the experience
of the present writer there was some years ago a house cat in a station in Bornu (Nigeria)
which clearly exhibited the key markings of lihyai but had the passivity of temper
associated with an ordinary domestic pussy.
Descriprion. This (Plate lo) is one of the two smallest of West African cats though,
so far as can be judged trom the few study specimens with any live data, of pretty diverse
size from 400 to over 600 mm for head &: body length, and a weight usually quoted as
of the order of 4-5 to 5-5 kg though one reliable collector in West Africa has given a
figure as low as 2-7 kg for an animal said to stand 320 mm at the shoulder, which is
big for a cat. It has already been pointed out that the pelage colour and general body
pattern of libycii are extremely variable, ranging from greyish to reddish, with or
without markings of small spots which when present may or may not tend to rim into
transverse stripes. In known West African specimens this body pattern is absent or
very obscure; but in some extrahmital races it is a pronounced feature. In Asiatic forms
the spots are independent and veiy strong giving a quite different overall impression
from the more typical African animals, the appearance recalling rather that of a small
serval.
In accordance with the and terrain most commonly favoured by this cat the pelage
colouring is often basically of a sandy or buffish nature, the belly mostly whitish ; but
animals from the rather damper woodlands are distinctly more rufous. Despite this
overall variety of colouring and pattern there are seven points of recogmrion of the
species no matter where it occurs throughout its wide range, at least four or five of
which are always detectable in any given specimen. These are, roughly in their order
of constancy and importance:
Tail There arc always a short black tip and two, sometimes three black subterminal
rings or partial rings. Its length is usually roughly two-thirds of that of the
head & body.
Fed These are entirely hairy below except for the actual pads; and they are always
black below, this colour being sometimes linnted to the plantar area but,
on the hindlimbs, often extending some or all of the way to the heel.
386 THE CAUNI VORFS O I- WEST AH(K:A
Hiir.v' These .lie subtriangularly pointed and always plain red nr ochre on their
hacks and mostly with a short apical tuft of the same colonr.
/•'lice This IS marked with a narrow, mostly ginger, Ime from the outer corner of
the eye towards the angle of the jaw, and with a similar, shorter one from the
inner corner of the eye to the rhmarium. There is often, but not always,
a third line nearly parallel to the first of these and running lower down across
the cheek.
Foirlcfis At maximum development there are two complete black rmgs ("bracelets")
encircling the arm, one near the top, one about half-way down; but these
show various degrees of fiding, being sometimes visible only as half-rings
on the msidc ot the limb, or sometimes one or both lacking; and the colour
may decline from black through blackish-brown to ginger.
I'liroat This exhibits two, mostly ginger-brown, narrow transverse bands, of wliich
the anterior may be indistinct or lacking, but the posterior nearly always
perceptible.
Spine From just posterior of the shoulders to the root of the tail there is generally
a band of more pronounced, mostly golden-brown colour, never very sharply
defined and sometimes very obscure.
One of the many factors that are inconstant in the African wild cat is the length
of the coat which, apart from increasing from youth to maturity, varies in accordance
with ditfering seasons as well as with general ecological conditions, hi Pocock's accounts
of the different races (1951), for example, he quotes extremes of pelage length of 22 to
44 nnn on the flanks and 34 to 55 mm for the spinal crest. The texture of the fur as
well as Its overall appearance varies with this length from moderately soft and loose
to slightly harsh and close. The pelage is composed of three types of hair: very fme,
fairly long, dense underfur; lengthy flattish-sectioned bristle-hairs which arc of con-
stant diameter throughout all their length except for the tapering tip, and which by
reason of their much greater stoutness and pure white or very pale basal regions stand
out prominently when the coat is turned back; and almost equally long sub-bristles
with appreciably finer proximal portions than m the bristle-hairs but expanded distal
regions. The bristle-hairs are markedly more abundant in the spinal band than on the
flanks.
The coloration varies considerably, ot course, from race to race; but m general
terms, and as far as West Africa is concerned, the underfur, which is of die order of
14 to 17 mm long on the flank and 20 to 25 nnn in the mid-dorsum, is white or pale-
coloured for the majority of its length but has a short terminal zone of slightly more
intense colour, and occasionally a faint intermediate band. The underfur is not entirely
concealed by the longer and stouter bristle constituents of the pelage. Average ranges
of lengths for these latter 111 West African forms are: for the bristles 30 to 35 mm in
the spinal band, 21 to 27 mm on die flanks; the sub-bristles 2 or 3 mm shorter. The
coloration of these two types of hair is very similar: there is always a slender black
tip 4 to S mm long; subteriiiinally there is a pale band, often narrowly golden yellow
3
1
.1
/^^^ ~ s
Afiic.ui Wild C.it (I-i'li> lihyui): Car.ic.il (/c/l> i.ir.i..)/)
FELIS LIBYCA 387
fading into white or pale yellowish and 3 to 7 mm in length; this is succeeded proxi-
mally by a black or deep-brown band 2 to 3 mm wide; and tliis passes into a long white
base. In extralimital greyish forms the subtcrminal white bands accord a "frosted"
look to the fur; but this is not apparent in the generally more butfish or sandy West
African animals though it does account for the "ticked" appearance of the pelage in
these.
Skull (figs. 51 and 52). According to Pocock (195 1) there is no constant difference
between the skulls and teeth of lihyca and those of the European wild cat, F. silvcslris
Schrcber. The profde is very rounded, falling away from an inflated frontal region
just posterior of the postorbital processes, the nasals descending very steeply to a short,
blunt rostrum, the length of skull lying in tront of the infraorbital foramen being only
a quarter, or less, of that lying posterior to it. There is no sagittal crest except for a short
and usually not very highly developed posterior section joining the supraoccipital
crest. Only in a very occasional extralimital specimen is there any sign of development
of a crest across the mam part of the cranium. The supraoccipital crest is variable, being
always present; but the posterior portion of the cranium is m some specimens much
more deeply excavated than in others. This is partially, but not entirely, to do with
age; no conclusion can be reached with regard to se.x. Apart from this backward exten-
sion the braincase is broad and subglobular. The postorbital processes are long but
never join with the very well developed jugular processes to form a complete ring.
The orbit itself is very large. The interorbital distance is narrow for the general width
of the skull; the postorbital constriction relatively broad and inconspicuous, usually
though not invariably at least twice the former.
The zygomatic arch is strong, the anterior, suborbital, portion much deeper than
the posterior half. The palate is almost equilateral in form; there is a short postdental
extension divided from the main area by broad emarginations wliich reach at least as
far forward as the level of the posterior edges of the carnassials, and generally rather
more, its posterior border double-concave with a small medial spine. The mesopterygoid
fossa is wide, parallel-sided, the posterior processes of the pterygoids (hamulars) very
narrow and sharp. The bullae are large, the anterior chamber much smaller than the
highly inflated domed posterior portion; the paroccipital processes pronounced and
spreading over the posterior aspect of the bulla wall but not, or scarcely, fused to it.
The dental formula is nominally j-pn = 30, but there are occasional aberrations:
m some specimens (3 out of 9 available West African skulls) p~ is lacking on one or both
sides having been shed and the alveolus closed over, an occurrence that can be observed
in process in some skulls. Li one specimen p'^ is also present on one side, of minute
size closely approximated to /)-. Wlien present, p^ is always very small, no bigger,
sometimes less, than the outer incisors, p^ is a narrow but much bigger tooth, sub-
triangular in lateral profile, with a pronounced cingulum, a main cusp usually slightly
taller than the carnassial, and a very small posterior cusp but no anterior one. The
antero-internal cusp of the upper carnassial is small, sometimes very small, and sharp;
in^, which lies at right-angles to p*, is always small or very small, generally about the
size of /)2 hut in some examples rather larger.
i!88
Tlin CAKNIVOIU.S or WEST AFIUCA
Habits. Very little indeed is known ot the life ot this cat either in the wild state or
111 captivity. Like most cats it is extremely secretive in its habits, the difficulty of study
being added to by the flict that it is very largely nocturnal in its activities, though not
entirely so for it may from time to time be seen on the move, or at least awake lying
along a branch, during daylight. Normally it likes to shelter in a hole of not too large
size, either m a tree or amongst rocks or in a small burrow made by some more fossorial
animal, not too fir below the surface where the soil is warmed by the sim. In Pocock
(1944) the burrows of fennccs receive particular mention. Like all cats /i/iyw seeks good
protection from the rain, and a semi-open sleeping-place on the surface amongst grass
Fig. 51. i't7i.( lihyat foxi: skull, B.M. Nn. 13.2.5.3, .?,
I ; Literal view
is thus not much favoured, as such sites otten are by the wild dogs or hyaenas. Since
practically no field study has been carried out upon these cats it is unknown how
constantly any shelter may be occupied; except, ot course, that at breeding tune a
permanent secure home is absolutely essential for a few weeks. Most of the time,
apart from the short period of actual mating or for the few months when a mother is
accompanied by her kittens, these cats appear to lead entirely solitary lives.
The African wild cat feeds upon rats, mice, gerbils, hares, birds of small to medium
size (up to francolins), and lizards. It has been said 111 the southern part of the con-
tinent, where the species often attains considerably greater bulk than in the west,
to tackle larger prey such as antelope fiwns, or domestic stock such as lambs and kids;
and it is reputed to be a continual threat in poultry yards. Like other carnivores it will
probably, on occasions of need or when good opportunity off"ers, turn its attention
to large insects or swarms of plump termites; but whether it will eat snakes, as some-
times claimed, seems never to have been clearlv established.
FELIS LIBYCA
Nothing appears to have been recorded of courtship and matmg; but these animals
certainly often frequent the outskirts of human habitations and couple with domestic
cats to the accompaniment of a good deal of vocal outcry. The period of gestation is
usually held to be normally 56 days, though the longer period of 63 days is sometimes
Fig. 52. Fclis Ubyca foxi: skull, B.M. No. 13.2.5.3, J, >; i; palat.il & dorsal views
quoted, and the reputedly conspecific silvcsiris (see the taxononuc section below)
has twice been observed to take 68 days (Cocks, 1876). The litter consists of from 2 to 5
kittens, born with their eyes closed. No details of development appear to have been
published.
390 THE CARNIVORES OE WEST AFRICA
Fills llhyca is certainly of a fierce disposition and it is often stated to be quite un-
tamcable; but it is pretty clear that it was, in fact, tamed in ancient Egypt and became
the chief ancestor of modern domestic cats, being carried thence eastwards to Asia,
and westwards and northwards to Europe, where it appears to have interbred with the
local wild cat, illvtstrls, and given rise m this way to at least one ot the two pattern-
forms of the common household animal (I'ocock, 191 1 and 1951).
Johnson & Lester (1929) reported fuiding one of these wild cats, under the then
commonly used name Fells ocnata, at Sherifuri (i i°5S N, io°oS E) to be infected with
'I'rypimosoma hniai.
Taxonomy. The taxonomy is uivolved and, like that of the Felidae 111 general,
the subject tif much disagreement and consequent abundance of literature. The trouble
stems from the fact that, while there is a strong fundamental resemblance current
throughout the entire subgenus that unites it and renders it a matter of ease to perceive
that any member can only be a Fells, there are a number ot basic pattern-forms that
on the one hand appear to establish sound points ot separation yet which in the sheer
abiuidance of their variation furnish such stages of intergradation between widely
diverse forms that it becomes ditiicult to know where to draw distinctions or whether
such distinctions have any real validity.
The problem starts at specific level. Ilalteiiorth (1953) and authors following him
now consider that the African wild cat, F\lis llbyci, is not a true species but nothing
more than a race of the Euriipean wild cat, Filis sllvcstrls Schreber. Pocock (1951)-
on the other hand, while admitting the close relationship of the two, held that they are
nevertheless taxonomically distinct. The fact is that on the existing evidence it is impos-
sible to be defuiite. If accepted in the wide sense now claimed for it, silrcsnis embraces
extremes of such vastly diverse appearance, between diose of northern Europe, southern
Africa and central India, that it becomes diliicult to credit that they could belong to a
single species, though there is admittedly a considerable degree of intergradation.
Pocock, nevertheless, with no apparent hesitation accepted half of this proposal, in-
cluding the spotted cats of Asia with the grizzled cats of Africa in the one species
llhyut; but while conceding that there is no constant difference between the skulls of
that species and sllvistiis he considered that the two are always clearly separ.ible by
features of the pelage pattern and the nature of the tail; and, what is much more, that
this separation is specifically valid. Such characters are indeed slender as the basis of
specific distuiction; and the two forms certauily breed with one another as Pocock
himself (191 1) practically demonstrated. The two are in fact held to be the common
ancestors of one pattern of the domestic cat, the blotched tabby or c.j/i/i-phase, pre-
sinned to have been brought about by the infusion of soinediing from sllvcsnis into
the basic striped tabby or torqtuUus-phAi.c derived directly from llhyui. Data concerning
the occurrence at the present time of the two reputed forms ot wild cat, silrcslrls and
lihyca, in the same localities in southern Europe or the Mediterranean islands, with
the consequent possibility of crossing in the wild, is sketchy and inconclusive.
It IS strange that Pocock, with his very great interest in and deep knowledge ot cats,
should in his last, detailed and most mature monograph on die Felidae (195 0 have
offered not the slightest hint of the possibility that lihyca and silnsliis might be one
FELIS LIBYCA 391
and the same species, though the probabihty of this had been suggested in the htcrature
for some years (Schwangart, 1928: 28 and 1932; Hakenorth, 1941). He apparently
did not think the suggestion merited mention even in the paragraph in which he
stated the two species to be closely allied and that there was no constant difference
between their skulls and teeth. Nor have others, for example Ellerman, Morrison-Scott
and Hayman (1953), thought fit to unite the two groups, Without doubt there is a
not inconsiderable case for such a union; but the matter is still every bit as much one of
opinion as the case for the opposing view, and on the existing data is beyond proof.
Such a problem, in common with similar ones elsewhere can yield only to the applica-
tion of more sophisticated and positive taxonomic techniques than mere estimation
from morphology and pattern, accompanied, perhaps, by a clearer defmition of
species. Meanwhile, there seems no incontrovertible reason for the rejection of a
convenient as well as possibly correct classification ; and lihyca is therefore herein treated
as a species in its own right.
Attention must now be directed to the question of subspecies. Pocock (195 1) accepted
and furnished very detailed descriptions of 25, of which 16 arc African, apart from a
number of specimens wliich he did not feel able to place with certaint)'. There is httle
doubt that some of these named forms could reasonably and advantageously be
synonymised. Once again the ditliculty arises from a wide range of individual variation
leading to a multitude of forms and intergradation with consequent doubt as to what
degree of difference merits a fresh name. This, of course, must always remaui purely a
matter of opinion. Study of Pocock's subspecific descriptions clearly shows a remarkable
degree of variation, both of colour and marking, admitted by him to a single form;
and many of the taxonomic diagnoses specifically state that there is intergradation
with one or more other forms. One is left with the impression that without knowing
the provenance of a skin it would very often be difficult if not impossible to assign it
with confidence to any particular race. J. A. Allen (1924) also made it clear that a series
of 15 from a single locahty (Faradjc, Congo) presented such a wide range of colour
variation irrespective of age or sex that the material helped to show the imcertain
basis of forms founded on single specimens, there being rvvo extremes, one distinctly
grey, the other rufescent, intergrading through intermediate stages. It is fair to add that
Pocock (1951) considered that this series may have been nothing more than pure or
half-bred feral house cats, though the fict that the skins were gathered together over a
period of many months, partly as freshly collected animals by the expedition and
partly from local African hunters would, if this were so, seem to be stretching co-
incidence a long way. Dorst (1950), also, foiuid a wide range of colour from red to
grey in a very useful series of 15 from a restricted collecting area 6 to 16 km north of
Lome (Togo). This series also demonstrates that there is considerable range of size
in a single population of these cats.
The practical value of the attachment of a third name in such circumstances becomes
questionable. There are indubitably recognisable distinctions: between the wild cats
of Mediterranean Africa and those of the temperate south ; between those of the arid
tropical pre-Saharan zones and those of the moister and denser Guinean %voodlands;
but to multiply such distinctions until they become so slender as to be incapable of
392 THE ( ARNIVORF.S OT WEST AIIUCA
more precise dctmition than at present achieved seems to lack point. Dekcyscr (1950)
made some attempt to rehire races of the African wild cat more or k-ss to dcfuiite faunal
zones as defined by Chapin (1932 : 90), and it seems that some such scheme is a reasonable
basis for the recognition of subspecitic forms, at least for West Africa. As far as this
latter region was concerned he postidated three : one in what is now known as the Sahel
and the northern portion of the Sudan woodlands, reaching eastwards to the Nile
and beyond; one m the southern portion of the Sudan, the Doka and the Guinea
woodlands as tar east as roughly the Beinie and the western shore ot Lake Chad; and
one in the same zones but continuing east ot this limit as far as the north-eastern Congo.
Enough has been said about intergradation to make it clear that no precise boundaries
are likely between forms; but recognisable breaks are more probable 111 a north-south
direction between different ecological zones than east-west within an unaltcring
vegetational continuum. From examination ot West African specimens it seems to
the present writer that they tall conveniently into two groups, one occupying the
northern Sudan, Sahel and Subdesert zones, and the other southwards of diis. The
firmer, m which the basic dorsal tone is whitish-grey or buff, and whatever red there
may be in the pelage — and there is sometimes a good deal — is contmed to the spinal
band and to any pattern of spots that may be developed, is Ihiussa. In this the belly is
fundamentally white though often marked with pale red, sometimes heavily. The
second group, /c)a/, is appreciably d.irker 111 tone, and red can be seen to suffuse the
entire de^irsal and flank pelage, the spinal band and any spots being correspondingly
deeper. All but one of the British Museum skins are locally purchased and incomplete
so that it is difficult to be positive about the belly colour, but this too seems to be
dark and reddish rather than basically white as in the other group. These matters are
entered into 111 more detail 111 the following accounts of the two forms.
Felis libyca haussa Thomas & Hinton Hausa Wild Cat
Distribution. This is mostly found m the Sudan and Sahel woodlands, but three
specimens were obtained by Angus Buchanan in Air in the Subdesert zone. The places
m West Africa from which specimens exist in the British Museum arc:
SutLvi :oiit : Farniso, Dan Kaba (Nigeria)
Salul zone: Zinder (Niger, type); Fort Lamy (Chad)
Siihdcstrr :oni: Tchsiderak (Air, Niger).
There is also a skin from an unspecified locality 111 Ghana; and a skull from the Gombe
River (Bauchi, Nigeria; Sudan zone); as this is without a skin it is subspccifically
indeterminable, but from its provenance should belong here, though this is discussed
later. This gives a total of 9 skins and 8 skulls, of which 6 match, I being very juvenile.
Taxonomy. Pocock (195 1) divided these specimens between hcuissii, lyiicsi and lowti,
the two latter with some uncertainty. The present writer questions the worth of the
distinctions of colour, pattern and size drawn. A good deal has already been said of the
capricioitsness of the two former characters; scarcely any two existing specimens arc
precisely alike, and m these circumstances, with no real knowledge of normal popula-
tion range, comparing a maximum of three skins from one locality with even less
from others, drawint; fine distinctions between shades of buff 111 the coat, or the amount
FELIS LIBYCA 393
of red along the spmc, or black on the feet, or the precise tint ot the ears seems to be
straiiiiiig after a precision that the existing material docs not in fact warrant. The same
applies to pelage length: for though valid differences may well exist between the
coats of temperate, Mediterranean, animals and those of warmer regions attempts
within the latter zones to make taxonomic deductions from this character, which is
variable m response to season, age and moult as well as temperature and humidity,
by the comparison of single specimens are unconvincing.
The form haussa was distinguished by Pocock from lynai and lowci solely by its
smaller skull. On the face of it, from the figures given for total length, there is some
support for this view; but once again we have no idea of the normal range of size,
and examination of the table given below on page 400 shows that the means of most
of the other cranial and dental measurements of such limited study material as exists
are fairly close and there seems no very strong reason at present for separating haussa
from the outwardly very similar lyiusi. Turning to the suggested form lowti which
Pocock erected from a single skin from Jebel Marra and to which he provisionally
assigned a West African (Fort Lamy) skin, the type is certainly a shade darker and
greyer than the haiissa-lyiicsi material; but it nevertheless belongs to the grey-buff
group, not to the red phase, and the erection of a separate subspecific name for it is of
doubtful justification.
If the reputed differences between supposed forms were eventually found to hold
true for long series from the given localities there might be sounder support for a
multiplicity of names; but there is an at least equal chance that more abundant study
specimens would demonstrate the futility of attempting to draw an excess of minor
distinctions. Meanwhile it seems most satisfactory to regard all the wild cats of the
transcontinental dry Sudan and Sahel woodlands, as well as those from the yet more
arid Subdesert areas, as belonging to a single race, haussa.
Description. Iii view of the range of variation no more precise description can be
furnished than that the overall impression is one of a palish cat, basically huffish or
light-greyish and with a little red mostly in the spinal band or such faint pattern of
spots or lines as may in different cases be developed. The whole underside from chin
to anus is long- and loose-haired, fundamentally white but sometimes faintly spotted
or suffused with very pale red.
The skull from the Gombe River (mislabelled Gonebc River) mentioned earher as,
vcgetationally, presumably belonging here is nevertheless, as the table on page 400
shows, in several cranial and dental respects appreciably larger than any of the iden-
tifiable haussa group of specimens, and much more nearly corresponds to foxi. Without
a skin it is impossible to know whether this indicates a wider range of size for haussa
or that foxi spreads deeper into the Sudan zone than hitherto supposed.
Felis libyca foxi Pocock Mid-belt Wild Cat
Distribution. The Doka zone may be regarded as the centre of distribution of this
form which, however, ranges a little firther north into the southern part of the Sudan
zone and, apparently less commonly, soudi into the Guinea woodland. Seven specimens.
394 THE CARNIVORES OF WEST AFRICA
all from Nigeria, exist in the British Museum but constitute a very imsatisfactory
set of study material consisting of hve skins without skulls and two skulls without
skins. Of the former only the type appears to have been obtained at first hand by the
collector, and that is in only fair condition; the others are mostly incomplete and
give the impression of having been purchased from African hiuiters or in local markets,
their exact provenance being thus open to some doubt. Three of the seven specimens
here attributed to foxi, the rv-pe and the two skulls all from Kabwir, originate from
the Doka/ Sudan interchange zone: one, said in general terms to be from Zaria district,
is probably from the Doka zone; two others from the Kode area 24 km N.-E. of Lau
(on the R. Benuc) and from Langa 24 km N. of Lau, would thus come from the
Sudan woodland; and one from Mkp.mi, Obubra. This last is somewhat of a surprise
and has the interest of being by a very long \vay the most southerly of all known West
African examples. This village is nominally in the high-torest belt but lies opposite,
across the Cross River, a large expanse of Guinea-type woodland. It is presumably
from this last that the specimen, part of Sanderson's collection but obviously of local
purchase, may well have come — if, indeed, it did not enter the area from a distance by
way of trade along a well-established route, the river. Of this set Pocock dealt only
with the Kabwir skin and two skulls, and the Zaria district skin, referring them all
to fiwi, the last somewhat hesitantly. There seems little doubt that this was the form
that Dckcyser must have had in mind when he, apparently unaware that foxi had
already been erected, described sauanicohi from Messirah, Senegal, in Guinea woodland
vegetation, and projected its range from Senegal to Lake Chad.
Description. The external character that differentiates this race from liiiiissa is the
deeper, sometimes much deeper, red colour which suffuses the whole coat. There is,
however, the usual variety of tone and pattern. The type shows no pattern of spots
whatsoever, the only disthict mark on an otherwise plain ticked coat being the deep
red spinal band. The other specimens all exliibit spots in a greater or less degree, tending
to coalesce iirto transverse bars, this fusion being complete m the Zaria district skin.
The miderside is paler than the back but in no sense fiuidamentally white, as in lunissii.
being at most a light brown. It will be seen from the table on page 400 that the skull
and teeth o£foxi, as deduced from the two Kabwir skulls devoid of skins, are both quite
appreciably greater than those o{ luwssti.
The two skins from near the Benue north of Lau present some difficulty. They are
incomplete lacking the heads and parts of the legs, feet and tail; but though they have
all the appearance of pelts originally inexpertly prepared they have subsequently been
soft-dressed and their places of origin given with some precision by the collector —
of whom no particulars beyond the bare name have been discoverable. The two
localities he within a few kilometres of each other and can with some certainty be
regarded as lying within a uniform biotope; but the skins differ fairly markedly in
colour. That from Kode is not very dissimilar from the Cross River skin; the one from
Langa is of a distinctly paler tone, though still much redder than any luuissn specimen.
The imdersidcs, too, are different, though in the incompleteness of the skins it is impos-
sible to be precise. The darker, Kode, skin has the chest and at least the after part of the
belly pure white; the paler, Langa, skin appears to have very little white. It was the
FELIS MARGARITA 395
whiteness of some of J. A. Allen's reputed nihiihi specimens that caused Pocock to
suggest that they were pure or hybrid feral house cats; and similar considerations
may be applied to the two Benuc animals. The scries of 1 5 skins from north of Lome
(Togo) reported on by Dorst (1950) obviously belong to foxi; but they demonstrate
that there is a considerable range of colour in animals from a small uniform area; and
Dorst concluded that it is the greater intensity of pigmentation rather than actual
colour that distinguishes foxi from haiissa.
A problem of a somewhat different nature is presented by the skull from the Gombe
River, cited above under liaussa. This locality lies some 100 km further north than the
area from which the two skins just discussed came, rather deeper into the Sudan zone.
It might thus be assumed to have come from a luuissa type animal; but, as the measure-
ment table shows, it is of a size much more representative of foxi. However, Dorst
(1950) came to the conclusion that size was an unrehable taxonomic character in these
cats since there was wide variation in specimens of apparently equal age; and all his
Togo skull material of foxi was more comparable in this respect to hmissa rather than
to the type of that race. The skull is illustrated in figs. 51 and 52.
Other forms. Finally, Dekeyser's suggestion that the West African Guinea form
(which he called savanicola) is replaced eastwards of Benue-Chad by the north-eastern
Congo ruhida Schwann must be examined. This latter is a red form in the type skin of
which the pattern is not only unusually clear but also consists, exceptionally, of spots
that show no tendency to coalesce into transverse bands. Pocock suggested that this
was probably nothing more than an individual idiosyncrasy; but other skins with
the same peculiarity have since come to light from the southern Sudan. The only
other skins in the British Museum collection attributed to ruhida by Pocock are all
too young to be properly comparable. J. A. Allen's (1924) series of reputed ruhida
from the north-eastern Congo, on which Pocock cast doubt as possibly pure or half-
bred feral house cats, shows that the form is widely variable and that one cannot take
any particular colour or pattern as narrowly representing the race. Taking this much
less restricted attitude than that adopted by Pocock there is little doubt that ruhida
closely resembles some of the West African skins herein considered to htfoxi, and there
is a considerable possibility that a single variable, constantly mtcrgrading form — but
not a cline — ranges across the continent in the Doka-Guinea zones from Senegal to the
north-eastern Congo ; and that^o.v/ in consequence may eventually prove to be nothing
more than a synonym of ruhida. This last position is not adopted in this present work
because there are factors of uncertainty : the material from both sides of the range is
quite inadequate; the lacimae in collecting localities great; and such figures as exist
for cranial and dental measurements — two skinless skulls from the west and Allen's
slightly doubtful scries from the east — indicate that_/t).vi might be appreciably larger.
FELIS MARGARITA Loche Sand Cat
Felis margarita Loche, 1858, Revue Mag. Zool. (2) 10: 49-50, pi. i. Near Negousa (misprinted Ncgon(;a),
Algeria. The type has in all probabilit)' been destroyed (Haltenorth, 1953: 63 f.n.). The spelling of
the specific name as given here is retained in order to avoid confusion in view of its long and wide
usage. Nevertheless, it seems to the present writer that it was clearly originally a lapsus on the part of
396 Tin: caunivoiils oi wesi aiku.a
the author or, more prob.ibK, a printer's error. The species was dedicated to Cominaudaiu (sub-
sequently Cicncral) Marguentte to whom Loche went to some length to express his gratitude for
constant aid as W'cll as the hope that natural-scientists would always preserve this helper's name. It is
thus unthinkable that Loche would have nisulted a high-ranking soldier and friend by deliberately
attaching a tenininie forename to an animal which he hoped would lastingly connnemorate this
officer's kindness. It is unfortunate that yet a second error crept into Lochc's description when one of
the 't's from Margueritte's name was dropped later in the text; this, however, is no sanction for the
spelling iiiargarila or iiicir^ucjiui, and onU' Trouessart (see below) eventually grasped and recorded
the author's true intention. Jcan-Auguste Margucritte published a work on hunting in Algeria and
earned fame as a general in the Franco-Prussian war; it is to be regretted that mischance and con-
venience of usage deny to his name the honourable position that Loche obviously desired.
Felis inari^inala Loche (Gray, 1S67, Proi. zocl. So{. Loud.; 275. Printed in error tor the above).
Felis lalif^aia itiarf^aritac Loche (Trouessart, 1X97, Calalcoiis Mammalinm, I: 363. An emendation of Loche,
putting the name in the more usual genitive case rather than simply in apposition).
Felis lihym iiiar::
T' ? ? ^
6 - rh i
I I I
t^ Tl- \0 r^ "^ ^C
:c r^ — :.c
C '-. "-, o
— O -^i-^. V-,r^r] 0>^ O
- ^, -t '1 r|
>c r- "- c
o r- 7-> o ■_'"■ r-- --1
'' ? 9
r;, n S --'-
ri •y-. r] O C' X T^
■^. -^ c ^ '^ b b--
O --l DC
-*-«-,_ '^*
t -- r- -^i r^ 1^ -t -t
1-1 -+ o o
3C r- r', c
y: o c f~~
nr-, r| -t .
o ;>;
2 Tj. ^-f -g lv.± i J -6 -
, _o O T3 c ^ c
^I^ -^ t- (J N -il. /^
o s p ,^ ;? != -s =
FELIS (caracal) 4OI
liandlc it. During its captivity it developed little real response to domestication though
it would allow its head to be scratched and closed its eyes in satisfaction while this was
being done; but it would never voluntarily approach with the object of receiving such
attention as a domestic cat, or some captive animals, would.
It is interesting to note that the dense mat of hairs covering the soles completely
obscures the pad marks in the sand so that the spoor is nothing more than the plain
outline of a cat's foot, claw marks only showing when the claws have been deliberately
extended prior to making a spring.
Taxonomy. Fclis margiwita seems fnrly clearly to be a species in its own right,
the differences of external form between it and lihyca, concerning the ears and feet,
as well as of the skull and teeth being of a specific rather than racial order. Nevertheless,
it does in a degree share some of the pelage markings odibycn and it has been suggested
that it is, m fact, nothing other than a subspecies of this — itself often held to be only a
form of sih'cstris; but this view is not much favoured today, though margarita may
well be a specialized desert derivative of lihyca. Animals at the eastern end of the
known range, in Turkestan, have not only been placed in a different species, thinobia
Ogncv, 1927, but also in a separate genus, Ereiimehirus Ognev, 1927; but while Haltc-
north (1952) accepts the species, but not the genus, both Ellerman & Morrison-Scott
(195 1) and Pocock (195 1) regarded this as nothing more than a race o( in(ir<^(irita. The
point IS of no particular concern to West Africa.
Regarding subspecies, since the type of margarita si iisii striae from Algeria is un-
known It is difficult to say whether Pocock's proposed mcincrtzhagciii, also from Algeria,
is materially different and therefore valid as a race. But while this is of little direct
moment to this present work it seems evident from the single specimens available,
that is to say the two types, that nirciisis is sufiicicntly distinct in size and colour from
mcimrtzluigcni to merit separate recognition. The West African animal should therefore
be referred to as given below.
Felis margarita airensis Pocock Air Sand Cat
The form, which is known only by the type skin and skull in the British Museum
from In-Abbangarit, on the extreme northern edge of the Subdesert zone, has been
sufficiently dealt with above. A second skin from 300 km further north, from Touaret
(northern Niger) in the Desert zone, might also be looked upon as belongmg to this
race; but it is not only paler but also of a very much more lively light red colour.
Subgenus CARACAL Gray, 1843
Caracals
This monospecific subgenus differs from Fclis sciisii srricio partly in its larger size but
chiefly in the possession of relatively much longer, very pouited, long-tufted, blackish-
backed ears and a very short tail measuring little more than a third of the head &.
body length. The webs between the toes are also markedly shorter. The skull differs
402 THE CARNIVORES OF WEST AFRICA
111 h.ivine; oiilv two upper premolars, a thud anterior one being only exceptionally
developed. In addition to this the nasal branch of the preiiiaxilla is a narrower and longer
bone than in Filis. wedging much further between the nasal and the maxilla so that
contact between these two latter bones is very much shorter and sometimes almost
non-existent, the premaxilla practically abutting onto a narrow tongue of the frontal.
This feature is, however, shared with L pliiilnnis. Cdnical is of wide distribution through
most of Africa and much of western Asia.
FELIS CARACAL Schreber Caracal or Desert Lynx
fi'/is Ciiracal Schreber, 1776, Dii- Sinn^ctlncre . . ., pi. no; 1777, text 3: 413 and 5S7. Type locality Table
Mountain, Capetown, South Africa. The priority of this over Miiiler's identical naming, which follows
below, and a similar one that has wrongly sometimes been attributed to Gueldcnstaedt (1776) has
been fully discussed by J. A. Allen (1924: 279-2S1). Allen's conclusions were either unknown to or
ignored by Pocock (1939) who continued to follow Matschie (1912) in citing Miiller as the prior
author; biit they were at a later date accepted by Ellerman & Morrison-Scott (1951). The derivation
of this name has been given above in the generic synonymy.
R-lis caracal P. L. S. Miilicr, 1776, Dcs Riticrs Carl von Linnc . . . NalmsyMm . . . Supplement: 30. Type
locality given as Arabia but was more probably Constantine, Algeria.
Caracal welaiiotis Gray, 1843, List of the specimens of Mammalia in the collcciiou oj the British Museum: 46.
A renaming of Schreber's species on transferring it from Fetis to a new genus Caracal. The name is a
combination of the Greek im7i7t, melauos black, and ous, otos ear, with reference to this diagnostic
character.
Caracal caracal pocciloiis Thomas & Hinton, 1921, Kovit. Zool, 28: 3-4. Type locality Mount Baguczan,
Asbcn, Niger. Type m the British Museum, No. 21. 2. 11. 19, V; skin in good condition, skull good
except for the left 1' and p^ missing and c broken. This name is derived trom the Greek poikilos vari-
coloured or pied, and ous. olos car, from the "frosted" nature of the back ot the ear in this torm.
Distribution and generaL Outside Africa the caracal ranges across the Arabian
peninsula, Palestine, Syria, Iraq, Iran and Turkestan as far east as northern India.
In Africa itself its distribution once embraced the greater part of the continent from the
Cape to die Mediterranean with the exception of the closed forest block and central
Sahara; but while it has never been a common animal it has now like so many other
mammals in the ftce of advancing development become much rarer than it used to
be and is now absent from the southernmost parts of the continent and is fairly unusual
m Morocco, Algeria and other north African countries. However, it still occurs widely,
though sparsely, in the tropical open woodlands and is at its most plentiful in the
eastern territories. In West Africa it manages to retain a foothold but in many areas
this is now very tenuous. The only material from this region in the British Museum
consists of 6 skins, mostly incomplete and without skulls, from Gambia, Lake Chad,
Ghana (Ganibaga) and Nigeria (Kode and unspecified); one skull without a mandible
from Nigeria; and a single matching skin and sktill, the type o£ pwcilotis, from Mount
Baguezan, Ai'r. In so far as can be judged from the localities cited, the vegetational
zones which these specimens inhabited were the Sudan, the Sahel and the Subdesert;
but the Gambia animal may have come from the Guinea woodland; and G. S. Child
(private communication) says that it occurs in the Borgu Game Reserve, which lies
in the Dok.i zone.
Besides this museum material there is a very interesting record supiported by a photo-
FELIS CARACAL 4O3
graph {Aniiihils, ]m-i. 1970) of two cubs taken from a cave near Ado Ekiti (Ondo, west
Nigeria), a place which lies close to the high forest; but it is now largely open woodland,
and being rocky country' offers attractive refuges. The literature contains several
generalised statements asserting occurrence of this animal in most of the West African
territories from Senegal to the Central African Repubhc without citing particular
specimens; but Frade (1949) published a defmite record for Bafata, Portuguese Guinea,
apparently in the Guinea woodland.
Descriprion. Fclis caracal (Plate 10) is a medium-sized cat, that is to say it is appre-
ciably larger than lihyca and iiun(;arita but very considerably smaller than "the great
cats" — in Africa, the lion, the leopard and the cheetah. There is, however, some con-
siderable range in size though satisfying figures for this are difficult to come by. Only
one West African specimen has any collector's measurement data. These relate to a
fully adult animal, the type o( poccilotis: but since this is both female and an inhabitant
of difficult subdesert terrain it is expectedly of less than average size. Nevertheless,
the dead weight given by the careful collector, Buchanan, 5-9 kg is vastly mferior to
that indicated by Shortridge (1934) for South-west African animals, 18 kg, or even
that given by Wilson (1968) for Zambian specimens, 12 kg for a female to 14-5 kg
for a male. Buchanan's type was recorded as having a shoulder height of 400 mm, from
which Shortridge's range, 406 to 457 mm is not so widely remote.
Colour, too, IS a matter of considerable variability. There are dark specimens with a
good deal of black mixed into the fur, pahsh sandy-grey specimens, and those of a
lively red. Shortridge (1934), indeed, speaks of the intensity of colour varying within
the same district. No albino seems to have been reported ; but, as in a good many other
of the Felinae, melanos are known; there are two Kenya skins in the British Museum;
Pitman (1949) recorded one for Uganda; and one of the two cubs from west Nigeria
referred to in the previous section was, from the colour-photo, very dark though
apparently not a true melano. With these reservations the caracal can be described in
general terms as dorsally of a more or less uniform greyish-sandy or reddish colour
from the crown to the end of the tail. There is usually a faint and very ill-defmed,
slightly darker spinal band. Much of the upper side is often "frosted" to a greater or
less degree, this feature showing up better on the darker, redder coats than on the pale,
buffer ones. The "frosting" is due to longer or shorter subterminal regions on many
of the bristle-hairs and sub-bristles, interesting in that they are not really complete
encircling bands as in most "ticked" mammals but are pure white on their upper sides
and of the normal pelage colour below. The dorsal pelage, as exemplified by the few
West African specimens, is variable in texture, density and length, possibly as much
according to age, moidt and season as to locality. In some the fur is moderately long and
soft, in others short and rather harsh. It consists of abundant long or fairly long underfur
which is colourless or very pale brown with darker tips and amongst which are scattered
bristle-hairs slightly oval in section, with long white bases and black tips; and there
are also less frequent sub-bristles with long very fuie petioles and flat-sectioned blades.
Average lengths for these components are: underfur 9 to 11 mm, bristle-hairs 13 to
20 mm, and sub-bristles 13 to 18 mm, the petiole measuring 10 to 12 mm, the flattish
blade 4 to 6 mm.
404
TUP. ( AUNIVOKIS Ol WHST AlUICA
The riaukb .uc soiiKwh.it p.ilci tli.m the b.ick. Yhc bell), lIk'si .md smnctinics the
throat arc pure white or whitish and bear, especially on the belly, pale red spots wliich
vary m their distinctness, their abundance and the amonnt ot the underside they cover
accordhig to dilTerent specimens. There are distinctive fiicial markings, one being a
fairly pronounced dark line from the inner corner of the eye to the rhinarium. Above
the inner corner anti below the whole of the eye are white patches which are sometimes
surticientlv extensive to give the impression of a complete encircling ring. Tliere is
also a prominent white patch cither side of and below the rhinarium, continued
postenorlv along the edge of the upper lip, at first narrowly and partially interrupted
bv a blackish patch from which the nnstacial vibrissae .inse and tlien expanding onto
Fic. 54. Filii ciViunl: skull, Type ot pociilinis, H.M. No. 21.2.1 i.n).
I ; l.itor.il vic\
the cheek. The lower lips are wholly white. Tile eyes are greeiiisli, the pupil an upright
ellipse which never narrows to a mere slit as in the typical cats.
The ears are very characteristic. They are of a tall triangular shape, quite different
from the rounded pinnae of the serval or rather squat triangle of the wild cat; and they
carry at the apex a long pencil ofhairs, by f^ir the longest tuft ofall the felines, measuring
about half as long as the pinna itself. The backs are covered with very short close-lying
hair, their colour often sweepmgly stated, as a diagnostic character, to be black. Yet
this last IS true only in a general sense, the overall impression in sharp contrast to the
otherwise sandy pelage being broadly of that colour; but a closer look shows that there
is invariablv some white mixed with the black, and in a large number of cases a con-
FELIS CARACAL
405
Fig. 55. Felis caracal: skull, Type o( poecilotis, B.M. No. 21. 2. 11. 19, 9i x i; palatal & dorsal viewi
406 THE CARNIVORES OF WEST AFRICA
siderable amount, luitil at its extreme the colour becomes more silvery than black.
This latter character was largely relied on by Thomas & Hinton for the differentiation
of the West Atrican race [UHcilolis; but it is by no means so uncommon as the description
of that form might lead one to suppose. The base of the piima together, often, with a
small surroimding area on the head, is of a more intense black which not uncommonly
shows up very conspicuously from the side and behind in the living animal. It is
interesting to note that the greying on the backs of the pinnae shows up most pro-
minently 111 melano animals where it forms the only cemtrast to an otherwise wholly
black pelage. In all cases, except melanos, the extreme edges of the ears are narrowly
white; the front face carries a growth of upwardly directed long white hairs along the
inner margin, and shorter white hairs along the outer margin; and there is a tuft of
similar hairs arismg from the head and obscuring the orifice. The apical pencil may
be pure black or, more usually, have a greater or lesser mixture of white hairs with it.
In older animals when the cars arc not being actively pricked their tips together with
the apical pencils hang down in tassel fashion and waggle when the pinnae are twitched
— as they can be independently.
The tail is remarkably short being merely about a third of the head &; body length,
reaching only to about the hocks when the animal is standing and the tail hanging. It is
subcylindrical, the hairs coming to a slight point at the tip, a little darker above than
below but devoid of black rings or other markings. The legs are moderately long but
relatively stout compared with those of the serval, the hinder ones rather longer
than the front ones, so that the rump stands higher than the shoulders, the whole
hindquarters being somewhat more heavily built than the forequarters. The feet are
abundandy hairy below between the pads but do not carry that dense mat of stiff hairs
characteristic of the sand cat. The interdigital webs are short; the claws retract into
sheaths in the normal cat fashion.
Skull (figs. 54 and 55). This is of roiuided but irregularly curved profile, being
most elevated across the frontals at the uiterorbital constriction, tailing away gradually
posteriorly but sharply anteriorly, the nasals descending abruptly to the short bhmt
rostrum. The frontal region itself is slightly hollowed medially. The nasals are, in
comparison to the most typical Filis, broad, parallel-sided at about their mid-length,
tapering to a short blimt point posteriorly. Attention has already been drawn above
to the narrow and intrusive nature of the nasal branch of the premaxilla which reduces
the length of contact between the nasal and the maxilla to a short, or sometimes
extremely short, distance — though this reduction is not very marked in the type of
pcccihtis, shown in the figure. The posterior part of the sagittal crest, joining the
broad, sharp supraoccipital crest in a T, is always well-developed; anterior of this there
is sometimes a low continuation across the main body of the cranium even in females.
The orbital ring is never complete. The bullae are fiirly large, the exterior chamber
occupying a quarter or less of the whole. The post-dental palate is short and wide;
the mesopterygoid fossa also broad and open, the hamulars curved and sharply pointed,
the external wings of the pterygoids well-developed. The main palate itself is short and
broad, its posterior margin shallow open curves, not narrow notches as in typical Felis.
The mandible is strongly built, the rami deep, sharply upcurved anteriorly, the
FELIS CARACAL 4O7
posterior blade deeply excavated and ridged to accommodate powerful miBclcs. The
coronoid process is tall and narrow; the angular process short and blunt, incurving.
The dentition is powerful. The cheekteeth are ^q, p^ and;'^ being normally lacking,
though the latter may exceptionally be developed. With the loss of these teeth the
jaw has become shortened and the distance between the back of the canine and the
front o{ p^ is unusually short. There is, nevertheless, a clear, if narrow, post-canine gap.
Both upper and lower carnassials are large; p^ has a small anterior as well as a posterior
cusp but it is very often worn away and luidetectable. »ii is not only very small but
transversely orientated and bears against the posterior face of //ii.
Habits. Some observations of the habits of caracals in captivity have been recorded
but not much is postivcly known of their very secretive lives in the wild. Without
question they are solitary animals associating in the adult state only for brief periods
of courtship and mating. Even the nature of their nocturnal or breeding shelters is
not clearly known. For this purpose they certainly take advantage of natural caves,
fissures in rocks or cavities amongst boulders; but relatively little coimtry providing
refuges of this kind exists in West Africa, and though caracals are very good climbers
and no doubt often rest and possibly sleep in trees such locations are wholly unsuited
to raising a family and it seems probable that holes in the groiuid, originally excavated
by other animals, are most commonly resorted to as homes.
It is obvious that small to medium-sized mammals and birds form the staple diet;
rats, gcrbUs, hares, dassies, monkeys, young baboons, francolins, guinea-fowl and
the like. But these very active cats also kill small antelopes and snatch the fawns of
larger ones; Bothnia (1965) records the remains of a Grimm's duiker found in stomach
contents. Caracals also have a reputation for raiding domestic stock and poultry,
and for this reason they are accounted vermin in South Africa. They are such accom-
plished climbers that they are difficult to fence out. Whether these cats regularly take
other kinds of prey than mammals and birds docs not appear to have been clearly obser-
ved in the field; but Shortridgc (1934) mentions that a specimen in an Asiatic zoo killed
and partly devoured a cobra that had entered its cage. As in other carnivores, a certain
amount of vegetable food is taken; Bothma (1965) notes that in South Africa grapes
were found in a stomach, and a small quantity of green grass.
Caracals may be on the move at night but chiefly hunt by day. Two methods of
capturing prey are employed : running it down or leaping upon it by surprise, a method
that must most especially be used, of course, for birds. With regard to the former, the
caracal has some reputation for speed over short distances, though in this it can hardly
be in the same class as the cheetah. Indeed, with its not particularly long, rather sohdly
built legs it is difficult to see how it could establish itself as an outstanding runner;
yet, nevertheless, it was up till recent times tamed and kept by Indian princes for the
hunting and capture of small game, and Vigne (1842: 42) who wimessed such hunts
considered that its speed "was quicker in proportion than that of the chita". The kill
in the case of larger animals is made by a bite in the neck severing the jugular vein.
The caracal's real abihry, however, lies in its leaping capacity, its powerfully built
hindquarters enabling it to make jumps of surprising range. These are most spec-
408 THE CARNIVOHES OF WEST AFRICA
taciilarly performed in tlie killing of birds, the caracal making a standing jump ot as
much as a couple of metres clear into the air to bring down its victim with a clap
between us two front paws. Almost every cietailed account of this animal refers to the
statement attributed to Dlyth, the prime mid-igth century authority on Asiatic
mammals (original reference not traced), that Indian potentates customarily pitted their
tame caracals against one another, wagering on the number of birds that would he
killed by each when the competing animals were allowed to leap into a flock of feeding
pigeons. A skilful caracal could knock down nearly a dozen on the ground and in the
air before the remainder could make their escape. This deliberate act of "sport" with
its resultant fluttering confusion must with little doubt be the true origin of the expres-
sion to "put the cat amongst the pigeons" — a saying that has, however, somewhat
unaccountably come into common use only in very recent years.
The habit of stalking and leaping upon a flock of birds is illustrated by an experience
of the late J. T. Davcy (personal communication). While he was travelling in a
lorry in northern Nigeria in the early morning the driver sighted a flock of guinea-
fowl in the road ahead and, as die custom is in such circumstances, accelerated rapidly
in the hope of killing one or two. At that moment a fully-grown caracal leapt from the
grass at the roadside and seized a bird, with the consequence that both it and the guinea-
fowl were immediately killed by impact with the vehicle. Either the animal's concen-
tration upon its prey must have been so intense as to render it unaware of the clatter
of the approaching lorry, or it failed to connect such a noise with danger. A young
caracal's attitude towards motor-cars was noted by the same observer on another
occasion. When a caracal cub emerging from the bush tust sighted his car parked at the
side of the road it hastily hid behind a tree, from which position it played "peep-bo"
for about ten minutes, repeatedly peering cautiously round the edge of the bole and
withdrawing again until it was satisfied that no danger existed, when it came out into
the open, examined the vehicle and finally trotted off.
Most cats are of an independent nature but the caracal has a stern aloofness of look
and bearing that sets it apart even from the others. There is no doubt that it is not only
very self-suflicient but also of an extremely fierce disposition, becoming a formidable
antagonist if cornered. Some authors, indeed, flatly assert that it is quite untameable;
but though this may often be the case it is not always so; for animals, as humans,
differ in their temperaments and adaptability and, apart from caracals formerly habitu-
ally domesticated in India, there are more modern well-authenticated cases of com-
plete taming (c.i;. Petzsch, 1939). In its natural surroundings the caracal is one of the
most briskly active and nimble of cats, with its shc^rt bursts of high speed sprinting,
its prodigious leaps and its ficile climbing. So powerful is it in the last that, like the
much bigger leopard, it is said by Roberts (195 1) to carry its kill up into the forks of a
tree. There does not appear to be much range of vocalisation beyond alarm notes
consisting of a low growl, that sometimes amounts to little more than a hard outward
breathing, and the usual feline hissing. Kralik (1967) noted, however, that if a male
and female are kept in adjacent cages they communicate with each other by "peculiar
barking signals".
Observations on breeding and the raising of young in captive animals have been
FELIS CARACAL 4O9
recorded by Kralik (1967), Krishnc Gowda (1967) and Cade (1968). Tliese differ
slightly in some of their fuidmgs but present the following general picture. Coupling
according to Kralik is carried out at night, but Cade says the act is performed several
times a day, the male gripping the female m the back of the neck with his teeth. There
is some preliminary play, and coition lasts for about 10 minutes. The period of gestation
has not been determined with great accuracy but appears to be between 69 and 78 days,
the former of these probably being more nearly correct. As the time of parturition
approaches the female cats more and at length prepares a nest for herself of hair and
feathers obtained from her prey; but she does not feed on the final day of her preg-
nancy. The litter may be of from i to 6 cubs but 2 or 3 are the commonest numbers.
The newly-born young are closely similar to the adults in appearance with the normal
sandy or slightly reddish fur, unspotted except perhaps on the belly, black-backed,
tufted ears and the usual facial markings. They arc born blind, the eyes opening on the
9th to loth day — though Cade records that in his litter they began to open on the
4th day and were fully open by the 6th. The cubs start to crawl at the age of about
3 days, begin to learn to walk as soon as their eyes are functional about the 9th day,
and become steady on their feet at about a fortnight from birth. A week later they
regularly leave the nest of their own accord and start learning to chase moving objects;
and when a month old they can run fairly rapidly. However, at this time they are still
suckling though just learning to take solid food. They cat meat regularly at about
i-J months and are weaned about a month later. During babyhood they indulge in
lively play. What happens in the wild state is quite unknown; but in captivity if the
male is near and unrestrained he at once attacks and devours the newly-born cubs;
and unless she has an atmosphere of quiet security free of all disturbance the female
also may eat her young, even up to a couple of weeks after birth (Petzsch, 1939).
Krishne Gowda's pair of adults produced a second litter 7 months after the first; but
Petzsch records an interval between births of just over 3 months. According to the
dates given by Krishne Gowda his caracals bred at the age of little more than 6 months;
but Cade found, from his own and communicated records, that females can be expected
to become sexually mature at about 2 years old. As for longevity. Flower (193 1)
cites a pair of caracals each of which attained an age of between 16 and 17 years in the
Dublin zoo, though the normal expectancy of life in captivity is more usually of the
order of 8 or 9 years.
Taxonomy. There is no question concerning the vahdity of caracal as a species,
it being amply different both externally and cranially to distinguish it clearly at this
level from all other felines. Subspeciation is, however, another matter. The variations
in colour found amongst study specimens have inevitably given rise to the erection
of a number of presumed races; but the relatively few skins that exist from any one
locality give little or no idea of what might be regarded as a normal range of variation
within a single population. This docs not apply solely to West Africa; Ognev (1962),
indeed, drew attention to the fact that the material at present available in collections
was extremely poor and did not permit the drawing of any conclusions. The trouble
starts, as so often, with the proper determination of the nominate race. In this case
there are two contenders for that position: Schrebcr's animal from South Africa and
410 THE CARNIVORES OF WEST AFRICA
Miillcr's from the opposite end of the continent, Algeria {vide the synonymy above).
It is difficult to say, therefore, how or how much any suggested forms differ from that
which might be regarded as the typical race.
Only one race has been ascribed to West Africa, poicilotis. All other named forms
concern either northern or southern Africa except one, nuhicus (J. B. Fischer, 1829)
which, from its name alone, must be assumed to have its origin in or near the Nubian
Desert, Sudan. Fischer's diagnosis is without any value whatsoever. As far as West
African specimens are concerned there appear to be, from the 7 skins available, two
extremes of colour, at one end very red, at the other only slightly so, these being
basically more buffy-grey, less warm in tone. The two categories, however, do not
appear to relate to particular vegetation zones. Considering the greyer material first,
this includes the rj-pe oi poccilotis from Moiuit Baguezan, in the Subdesert, and a skin
reputedly from Kode, 24 km north-west of Lau on the Benue (Nigeria), situated in the
Sudan zone, but from its history and appearance quite possibly bought in a local
market. This, a poor specimen, is very little darker or different from Thomas & Hin-
ton's "desert-coloured form", the ears perhaps a shade less silvered. Tliere is also
one of two skins purchased in Kano market (No. 71.758) which, though belonging to
this category, is vers' noticeably darker than the others, appreciably different, in fact,
from any of the rest by reason of a plentiful admixture in the pelage of bristles with
much longer black tips, giving a deeper greying to the basic buff. The blackening is
extended onto the upper surface of the tail. It resembles skins not only from Sudan
but from Ethiopia and Algeria as well; and though it might be local it could just as
easily have travelled across the desert by one of the many camel trains which regularly
come to Kano market.
The remaining 4 skins of warmer tone come from Gambia (locality unspecified but
possibly Guinea woodland), Gambaga (Ghana, Sudan woodland), Lake Chad (ex-Zoo,
almost certainly Sahel woodland) and a second skin from Kano market (No. 71-757.
real provenance uncertain). These are all fiuidamentally of similar colouring, con-
trasting in their red tone, to a greater or lesser degree, with those dealt with in the
previous paragraph, being, moreover, essentially dit^erent from anything else in the
collection. They vary amongst themselves both 111 the intensity of their redness and
in the amount of "frosting" on the coat brought about by white subterminal regions
to the bristle-hairs. The least intense in its redness is the Kano skin; the Gambia skin is
several shades redder and also has the greatest amount of frosting. The remaining two
skins are both of a richly warm colouring, differing from each other only in the some-
what greater frosting to the Lake Chad pelage.
The 7 West African skins all differ from each other in some degree. At the very
minimum it would seem admissable to allot them to four different forms. But 111 view
of the poor quality' and incompleteness of the material available, the doubtful proven-
ance of most of the specimens and our virtual ignorance of normal population variance
the drawback of adding further burdens and uncertainties to the nomenclature seem
to outweigh any possible slight advantage. Nominal differentiation of these putative
races is therefore postponed till further, better documented and more complete collec-
FELIS CARACAL 4.II
ting has adequately demonstrated that the creation of new names is justifiable and
beneficial.
Table 25 : Numerical data for Fclis caracal
poecilotis, Nigeria
type
Subdesert
specimen
Vegetation
?
Number in mean
I
I
Condylobasal length
105-7
113-0
Basilar length
94-3
I0I-6
Palatilar length
43-4
42-8
Zygomatic breadth
8i-4
83-6
Upper cheekteeth breadth
49-4
50-5
Nasals, length
33-8
—
Interorbita breadth
22-0
23-5
Postorbital constriction
30-0
27-7
Braincase breadth
49-9
50-3
Toothrow (c — m^)
3ri
39-4
p* length
I5-I
H-4
ml breadth
4*2
(4-6)
Hil length
II-5
—
Head & body
594
—
Tail
2S4
—
Hindfoot
ISS
—
Ear
76
—
KATIOS (per cent)
Tail/head & body
43
—
Zygom. br./condylob. 1.
77
74
Braincase/condylob. 1.
47
45
Braincase/rygom. br.
61
60
Palatilar l./condylob. 1.
41
38
Interorb./postorb.
73
85
p^lc—in^
407
36-6
Subgenus LEPTAILURUS Scvertzov, 1858
Serval Cats
Taxonomy. This is another subgenus most often now considered to be monospecific
though at one time it was commonly held to comprise two species. This latter view
was due to there being two apparently distinct pattern forms, one, the typical serval
cat {serval Schreber), in which the black spots are relatively large and relatively few,
and the other, commonly called the servaline cat {brachyurci Wagner, sert'aliua Ogilby)
in which the maculation consists of innumerable very small spots. DisbeUef in the
specific disunity of the two forms was expressed by Elliot as long ago as 1883; but the
controversy really centres round Pocock who in 1907, having gone into the matter
at some length, at first accepted that the two forms represented separate species. Ten
412 THE CARNIVORES OF WEST AFRICA
years later (1917b) he had become convinced that tliis was not so. This change of view
was brought about by a statement made at a meeting of the Zoological Society of
London in 1915 [I'iJc Proceedings: 154) to the etiect that in Sierra Leone kittens of the
two diverse forms had been brought by a local African to the speaker "almost certainly
from the same litter", thus showing the pattern to be dimorphic in a single species.
J. A. Allen (1924) was, with justification, somewhat scathing about the scientific value
of this "evidence", which was in f\ct nothing more than a supposition. Nevertheless,
Pocock retained the same monospecific standpoint, and in what was practically his
last paper dealing with the Felidac (1944: 696 fn.) asserted that since he had first
expressed his conviction of the two forms being but a single species "considerably
more evidence in confirmation of that conclusion has come to hand", though he
omitted to indicate what this evidence was. It must, however, have been very largely
that contained in Pitman (1934) where a specially assembled collection oi^ Lcpttuhirus
skins is described as showing "every stage of variation in size of spots betAveen the
large spots hitherto considered as typical of svrviil scrval, and the (almost) pin spot
Cj'pical of scnuil scrvdlind". Other, less important, notices of the wide variability of
maculation have also been published and the synonymy of the two reputed species is
now generally imquestioned. The incontrovertible evidence indicated by Pocock
himself (1907) as desirable "the occurrence of the two types in the same litter of kittens
known to be the progeny of parents resembling each other in pattern" is, however,
still not forthcoming.
General description. Lipttiihinis is clearly distinct from all other African members
of the genus Ftlii and is only possibly to be confused by the inexperienced with the
cheetah [Acinonyx) because both, typically, have a pattern of single, smallish, jet-black
spots on a sandy ground. It is this coloration and pattern that immediately distinguishes
the scrval from the other West African felines, the wild cat, the sand cat, the caracal
and the golden cat. Other quite characteristic features arc found in the very prominent
cars. These are very tall and large, oval, not triangular, in shape with broadly roimded
apices, and sited unusually close together at the top of the head: and on their backs their
distal half is black enclosnig a somewhat elongated white patch. The head is small for
the general size of the animal, the legs exceptionally long, and the tail short, only a
quarter to a third of the head & body measurement. The skull is very similar to that
of Ciiracil including, in a number of instances, the long, narrow intrusion of the
premaxilla between the maxilla and the nasal. The bullae are generally rather larger;
but the most immediately observable difference between the two skulls is the presence
in Lcpliuhinis of a small anterior premolar, in the upper jaw very largely filling the
space between the canine and ]'^.
Distribution. The subgenus is solely African in its distribution, ranging from Algeria
to southern Africa, now to only about 30'S. though once much nearer the Cape.
FELIS SERVAL Schreber Serval Cat
Felis scri'al Schreber, 1776, Die Sdu^cthicrc in Ahbih]uii(;i-ii . . ., pi. loS; 1777, text 3: 407 and 587. Cape of
Good Hope. This name is supposed to be derived from tlie Portuguese lolh^-ceri'al, a lynx.
Ft'lis capvnsis j. R. Forster, 17S1, Phil. Trans. R. Soc. 71: 4, pi. I. Cape ot Good Hope.
Felis gdlcoytirthis Desniarest, 1820, Encyclopalie McdioJiqiic . . . Manimalo^ie, I: 227. No type locality.
FELIS SERVAL 413
This was based on "Le Serval" of F. Cuvier(i8i8); J. A. Allen (1924: 269) found it to be indeterminable
and hence unavailable either as a valid species or as a prior name for saicgalettsis Lesson, as has some-
times been argued. The term was coined from gale the Greek for a weasel, and the Latin parilus leopard,
referring to its similarly spotted coat but relatively insignificant size.
Felis scnegok-nsis Lesson, 1839, Magasin Zoo]. Paris {2), classe i Mammiferes, I: pi. 10 and two pages of
le.xt. Banks of the River Senegal. Preoccupied by Felis leo seiiegaletisis Meyer, 1826.
Felis servalitta Ogilby, 1839, Proc. zool. Soc. Loud. : 94. Sierra Leone. Preoccupied by Felis tcrrfl/iiia Jardine,
1834, = Felis libyca omata Gray, 1830, [vide Cabrera, 1910: 426). This name is a dimunitive of serval
given with reference to the smaller spots.
Felis brachyma]. A. Wagner, 1841, in Schreber's Die Saugethierc, Supplenientband 2: 547. A substitute
for Felis servalina Ogilby preoccupied by F. sc rra/i'im Jardine. The derivation of this name is from the
Greek words hrachys short and oiira tail.
Felis ogilhyi Schinz, 1844, Synopsis Mammaliiim, 1 : 469. Sierra Leone. A renaming of Felis servalina Ogilby,
1839, preoccupied.
Felis serval pococki Cabrera, 1910, Bohi R. Soc. csp. Hist. nai. lO: 427. Senegal. A replacement for Felis
senegalensis Lesson, preoccupied. This was called after R. L Pocock who had first demonstrated the
true position of Lesson's naming.
Felis {Serval) togoensis Matchie, 1893, Sher. Ges. natnrf. Frennde Bcrl.: 109-110. Bismarckburg, Togo.
Distribution and general. Since there is only a single species the distribution of
the serval is that already broadly indicated for the genus, that is to say, it is whoUy
African, ranging in the north from some of the Mediterranean countries to southern
Africa, where it once occurred not far from Capetown itself but has now been forced
to withdraw much further north. At the other extreme of its range it is also losing
groimd; but it may still be encoimtercd in the northern parts of Tunisia (Gouttenoire,
1954), and it was, at least until recently, to be found in Algeria. The species is known
to exist in all West African territories, inhabiting all zones from the Sahel to the closed
forest, being recorded even from a coastal island; and it ascends to moderate altitudes
into the mixture of forest, open woodland and pure grass characteristic of hilly parts
of the region at about 1000-1500 metres. The serval appears to avoid the desert;
indeed, it demands two conditions, dense cover and fair proximity to water. This
means that while it may be at home almost anywhere in the forest or Guinea zones,
in the drier sorts of grass-woodland, the Sudan and the Sahel, it fmds suitably dense
vegetation only near the banks of rivers or marshes where there is sufficient tellurian
moisture to support lusher growth. Nevertheless, as this is a predominantly nocturnal
animal it will, of course, in order to hiuit emerge into scantier vegetation under cover
of dusk or darkness.
The serval is a not uncommon animal; there are, in fact, 34 West African skins and
12 skulls in the British Museum as contrasted with only 7 caracal specimens. Yet as it is
only exceptionally on the move during the day, and then mostly skulkuig in luider-
growth, it is very rarely seen save in car headlights or by night himters with powerful
lamps. Of the British Museum material, 9 specimens come from Senegal, including
Thies and Bakel; 9 from Sierra Leone, including Makali, Makeni, Karina, Nerekoro,
Bonthe, Gberia Timbako and the River Moa; a single skin from Liberia, 80 km inland
of Monrovia; 5 from Ghana, including Ejura and a site 48 km west ofjuaso; a skin
from an unnamed locahrj' ui Togo ; 8 specimens, but with only 2. skulls, from Nigeria
between nearly the coast and Lake Chad — Gombe River, Zo 32 km north-east of
414 THE CARNIVORES OF WEST AFRICA
Lau, 48 km cast of Bauchi, 20 km south-west of Jaliiigo, Pctico 24 km north-west of
Lau, near Sapclc, and an unspecified locality at Lake Chad (ex-Zoo) ; two examples
from upper Camerouii at Njawbaw and Olulu both situated in Assumbo north of
Mamfc; and one skin from Fort Laiiiy. It is also reliably reported (in personal com-
munications) by A. J. Hopson as occasional near Lake Chad, one having been killed
in a compound at Malamfatori; by the late J. T. Davcy as common in Macina (Mah);
and G. S. Child as occurring in the Borgu Game Reserve (west Nigeria). Published
records are: the environs of Timbuctu (Zimara, 1935); and three places in Portuguese
Guinea, Catio (Frade, 1949), Contubo-el and Pitche (Moiiard, 1940). This is an extremely
wide range and varied habitat, from mangrove and coastal scrub (Bonthe), or dense
high forest (Sapelc), to widely open Sahel-type woodland not very far from the Sub-
desert though in swampy conditions near the Niger or a lake. However, the position
with this apparently abundant London study material is not very satisfactory since
only 6 skins arc matched with adult skulls; and not a single skin carries any field
measurement — a point held in common with the vast majority of British Museum
specimens from anywhere m Africa, there being, all told, only some 3 or 4 bearing
such data. A very high percentage, in fact, appear to have been purchased from African
hiuiters. Only 7 specimens have the sex indicated; and a quarter have no precise locality,
and, indeed, even the broadly named provenance of some of the older material is
suspect. Such of the British Museum specimens as can be determined show that 3
specimens come from the Sahel zone, 5 from the Sudan, 1 1 from the Guinea, 6 from
the forest, I from mangrove and scrub, and I from mountain forest and grass.
Description. Because of the existence of very diverse types of maculation in this
species the account which follows deals initially with points of morphology and those
few markings which are completely common to the various forms, leaving description
of the pelage and its distinctive patterns to the end. The serval (Plate 11) is a much
more elegant cat than the caracal being more slimly built, narrow from side to side,
and with longer far more slender limbs, which differ from the other species also in
that the forelegs are longer than the hindlegs. Although it is of the same order of
size as amiail — both being in the category of "medium-sized cats" — and actually
stands higher at the shoulder, that is to say between 500 and 550 mm as compared with
400 to 450 mm, its weight, at 10 to 13-5 kg, tends to average out at rather less. Like
caracal, serval is somewhat more heavily built in the hindquarters than in front, and the
long slender neck and disproportionately small head serve to exaggerate this inbalance.
In comportment the serval is a fme upstanding, alert cat, not one given to cautious
slinking; and even when it sits up on its hindquarters it holds itself erect and looks
very tall. In this position the two parallel black bars on the inside of the upper end of
each foreleg are very conspicuous; and, also from the front, a narrow black line or
row of spots can be seen forming a "necklace" across the lowest part of the throat —
though this is mostly very obscure in the small-spotted forms.
The front view of the serval, however, is dominated by the remarkable upstanding
cars which are so large that they measure from base to summit almost as much as the
distance from the base to the large rhinarium. Apart from their size they have several
other notable features. They are set unusually high on the head and when actively
Scrval (hchs scrfalj: Atric.in C.oldcn C.it (l-flis mmua)
FELIS SERVAL 415
pricked arc brought exceptionally close together. They are ovoid shell-like in form,
the pinnae being deeply concave, broad at the base with the margins rising with very
gradual narrowing to a broad arc at the apex. There may be a slight apical fringe but
no tuft. The front aspect carries a dense cover of white hairs; the back is quite charac-
teristic with a black distal half wholly or partially transversely divided by a broad pure
white patch. This very noticeable mark forms one of the most immediately con-
spicuous features of the living animal seen from the rear or the side. Facial markings
consist of a dark patch at the inner corner of the eye reaching some way towards the
rhinarium; and a solid or broken line, generaUy much fainter, from the top of each
eye to the crown. There is a broad white streak under the eye, and a shorter white
patch above towards the inner angle. There is an area of white, sometimes considerable,
to the side of the rhinarium continuing narrowly along the upper lip and spreading
somewhat onto the check posteriorly. The lower lips and chin are wholly white.
Irrespective of the size of maculation on the back and flanks the outsides of the upper
parts of both fore and hindlimbs are prominently marked with large black spots,
which abruptly diminish into small black spots below. The inner faces of the limbs are
white, mostly black-spotted, the foreleg with two very conspicuous black bars. The
soles of the feet are densely clad between the pads with longish, deep blackish-brown
hairs; the claws are fully retractile, that of the pollex being appreciably larger than the
rest. The interdigital webs on the forefeet are fairly extensive reaching to about the
bases of the claws. On the hindfeet they are somewhat narrower. The serval is one of
the short-tailed cats, the tail measuring little more than a third of the head & body
length. It is of the common feline subcylindrical form, of the basic pelage colour with
black rings, or partial rings since they do not always join below. There are generally
about 6 or 7 of these, and the tip is always black. There is also, very clear in some
specimens, obscure in others, a black Une, continuous or interrupted, running down the
upper side of the tail to about the second ring from the end. In the eye, the iris is
golden, the round pupil contracts to a broad upright oval.
Turning now to the pelage, this is fairly soft and of moderate length, its composition
of the usual three elements: very abundant, long, fme underfur, mostly very pale buff
with coloured tips ; stoutish bristle-hairs of terete section, either all black or black-
tipped with or without a subapical band of gold; and sub-bristles with a fine petiole
and expanded flat-sectioned distal region which is mostly golden-yellow with a black
tip. The lengths of these components vary with different specimens but generally
fall within the following hmits: underfur 20 to 26 mm; bristle-hairs 27 to 31 mm; sub-
bristles 27 to 35 mm, the flat blade measuring some 7 or 8 mm. The black spots are
composed of similar elements which from pale bases become distaUy progressively
darker, passing through sepia to black.
In large-spotted, "serval", specimens there is a good deal of variation in pelage
colour in animals from different areas or sometimes from the same or closely proximate
localities. The general ground-colour is mostly some shade of sandy-buff, a httle
lighter, a little darker; but skins from the closed forest exhibit a much richer red colour.
Whatever the colour it is almost invariably a trifle more intense dowTi the spine, and
usually a little paler on the flanks. On this ground-colour is imposed, in the "serval"
4l6 THE CARNIVORES OF WEST AFRICA
form, a bold black pattern of continuous linos and independent spots, variations in the
size, shape and number of both of which iiave been the source of siibspecific naming.
The longitudinal pattern of continuous, or almost continuous, lines follows the
following basic scheme. It starts at the back of the head as four slender lines; these all
fin out over the back of the neck, the two inner ones remaining fine, the two outer
becoming rather broader and bolder. At the base of the neck the divergence continues
rather more sharply, the inner pair of lines becoming as bold as the outer. The latter
curve round the shoulders toward the flanks where they peter out as continuous
stripes and become the flank spots; the middle pair also break up into the innermost
row of dorsal spots at about the same level. Before they do so a third pair of narrow
lines appears between them, sometimes with the faiiit indication of a central spinal
line, and these after slight initial divergence continue back in sub-parallel fishion and
often slightly broken almost to the root of the tail, with the flint medial line between
them soon becoming equally pronounced and sometimes itself closely dividing into
two posteriorly. Over the main part of the body, cither side of the medial striation,
is a pattern of roundish, oval or oblong spots not very precisely disposed in longitudinal
lines. So fir as they can with any accuracy be determined there are at the posterior end
of the body commonly about 6 such rows either side ot the spinal lines; but there may
be one more or less. All this pattern, in the matter of continuity of the stripes and,
most especially, the size, shape and number of spots, is subject to v^nde variation,
though us main features are always clearly observable. As for the underside, the chin
is white with a chain of small spots across it posteriorly; the throat huffish, mostly
also with a chain of spots, sometimes confluent, across it; the chest and belly white,
or to a greater or less extent suffused with buff", well-covered with long and loose hair,
and mostly, but not invariably, marked with rather obscure dark spots.
In the small-spotted, "servahne", forms the whole dorsal ground-colour becomes
dulled so that there is little or nothing of the bright buff"y or sandy hue of the r)'pical
scrval. This seems to be due to a more general spread of dark pigment, which instead
of being intensely concentrated in small areas, giving black spots on a pale ground,
diffuses throughout a high proportion of the hairs of the back so that the underfur
together with the petioles of the sub-bnstles become a dull sepia instead of whitish or
pale buff. Extralimitally, the fur may even assume a grey tone rather than huffish. The
size of the black spots is very much reduced and their number vastly increased; but
in at least one known example the reduction is carried to the extreme of complete
disppcarancc of any dorsal maculation whatsoever. In all cases any trace of the dis-
tinctive pattern of^ continuous lines so characteristic of the large-spotted animals
completely, or almost completely, vanishes. The luiderparts, however, together with
both inner and outer aspects of the hmbs remain much the same in both phases.
This slight suffusion of the dorsal pelage with dark pigment is a long way from
intense general melanism — which, as in other felines, is not uncommon in this species
too, though no specimen has so fir been recorded from West Africa.
Skull (figs. 56 and 57). In size and overall appearance this is very similar to that of
the caracal, though the profde does not descend so abruptly over the nasals. Com-
parison ill this lateral aspect (e.g. in figs. 54 and 56) clearly shows that the zygomatic
FELIS SERVAL 4I7
arch is considerably flatter in its curvature; that the suture between the jugal and the
maxillary process is more nearly parallel to the line of the teeth instead of ascending
sharply; and that the anterior edge of the latter bone projects further forward in the
serval and overhangs the infraorbital foramen. The prcmaxilla intrudes narrowly
between the maxilla and the nasal as it does in canicnl, but on the average not quite so
far as in that species though in some young skulls it nearly reaches the long forward
angle of the frontal. The sagittal crest is as in CunKdl, developed only posteriorly;
but differences to be observed m the dorsal aspect are that the serval skull is narrower
in that the zygomatic width is relatively rather less; the difference between the inter-
orbital and postorbital widths is more marked, the former being somewhat less than
in caracal, the latter appreciably more, this last according the whole braincase a rather
larger appearance. One Sierra Leone skull (No. 50.2042 from the coastal scrub of
Sherbro Island) affords a striking exception to the general run in this character: in it
the postorbital processes are much longer than usual, there are well-developed post-
orbital ridges that markedly overhang a postorbital constriction that measures only
no per cent of the interorbital width instead of the usual 160 per cent. This skull is
in all other respects typical and is accompanied by a normal large-spotted serval skin.
Li the palatal view the bullae in scrral, though somewhat variable in size in different
specimens, are in general clearly larger than in caracal; the mesopterygoid fossa is
markedly wider — 14 mm or more as compared with 12 mm; and the lateral wings of
the pterygoids are broader, hi the mandible the coronoid process is not so tall; the
condylar process less strongly built; the rami are not quite so deep, the jaw thus giving
the impression of being rather less powerful. Anteriorly, the rami do not curve up
quite so steeply; and this together with the lesser angle of descent of the nasals brings
about a rather less blunt rostrum.
The cheekteeth formula is ^i there being 3 premolars on each side of the upper
jaw instead of 2 as in caracal; 30 skulls examined furnished only a single exception to
this, one young skull having the anterior premolar lacking on one side. The carnassials
both above and below are noticeably smaller in serval than in caracal and aurata, and the
toothrow, in spite of the extra premolar, is shorter; p^ has both a posterior and anterior
cusp, the latter mostly quite distinct but sometimes rather obscure. Most of these points
of comparison of the skulls of the species are brought out in the tables on
pages 411, 423 and 436, in so far as it is possible to judge from such hmited data
as the few available specimens provide.
Habits. The serval is one of those animals which are rarely seen and whose habits
have in a large measure been deduced from occasional and sometimes fleeting observa-
tions. This is mainly due to the species being nocturnally active; and even if these
cats are awake and alert during daylight they are more often than not hidden from
observation by the density of the undergrowth in which they take cover. Even though
they may possibly start their nightly roiuid of foraging before darkness has properly
set in they rarely seem to emerge into really open coimtry where they could be easily
seen. The difficult task of making a set field study of the life of these secretive animals
in the wild has not yet been attempted; and though servals have commonly been
kept in captivity very little of a detailed nature has been pubhshed relating to them.
4lS
THE CARNIVORES OF WEST AFRICA
The precise nature of the ground vegetation in which these cats find shehcr is
immaterial, whether it be forest imdergrowth, grassy tangle or palustrine reeds; but all
observers are agreed that the proximity of water seems a desirable factor; and it is of
course true that it is near water that the ground cover is generally most dense, especially
in the drier vegetation belts. If necessary, the serval shows itself to be an efficient climber
but it is predominantly a terrestrial animal. For breeding purposes it resorts to holes,
not made by itself since its digging powers are nil but those originally excavated by
aardvarks or porcupines, or at the bases of termitaries, or in hollow logs, or amongst
Fig. 56. Felis serval: skull, B.M. No. 99.10.23.3, ^, x i; lateral view
rocks. Whether such places as these are used at other times as ordinary day to day
refuges or whether these animals are normally content to sleep curled up 111 ground
vegetation has not been determined.
Since hunting or at least the greater part of it is carried out at night good nocturnal
vision IS indicated as in most cats; but from the enormous external ears backed by
unusually large bullae it may be inferred that hearing also plays a prominent role in
this animal's activities. The prey sought after is that common to all these small and
medium sized cats : rodents of all kinds, including the larger ones such as the cutting-
grass [Thryoiiomys) and the giant rat (Criatomys), squirrels and hares. The smaller
FELIS SERVAL
419
Fig. 57. Felis servai: skull, B.M. No. 99.10.23.3, t only short but stout and strong; the round heat! appears rather small, as in the other
two cats, but ditters from them most markedly in the very short ears which have a
height of only about <,<, mm as contrasted with 75 to 85 mm in the other species. Their
outline is rather roundly triangular with a blunt tip which has no tuft; and they are
almost wholly shiny black on the back except for a greyish patch at the lower outside
edge.
There are small white nr whitish patches .ibove the eyes, especially at the inner
FELIS AURATA 43 I
corners; and the lower part of the checks is also white; but there are otherwise no very
obvious facial markings. The iris is brown ; the pupil upright spindle-shaped. The tail,
which is soft-haired, smoothly subcylindrical or slightly tapering in form, has a dark
dorsal line throughout its length and a number of more or less obscure transverse,
similarly coloured bands, sometimes very ill-defmed. The extreme tip, when present,
is dark. The structure possesses the typical felmc mobihty. The feet are densely long-
haired between the pads; the interdigital webs broad; the powerful claws fully retractile
into sheaths.
Whatever the colour and marking of the dorsum the pelage always exliibits certain
pretty constant characters. The underparts are in a varying degree white or whitish:
the chin and throat are mostly pure white, but at the anterior end of the latter are often
to be detected one or more famt transverse chains of very small spots; the lower throat
may be white or more often suffused with ground-colour and spotted; the chest and at
least the medial part of the belly white, the sides of this last frequently being suffused
with colour; and the whole chest and belly is always heavily spotted with large blotches.
On the dorsal surface, throughout all the various forms, the coat always displays a very
distinct and characteristic reversal of direction between the shoulders and the crown, the
hair in this area pointing forwards. The change is marked at shoulder level either by a
single medial whorl or two lateral whorls; and by low crest-hke ridges along the sides
of the neck and on the crown just anterior to the ears where opposingly directed hairs
meet.
The dorsal area is subject to so many variations of colour and pattern that it is
pointless to attempt to give a detailed description of any particular form or forms.
Pelage texture, too, is inconstant. The coat may be either short, close-lying and rather
harsh, or fiirly long, loose and soft; and there are no data reliably relating such differ-
ences to any particular factor, whether age, season or climate. The fur is composed of
the three usual elements, undcrfur, bristle-hairs and sub-bristles. The underfur is always
dense and very fme; in some cases it is wholly concealed by the abundant bristles, in
others only partly so. The bristle-hairs are of slightly oval section and broad to the base;
the sub-bristles have long fme petioles. When the fur is turned back it is seen that the
coat colour resides mostly in its distal part, the base being pallid or even pure white.
The underfur is usually either very pale or wholly white but may occasionally have
faintly coloured tips. The bristles are always dark-brown tipped: in reddish skins this
apical zone may occupy only some 3 to 4 mm and be succeeded proximally by a narrow
orange zone and a faintly browii ring, the rest being white; in sepia-grey skins the
terminal zone may measure 9 to 11 mm, the whole of the remainder being white. How
variable coat length is, is illustrated by the fact that in some animals the undcrfur is only
10 to 12 mm long, the bristle-hairs 15 to 16 mm; while in others the underfur is 16 to
18 mm and the bristle-hairs 22 to 23 mm.
The overall colour ranges from a bright marmaladc-orange-red to a deep dusky
sepia-grey; but there are many intermediate shades both of red and grey, and it is
sometimes ditlicult to say to which basic colour category a skin belongs. Some of the
greys are cold, some tinged with a warmer hue. The dorsal pattern is also highly
variable, not only as regards its presence or absence but in the size and character of the
4.U
Tilt CARNIVORtS 1)1 WHST AIKICA
mai'ul.inon as well. In the most rv'pical West African form — the subspecies cilidoi^aslcr
according to van Mensch & van Bree — the dark spots are mostly large, roundish and
clear, whether the general colour be red or grey: but they may be smaller or of
elongated shape and sometimes rather obscure though unquestionably present. Largish
flank spots are always clearly observable. Over the back of the neck, in the area of
reversed pelage, the spots become very narrow and long, medially often joined into
more or less luiinterrupted lines. These stripes are not always clearly separated but are
Fig. jX. I-clis minva: skuil, B.M. No. 25.10.7.13, j.
latcra
appro.ximately four in number, apart from spots lateral to them. Between the shoulders
and the root of the tail there is a dark spinal band; and through this, in the majority of
cases, about three of these slender nuchal, yet darker, stripes extend backwards as fir as
the rump. In the Cameroun, central and east African form — reputedly the nominate
race iiiirata — there is no pureh' dorsal or nuehal patttrn of spots, but the dark spinal
band e.xists. Flank spots may or may not occur; but the belly and other underparts are
always spotted as described earlier.
Melanos are known, in West Africa as well as e.xtralimitally. In black forms of all cats
there is sometimes some difficulty, owing to obscurement of the pattern, m determining
the species; aiiratii presents du immediate means of reeognituin in the reversal of pelage
direction on the back of the neck.
FELIS AURATA
433
Fig. 59. Felis amala: skull, B.M. No. 25.10.7.13, ^, x ^^,; palatal &' dorsal views
434
THK CARNIVORIS ()!■ WIST A 1 RICA
Skull (tigs. s!< -iiul sy). Tins IS wn' likr tli.U ot scrr,'.! .uul thcrctorc not rcadilv
disnnguisliahlc tlKuigli on the wliolc slightly larger. Van Mensch i!\; van Brcc (1969),
who cxannnLti 46 skulls from all over Atnca, cite a niuiiber of characters serving to
diftereiuiatc between the two species; six of these are cranial or dental measurements
expressed as percentages of the condylobasal length: but the differences mostly seem to
be too slight to be meaningtul and in some cases rim counter to those emerging from
this present investigation of West African material. From the very wide size ranges
given it would seem likely that they must have included immature skulls, the propor-
tions ot which notoriously differ from those fully adult. It also seems to the present
author from an examination of all the British Museum African specimens concerned
that some of the characters from time to time quoted by .luthors to differentiate between
subgenera of [■clis are not constant and therefore unreliable — such as the shape of die
T.iblc 27: Conipai.itivc data tor /■. taiiual. scrval and aurata
Interorb./Postorb. widi
Maxillary process
Infraorbital foramen
Jugo-maxillary sutiUL'
Depth between jugo-
ma.xillary sutnrc and
alveolar line at the level
ol the toramen
Length of maxillary
process from the foramen
to its posterior edge
caracal sfn'al
'■ 75-S5 per cent (>$ per cent
only slightly overhangs sharply overhangs the
the infraorbital toramen
foramen
medium; broadly
elliptical; max. diam.
5-6 mm
aurata
c. So-90 per cent
sharply overhangs the
foramen
rises very steeply
anteriorly
usually not more than
14 mm
small, rather straight- large, more narrowly
sided and oblong; max. elliptical; max. diam.
usually well over 5 mm,
mostly r. 6*7 mm
diam. 4'0-4-5 mm
rises only a little
general
less
Iv 1 8 mm or
I i-i;
iS-io mm
lacking
small anterior ctisp.
3-4 mm frcmt to back
small anterior cusp
sometimes worn awa\'
the postdental iialatal extension.
rises very steeply
anteriorly
c. irt mm
20 mm or more
a mere peg, 2 mm
no anterior cuspi,
(or ver\' small)
and
the exterior wings o
f the
posterior margin ot
pterygoids.
In profile the cranium curves evenly piosterior ot the pc^istorbital pirocesses whereas
in other species it mostly dips slightly at about the level of the fronto-parietal suture,
especially in older skulls. There is in ProjcUs often, but not always, a fiirly well developed
sagittal crest extending from the postorbital constriction to the broad siipraoccipital
crest, at any rate in the males, there being no females in the British Museum by which
to judge this character in that sex. However, one old male as well as one medium-aged
male show no sign of a crest except posteriorly. The orbital ring is incomplete. The
zygomatic arch is of flattish curvature ns in scnuil; the jugo-maxillary suture first dips
down torming a distinct angle and then rises sharpily anteriorly, the height of the
maxillary process above the teeth being markedly greater dian in the two other genera;
FELIS AURATA 435
.ind it IS also longer from the foramen to the posterior edge. As in Liprailiiriis this process
sharply overliangs the infraorbital foramen, which is largest in this genus with a long
diameter usually of 20 mm or more. The prcmaxilla intrudes narrowly between the
nasal and maxilla but almost invariably leaves a long margin of contact between these
two latter bones. The nasals are fairly broad, often more or less parallel-sided posterior
to their suture with the premaxillary or tapering only slightly, their junction with the
frontals bluntly triangular or even rounded.
The bullae are fairly large to large; the mesopterygoid fossa relatively narrow (about
12-13 nim), its anterior margin generally an even elliptical curve; the lateral wings of
the pterygoids broad as in the scrval. The anterior end of the mandible is appreciably
less abruptly upturned than in ainicnl, the gap between the canine and fust premolar
longer; posteriorly it is deeply excavated externally, leaving a broad flat shelf for the
accommodation of muscle. Li this it differs quite clearly from scn>al and also, though
less markedly, from caracal. There are 3 premolars in the upper jaw, the anterior one
being a mere peg, of 2 mm or less from front to back.
Table 27 gives a comparison of various characters in fully mature skulls of caracal,
scrval and anrata.
Habits. Of this animal next to nothing is certainly known and not a great deal can
be inferred. Li the wild it leads a highly secretive and obscure existence. It has already
been pointed out that a very large proportion of museum specimens have been
purchased from African hunters or traders, and that few, if any, competent field
naturalists can ever have seen this cat alive in its natural surroundings. It is true that a
few golden cats, probably less than half-a-dozen in all, have been taken alive and
eventually exhibited in zoos; but this tells us almost nothing of their daily way of life,
and, moreover, regrettably little of the habits of these few 111 captivity has been
recorded.
It would seem that the golden cat not only secretes itself 111 dense forest growth but
is also very strictly nocturnal so that the chances of ever seeing it are slender, the more
so as there is no doubt that its colour and pattern, whatever form these may take, are
effectively cryptic. This difficult^' applies as much to African hunters as to European
collectors, and it seems likely that the majority of these animals killed have either been
trapped, poisoned or shot with the aid of head lamps — seen only as glowing eyes in the
dark. Only a single field note refers to a living animal having been seen at all, one of
Bates's specimens having been chased up a tree by a dog and shot there; and even this
is not explicit as to whether it was seen by the collector himself or by one of his hiuiters;
or whether the sighting was by daylight or by lamp at night.
It is not, indeed, clear whether this is primarily a terrestiral or arboreal cat. Its short,
powerful legs are ideally suited to climbing, the very reverse of the terrestrial serval's
long, slender limbs. Basilio (1962) professes a greater knowledge of this animal's habits
in the wild than any other author. He says that it spends its day hidden in the lower
branches of shady trees, and at night sets out to capture its favourite prey, birds such as
Guinea-fowl and francohns which also sleep in the lower branches, as well as small
mammals from rodents to little antelopes. This may well be so. Blonk (1965), who
recently kept a golden cat m captivity, found that, although it was not fastidious about
436
THE CARNIVORES OE WEST MRICA
food, It ate birds LMgcrly, alwavs plucking thcni first, however. A point which seems
to favour a largely arboreal life is the tact that nobody seems ever to have found a
breeding shelter in this species; for, unlike most other animals, there are no juvenile
specimens in the collection; and T. S. Jones, in a wide and varied field experience, has
never been brought kittens by African himters. One would suppose that if the nesting
refuge were a hole in the groiuid or other commonK- used terrestrial site it would from
time to tune have been come across, as in the case of other ground-breeding mammals.
On the other hand, if it is hidden high in trees, and if the kittens never descend to the
ground until they are well-grown this would accoiuu for the lack of juvenile specimens.
Accounts of this animal's behaviour in captivity are limited. Eliot (1873) recorded that
a specimen cxliibited in the London Zoo could be handled by its keeper even while
feeding, Blonk (1965), on the other hand, though characterizing his animal as relatively
tame found that it would not ever allow itself to be touched. This latter author also
noted that no attempt was made by the cat to cover up its droppings in the manner o{
Table 28 : Numcnc.1l data tor Ft/is mimtii
West
Lower
Africa
Cameroun,
etc.
Vegetation
Forest
Forest
Number in mean
4
3
Condylobasal length
115-4
126-0
Basilar length
104-4
IIJ-X
Paiatilar length
46-5
50-3
Zygomatic breadth
86-7
90-3
Upper cheekteeth breadth
5I-I
49-8
Nasals, length
33-8
35-6
Intcrorhital breadth
22-9
24-2
Postorbital constriction
27-7
27-9
Braincase breadth
50-6
5;i-7
Toothrow {i — H|l)
39-5
43-1
p'^ length
16-2
i6-o
tn^ breadth
4-8
5-6
nil length
12-1
12-2
Head {<.■ body
657
860
Tail
349
300
Hindfoot
164
155
Ear
54
55
RATIOS (pel cent)
Tail/head & body
53
35
Zygom. br./condylob. I.
75
71
Braincase/condylob. 1.
44
4^
Braincase/zygom. br.
58
58
Palatiiar l./condylob. 1.
42
40
Intel orb./postoib.
83
87
p'^k — 111^
41-0
37-1
PANTHERA 437
other species; there was only a shght scratching with the hindfeet before defaecation
or urination. This, too, would seem to postulate arboreal rather than terrestrial habits,
for if defaecation generally took place from a tree branch there would be no point in, or
possibility of, covering up the droppings.
Measurements. The table on page 436 is derived from 4 West African and 3 closely
extralimital specimens (i skin only of each); that is to say, broadly, a. cclido^astcr and
rest skins from [uaso (Ghana),
No. 1938. 7.25. 1, fig. 60b, than any Sudan zone specimen.
Since neither sktdls nor any body measurements of Guinea zone animals exist it is not
possible to assess size; nor does any constant distinction of colour or maculation between
Guinea and Sudan zone skins emerge from the existing material. It is conceivable that
leopards born within die grassless closed-forest respect the sharp natural boundary that
exists between it and the densely grassed contiguous Guinea zone and therefore rem.iin
within It; but that animals born 111 one or other of the grassed open-woodland zones
see no barrier to roaming indiscriminantly from iine mdehnitely bounded type to
another. Be that as it may, it is not considered in this work that die data at present
available justif)- anything more than a broad division of West African material into two
sections, closed-forest and open-country. This, in effect, is what Pocock (1932) did,
assigning die rwo races to li'opaiihis Schreber and nicli-.iioifi Cabrera respeciively.
Both of these would at first sight seem to be jiistifi.ible names; but, as already pointed
out, if hopaniiis is to represent die forest leopard, as it reasonably might, then the arbi-
trary choice ot Senegal rather than "Guinea" as the rv'pe locality is most inappropriate.
As tor nicluiuvi'i, Pocock assumed (1932: 577) that Bornu. whence one of his two
assigned specimens came, was "no doubt very similar to Yoko". the type locality, and
that there was therefore no ecological objection to lumping the Nigerian and the
Cameroun animals together. The veget.ilional assumption is, in fact, not vahd since
Bornu is in the dry Sudan zone and Yoko 111 the moister invasive Guinea woodland,
f.iter, Dobroruka (1966a), having examined specimens from northern Cameroun,
Mango mountains and Duma (Ubaiigi), including a topotvpical skull, came to the
conclusion that rriclinioii'i was identical with the leopards of the upper Nile, that is, in
his view, with the iiomin.ite r.ice, /'. p. pardiis. No one, liowevu', knows where pardiis
came from. Its type locality was given by Linnaeus (1758) as India but w.is fixed later
(1911) by Thomas as Egypt, and subsequently (1924) designated by |. A. Allen as
Algeria — thus establishing as representing the nominate r.ice .\n animal whose characters.
PANTIIERA PAROUS 457
range of colour and maculation have never been clear and, since it is iii all probability
extinct, are never likely to be known. Pocock by implication rejected Allen's designa-
tion; for in his 1936 paper he gave the upper Nile as the type locality — that is, none of
the hitherto postulated areas. Nevertheless, it was doubtless on the basis of this that
Dobroruka equated rckhcnowi with pardus.
Pocock himself indeed, had already half come to Dobroruka's conclusion, stating
(1936; 924) that in his opinion rciclunotri was only provisionally admissible since the
range of the race in question probably extended to the Egyptian Sudan and the upper
Nile and the animal was thus either pardus or else chiii Heller, 1913 — which in any case
Itself was hkely to prove to be identical ^\^th panius. Dobroruka (1966a), however,
differed from Pocock in beheving that the latter's description of the West African
specnnens which he assigned to rciclunowi depicted a paler, yellower animal that was
thus not of that race (or pardus) but required to be distmguishcd by a new name.
The present writer, having seen how much variation in ground-colour and in
markings regularly occurs within a limited area, does not place much weight on
assertions that leopards from given localities are truly characterized by specific shades
of colour and thus recognisably, and consistently, differ from those of other localities by
slight nuances — the more so where the evidence for such nuances rests upon written
descriptions by different authors. Further, he is of the opinion, as already mentioned,
that opcn-coiuitry leopards are in the course of their hves most hkely wide-ranging, in
respect of different woodland zones as well as of distance, resulting in considerable
interchange and interbreeding. The case for splitting them up into a number of vahdly
definable races is far from proven. In consequence it would be less confusing and just as
near the at present ascertainable truth to regard the open-coimtry leopards, at least those
of the circum-Saharan regions here under discussion, as all being, in contradistinction
to the duller-toned forest animals, of the nominate race pardus. This course is adopted
in this present work. In this connexion it is of interest to point out that Dobroruka
(1966b), investigating the leopards of South Africa, came to the conclusion from study
of the usual taxonomic characters of colour, marking and size that three good sub-
species could be distinguished: but that Hemmer (1967), was, however, as the result
of subsequent statistical assessment of these characters unable "to justify- sphtting up the
leopards of eastern, south-western and southern Africa in several subspecies".
There remain two specimens to be mentioned. The first is a skull. No. 49.604,
labelled merely "West Africa". From its size and structure this would seem without
doubt to be that of a forest female. More controversial are a skin and skull of
Sanderson's, No. 48.768 from Bamumbo (Cameroun), which lies at an elevation of
some 1000 metres in a region of high precipitation on the escarpment between Mamfe
and Bamenda. The vegetation is essentially heavy rain forest but has been much
destroyed and substituted by^ extensive stands of oil-palm and areas of invading grass.
The skin has, rightly, the characteristic dullness of a forest animal; but the skull, that of
a fully mature or even old male, is the biggest in the British Museum, far exceeding the
forest series and somewhat larger than the Sudan woodland male. The specimen is
therefore difficult to place. There is neither date nor any field note on the label, but
Pocock (1936) states, presumably from information supplied by the collector, that it
458 THE CARNIVORES OF WEST AFRICA
was shot on February i()th "in the palm bek at the junction of the forest country with
the moimtam grass". This does not necessarily mean that it was shot by Sanderson, who
did, in fact, obtain a good number of specimens from local hunters. The skin would
scarcely seem to have been prepared by, or under the eye of so experienced a collector
as Sanderson: and the bullet holes seem more consistent with those made by a Dane gun
than a modern rifle. Lastly, the skull has none of the appearance of havuig been
professionally collected and prepared, lacking its lower jaw and being smoked and
dirty as from long suspension in a ju-ju house. It may, thus, not belong to the skin.
Hiuiters in this border area coidd as easily obtain leopards from the adjacent very open
Bamenda district as from the dense Mainfe forest and may well have sold as a reputed
unit two separately and distantly derived specimens.
It should be added that in none of the British Museum skins from whatever locality
docs the length and texture of the pelage vary enough to be reckoned a usuable taxon-
omic character. Some seems a trifle less harsh than others, but this is probably as much a
matter of preparation as of anything else.
Panthera pardus pardus (Linnaeus) West African Open-country Leopard
The skull measurements ot this, the larger race, are shown below but there are no
collectors' body measurements. A specimen from Lake Chad is given in Rowland
Ward's Records as having a total overall length before skinning of 249 cm (8 ft 2 in).
As understood 111 this present work this form may occur throughout West Africa
except the forest belt. It is said to be rare in the Sahel zone of Mali, though it docs occur
there, more especially in the wooded islands (fO(_'/(m's) of the flood plain of the middle
Niger. It is common in the Sudan and Guinea woodlands both in West Africa and
extralimitally. As regards the former, British Museum specimens exist from Wulc
District (Gambia) ; near Sefadu, Kono District, and 5 skins from near Gberia Timbako,
Koinadugu District (Sierra Leone), 2 skins and skulls from Sicili River, Northern
Territories (Ghana): 2 skins from Boriiu, and a skull from Gorgoram, Yobc River
(Nigeria).
The skins in this race are vei'y variable but are all characterized by their bright, lively,
warm coloration with no trace of interstitial greying as in the following race. However,
the two pairs from Bornu and the Sicili River demonstrate how specimens can vary in
a single locality. The different maculation of the first pair is illustrated in fig. 6oc and
d: their colour is also appreciably different, skm No. 1939. 164S being considerably
redder than No. 7.12. 12. 2. The two Sicili River animals arc also prett)' distinct as
regards both markings and colour. No. 33. 7. 14. i having much sm-iller spots and a
more general redder hue than No. 35.10.22.72, both V.
Panthera pardus leopardus (Schreber) West African Forest Leopard
This is the smaller of the two West Atrican races, the skull sizes being shown m the
table that follows. It is, once again, not possible to give any figures for body size since
no field measurements exist and, as shown in Rowland Ward's Records (1928), a skin
may stretch anything up to 380 mm in dressing, so that measurements from
PANTHERA (lEo)
459
iiiusL'uni examples arc valueless. This form may occur anywhere in the forest belt, even
near centres of population, from Sierra Leone to Camcroun and beyond, specimens in
the British Museum coming from near Monrovia (Liberia); Goaso, Juaso, Bcssiadzi
and Foso (Ghana); Ikotmbo, Oban and Ogoja Province (Nigeria); various locahties
around Mamfe (Cameroun). Kuhn (1965) records it from the following Liberian
localities: Kpeaple, Freemantown, Grahnstown, Tappita, all in north central Liberia.
Eisentraut (1963) mentions early records of leopards at various points around the foot
of the Cameroun Mountain and up to 1500 metres; but he concluded that in recent
times these predators have there become extraordinarily rare.
As stated above, both coloration and maculation are very variable; but the back-
ground colour between the spots has a dull, somewhat greyed appearance. The spots
arc often heavy; and there are broad oblong spots forming interrupted parallel hues
down the spine.
Table 29 : Numerical data for Panthcra pardus
p. pardus.
p. pardus.
p. leopardus,
p. leopardus.
Cameroun,
S
'■f
0
0
Bamumbo
Vegetation
Woodland
Woodland
Forest
Forest
>
Number in mean
I
3
I
5
I
Condylobasal length
224.
i8s
204
172
230
Basilar length
205
174
186
156
(210)
Palatilar length
102
86
99
81
105
Zygomatic breadth
154
126
147
118
163
Upper cheekteeth breadth
83-4
74-9
76-1
70-5
89-8
Nasals, length
fi8-5
60-7
71-0
57-1
82-3
Interorbital breadth
45-4
37-2
39-8
32-1
43-3
Postorbital constriction
41-0
41-6
36-3
40-1
39-4
Braincase breadth
75
71
75
68
80
Toothrow (c — m^)
78
68
73
63
80
p* length
27-7
24-7
24-6
23-5
28-3
i»l breadth
8-8
7-4
(6-5)
5-8
8-5
(»l length
l8-2
i8-o
16-5
—
RATIOS (per cent)
Zygom. hr./condylob. 1.
69
68
72
69
71
Braincase/condylob. 1.
33
38
37
40
35
Braincase/zygom. br.
49
S6
51
S8
49
Palatilar l./condylob. 1.
46
46
48
47
46
Interorb./postorb.
III
89
no
80
no
p*lc — m^
35-6
36-4
33-8
37-4
35-4
Subgenus LEO Brehm, 1829
Lions
The characteristics ot this monospecific subgenus can be gathered from the account
of the species which follows. The name Lcc was originally used by Okcn, 1816; but
since that author's Lchrhuch dcr Naturgcschkhtc has been rejected by the International
460 THE CARNIVORES OF WEST AERICA
Commission on Zoological Nomenclature it can fortunately without causing any
confusion be better attributed to Brchm.
PANTHERA LEO (Linnaeus) Lion
Felis leo Liimaeiis, 1758, Systfiiia Xitturcu-. loth edition, I: 41. Constantine, Algeria, as designated by
J. A. Allen (19^4): 222. Leo was the Latin name for a lion.
Felis leo, race 3, sciie);aleiisis, ]. N. von Meyer, 1S26, Disserlalio iiiau^urahs aiiatoinico-ineJica de Gciiere
Felium: 6. Senegal, inferred from the given name. (Not Felis senegakiisis Lesson, 1S39, tor a small-
spotted serval).
Felis leo, B. seiieflaleiisis]. B. Fischer, 1S29, Synopsis Manimalium: 197. Senegal, inferred from the name,
which was based on the "Lion dii Senegal" of Ciivicr & GeofFroy, Histoirc Nciliirelle des Matnmifercs,
Part 9, 1819.
Leo gatitbiaiius Gray, 1S43, Lisi 0/ ilie SpcLiiiiciis of Miviinuilui »/ tlif . . . British Miiscmn: 40. West Africa,
interior of Gambia. A iioincu iiudtiiii.
Felis leo kmupizi Matschie, 1900, Sber. Gcs. iialurf. I'lennde BeiL: 92-^3, and Plate (skull). Yoko, upper
River Sanaga (Cameroun). This was named after Major von Kamptz, Commandant of the German
Imperial Troops in Cameroun, who made a collection of several mammals, of which this was one-
Distribution and general. The lion is certainly among the best known of all
animals, in popular imagination the King of Beasts — a title that has often been disputed
by animal lovers from Lidia who maintain that the tiger has far better claim to it
since it is somewhat bigger, yet more powerful and certainly more courageous. The
lion owes its fame to the proud bearing and impressive mane ot the male annual;
and its precedence in popular estimation over the tiger to having played a peculiarly
significant role in western civilization for many hundreds of years during which the
tiger was by comparison luiknown. This role may have been very closely real in the
Roman arenas but was mostly more remotely symbolic in a widely accepted folk
tradition, the lion having become since the most ancient historic times, from biblical
lands to northern Europe, a universally imderstood emblem of might, dread or endur-
ance, as exemphfied in poetic literature, statuary, heraldry or even quahrs* marking.
It remains so to this day. The reputed nobilit)- of the lion has influenced also the
collective noun coined to denote a congregation of them, a "pride" of lions — a term
dating from the middle of the 15th century. Its modern common use, however, is a
relatively recent cultural revival, "troop" having sufficed as the accepted description
m both literature and speech luitil the 1920s. Lions are, in nature, often far more
readily visible than almost any other of the wild cats, not uncommonlv roaming or
lying at ease in the open or only partly concealed; whereas the tiger, like the leopard,
is inherently far more secretive and is seen only when it breaks cover.
In prehistoric, neolithic, times the lion was spread not only over the whole ot Africa
and much of Asia but over a good deal of Europe as well. Indeed, in early historic
times it is possible that it still existed 111 Greece. It certainly then occurred around some
of the shores of the Mediterranean, not only in north Africa but without doubt m
Asia Minor and very probably in Syria and Palestine as well. Further, it must have
been abundant since it was presumably from these conveniently neighbouring sources
that, somewhat later, it was imported into Rome in exceedingly large numbers. The
a
PANTHERA LEO 46I
species continued to be plentiful m Africa and southern Asia until ven,- recently: but
within the last hundred years or so it has been extensively destroyed, a process which
reached a peak in the hrst half of the present century, when "big-game hunting"
became a fashionable pastime. This, together with the fact that the vital role of the
lion and other carnivores in preserving a proper balance of nature was not appreciated
and they were relentlessly destroyed as wholly lurdcsirable "vermin", has brought
about the present position where it has ceased to exist over vast stretches of its recent
range and occurs only in much reduced numbers m the areas in which it still maintains
a foothold. When one reads of the degree of destruction that went on unquestioned
the marvel is that the species has not followed the dodo into extinction. Without here
repeatmg well-known names, it is on record, sometimes from their own pens, that in
pursuit of "sport" or imder some seemingly more laudable-sounding pretext or another
good many hunters have individually killed or helped to kill scores of lions during
the course of their Uves — apart from the countless thousands of less prominent figures
who, following their owii personal satisfaction or gain, have achieved a lesser slaughter.
The result is today that the Hon is extinct in Asia except for one small district in
north-western India; and as fir as Africa is concerned its range is very greatly reduced.
From having flourished eveiTi-where m the contment except the closed forest it is
now to all intents and purposes confined to the tropics alone, and even there has become
extinct in many areas and rare in others. The species was virtually exterminated from
Egypt a couple of centuries ago; the last survivors were killed in Algeria and Tunisia
in the early 1890s, and in Morocco in 1920. At the other end of the range the hon
retreated from the Cape and Natal a very long time ago and exists now in only a few
of the more northerly parts of the Republic, its southern limit today being roughly
20 to 25° S. except in nature reserves or as an occasional stray. Lions are now at their
most plentiful in Tanzania and Kenya.
As for West Africa, there has been a very considerable recession, especially in recent
rimes. It should perhaps first be made clear that the lion's most favoured vegetational
types in West Africa are the Sahcl and Sudan zones. However, the species is also to
be found, less commonly, in the Subdesert and, during the dry season when the dense
grass has been burnt and the coiurtry assumes a relatively open character, as a tem-
porary visitor in the Guinea (or Invasive) woodlands, whither for a restricted period
it follows the game. It never enters the true high-forest except possiblv occasionally
to lie up in its extreme fringes abutting the grasslands; but it may shelter during the
day in the hght forest cover fringing river banks, hi East Africa Hons have been recorded
as ascending as high as 3500 metres into bamboo forest, but such elevations scarcely
exist in West Africa apart from the Cameroun Mountain. Nevertheless, lions are
known to occur at some 2000 metres or more in some of the Cameromi highlands.
There are in West Africa no extensive open plains such as exist, and arc frequented by
lions, in the more easterly and southerly parts of the continent; nor are there vast herds
of ungulates whose migrations in search of fresh grazing commonly, ui those regions,
determine the complementary occurrence of their predators. The lion in West Africa,
in fact, leads, and always has led, a far more restricted existence than in East Africa,
and one with considerably less abundant food resources.
4'>2 THE ( ARNIVCIRES OF WEST AFRICA
Rock paintuigs show that iii coninion witli several othcF now extinct species the
hon must once have dwelt in parts of what is now the Sahara (Lhote, 1951). It has been
absent from this area for very many centuries but continued to exist on the cdE^e of the
desert in Air up to some 60 years ago. South of this it occurs over a fairly wide area,
though nowhere abundantly and in rather scattered fashion. It is found along the course
ot the River Senegal and is probabK", in this region of Africa, at its most plentiful around
the great bend of the Niger, both nortli and south of the river. Thence it ranges across
to Lake Chad and beyond, taking in the northernmost parts of Gambia, Guinea,
Sierra Leone, Ivory Coast, Ghana, Togo, Dahomey, Nigeria and Cameroun.
Particular records for West Africa are few, and some are now so much out of date
that they may not truly reflect present distribution. According to a letter 111 the British
Museum from the late Sir Cecil Armitage, who was at the time Governor of Gambia
as well as a keen sportsman, lions turned up in 1926 within 20 km of Bathurst; and
later, in 1929, one was shot on the river bank at Basse near the extreme east of the
territory. One of the few West African specimens in the British Museum had been
shot at Tambana a few years earlier; and another near Bathurst. The situation today in
Gambia is unrecorded, but doubtless wanderers from further east still from time to time
enter the coimtr\-. Monod (1940), writhig of Portuguese Guinea, noted that at that date
hons had become rare and existed permanently only in the neighbourhood of Corabal
and Buba, where they were pretty common. From these places they sometimes visited
Chitoli and occasionally roamed uno other parts.
Stanley (in Goddaid, 1925: 227) recorded that lions were nothing more than rare
visitors to Sierra Leone at the height of the dry season from the neighbouring territory
to the north. One was shot in 1924 at Yiraia, north of the Loma Mountains (about
9"26 N, ir 16 W), while it was devouring a giant pangolin; and there is an account
of this together with a photograph of both animals in Stanley & Hodgson (no date
but C.1930). T. S. Jones (personal communication) was informed by a member of the
Veterinary Department that as recently as 1956 a lion had been seen in Sulima Chief-
dom, where it had killed cattle. There seems to be no later recorei than this for Sierra
Leone, nor is there any specimen in the British Museum from that country.
There is, however, a skin in the British Museum from Ghana (Tamale, Northern
Territories in Doka woodland, received in 1946); and one from somewhat to the north
of this, an unnamed locality in the Voltaic Republic, Sudan woodland, received in
1930. Ci. S. Cansdale, quoted in Guggisberg (1963) gives the impression of much wider
occurrence and more permanent residence m Ghana than is the case in Sierra Leone.
In this note he stated that hons were still found in many parts, as far south, indeed,
as the Afram Plains. The Northern Territories were their mam stronghold; but he
had also heard them roaring in the nordi-west corner of Ashanti near the River Volta,
where they were locally common. This lies in Guinea or Doka woodland; but whether
the lioiiN were there at all seasons of the year is not clear. Cansdale also said they occurred
as Well 111 thinly populated grass areas 111 the extreme south of western Togo. Lions
also exist at the present time m the i'are National of West Dahomey (Howell, 1968).
In Nigeria the species has never 111 recent times been numerous and is now much
inon uncommon than it w.is even 40 ye.ns ago. Nevertheless, in spite of a fairly dense
PANTHERA LEO 463
human population lions have continued to survive, though as somewhat of a rarity,
hi the drier vegetation zones, chiefly north of the Bcnuc; but they have also for long
existed west of the Niger, especially in the once remote and unpopulated disttict of
Borgu. Since the creation of the Borgu Game Reserve there is, in fact, some sign of an
increase in their numbers there. These were down to about 30 in 1962, but in 1968
they were estimated to be between 50 and 75, and with increasing and more stable food
resources they are themselves becoming less inclined to wander (Howell, 1968). They
may still exist a little further south, at least as seasonal visitors, in Old Oyo and Upper
Ogun. For the rest, the species may be come across in limited numbers, and in pairs
rather than in prides, throughout the open woodlands of northern Nigeria from Sokoto
and Argungu to Bornu; but the hon being often somewhat of a wanderer, especially
where game is scarce and sporadic as m much of this region, cannot always be definitely
related to a given locality and often suddenly appears 111 unexpected areas from which
it is normally considered absent. Some places in which it turns up may, in fact, be very
unexpected indeed. A score of years or more ago one lay for some time only partly
concealed in the grass opposite Yola police station watching all the comings and goings
until it was eventually shot by a police officer. Ajiother was killed not long ago by a
lorry on the road between Potiskum and Jos; and A. J. Hopson, m a personal com-
munication, records that as recently as 1968 two young lions were shot by Fulanis on
the Bama ridge between Maiduguri and Bama, having killed two people. With the
increase m prey species taking place in the Yankari Game Reserve (Bauclii Province)
lions may there, as well as in Borgu, now be becoming more numerous and sedentary.
Two skins from Nigeria exist in the British Museum, one without any localitv', the
other (Sanderson, 1940) from the Ogoja Province.
This last locality, which is contiguous with Camcroun, is nevertheless rather miusual.
In Camcroun itself hons still occur in many parts. Over a long period of years at least
one pride existed with little disturbance on the Mambila Plateau, 200 km north-east
of Bamenda, but is now in some danger. From there an occasional straggler has reached
Ndu and other places somewhat further south. But it is considerably further north of
tins that the lions of Cameroun and the neighbouring country of Chad become apprec-
ciably more common; that is, around Rei Bouba and other localities in the upper
Bcnue area and along the Logone and Shari Rivers to Lake Chad and fuither north
towards Wadai and cast to Sudan. One of the reputed West African races was described
in 1900 from a specimen from Yoko, which is in the Invasive woodland zone somewhat
north of the River Sanaga, one of the southern boundaries of the area taken m this work
as constituting West Africa. According to Guggisberg (1963) hons still exist in this
locahtv', or existed till recently, 10 specimens having been killed there in 1953.
Description. The lion is far and away the largest of African carnivores, a fully
grown animal weiglung between 170 and 220 kg, that is to say some three or four
times as much as a leopard. The males are appreciably bulkier than the females though
not so greatly different as regards the usual body measurements. It has been argued
that West African hons arc smaller than those from East Africa, and it is possible that
a far more abundant food supply and a less harassed, more stable, existence in the
east may lead there to the attainment of better growth: but the amount of com-
464 THE CAIiNlVOUrS OF WEST AFRICA
p.uMtivc in.itci-i.il and held daca available troin West Africa is realh- far too slia;ht to
make any reasonably reliable deduction possible. As so often, in nianimalogy as in
other matters, diere is a danger of seizing upon the exceptional as representative.
In Rowland Ward's Records the largest specimen is East African and is given as
measuring from nose to tail tip 332 cm; but the average is certainly very considerably
less. The three largest West African animals, from Lake Chad, Northern Nigeria and
Sierra Leone, in the same reference work range from 290 to 297 cm. The avera2:e
adult come across in West Africa would, however, probably be more in the nature of
270 cm, of which roughly 180 cm would be head & body and 90 cm tail. Shoulder
height might be of the order of 100 cm; the weight 190 kg.
Adult lions differ very markedly in appearance from juveniles in that the latter
are heavily spotted .ill over whereas the former, save in exceptional cases, are self-
coloured on back and flanks. The colour description of a lion's pelage is m general
terms given as tawny, that is to say a pale to medium brownish hue such as is com-
monly brought about by the tanning of leather; but, in fact, the colour of lions has a
fairly wide range between dull yellowish and medium reddish-brown, and may some-
times be considerably darkened by an abiuidance of black tipping to the individual
hairs. These difl^erent shades of brown have been used to diagnose local races but may,
indeed, commonly be found to exist within a single pride. It is not, therefore, possible
to provide a general colour description of West African lions, and the matter will
be gone into a little more fully in the t.ixonomic section which follows later. There
is usually some intensification of colour along the spine. The colour of zoo-kept lions
may change markedly.
The pelage is very sh.M t. 1 .ither harsh and close-King. It is composed of relatively
sparse, very fine underfur, 0 to S mm long, sometimes yellow-tipped; and of flattish-
sectioned sub-bristles some 10 to 14 mm long, the petiole being comparatively short,
about 4 mm, the blade quite long, 6 to 10 mm, and of whatever shade of yellowish-
brown the coat ni.iy be except fov a shorter or longer black tip. In the majority' of
cases the duection of the pelage is reversed on the b.iek from about the hips to the
•■houlders or a little further, there being whorls at each end where the change of direc-
tion takes place. This is so in four of the West African specimens in the British Museum,
but in the fifth, that from Gambia, there is no sign of this character.
In the lioness, where it is not obscured by the existence of a mane, the reverse of
the hair is usually continued forward as a lo\\-. narrow crest along the back of die neck
to the crown.
Li the adult male lion there is an .ibiuidance of lengthy hair, termed the mane,
around the sides of die face, over the top and sides of the head, neck, chest and between
the shoulders. The ears are only partly or not at all concealed. In some reputed races,
or at any rate individuals, the mane extends further along the back and in some cases
along the underside of the body; and there may be tufts on the elbows. Apart from
the extent of the area it covers, the mane vanes a good deal m length, abundance and
colour, this last ranging from black to bright gingery-yellow. All manes have at least
a few black hairs amongst them and blackness comes about when that colour pre-
donnnates over tawnv. Matiiiitv has a i/ood deal to do with the existence or size of a
PANTHERA LEO 465
mane, and reports of nianclcssncss or of poorly developed specimens may be due to
relative youthfulness. Reliable evidence of age rarely exists in these cases; for it is
often judged at some distance in the field or, in study specimens, may be impossible
to come by since, often enough, no skull accompanies the skin. However, Mazak
(1964b) has studied the colour of manes in relation to age and fuids that in their early
stages they are always yellow or tawny, and only much later develop a dark colour,
eventually black. This explains why a single pride may contain adult males with
cither black or tawny manes, as commonly observed. This author found blackness to
have no connexion with true melanism but to be the result of age and individual
idiosyncrasy; and it was not a racial characteristic. There is, indeed, httlc evidence
that mane colour or size have any of the significance once accorded them by taxono-
mists. Tliis is borne out, for instance, by both text and photographs in Guggisberg
(1963 : 47), clearly demonstrating that colour ar.d abundance of growth differ widely
in a single locahry. There may be tendencies in a given area but certainly no con-
stancy— the black-maned lions once held to predominate in South Africa, but now
extinct, being a possible example (Mazak, 1964a). Nevertheless, the fact remains that
despite the thousands of lions that have been killed there docs not exist enough concrete
evidence to lead to any certain conclusion.
Interpretations of what is characteristic of an area differ, hi the relevant section,
pubhshcd in 18 19, of Geoffroy & Cuvier's Histoire Naturelle des Mammifercs, on
which ten years later Meyer's scncgalcnsis was based, the Senegal hoii was said in
the text to differ from the Barbary lion by its yellower, more brilliant coat and a
mane that was neither so long nor so thick; but the accompanymg Plate depicts a
pretn,- abiuidant blackish mane, and blackish elbow tufts. Matscliie (1900) reviewing
the various forms gave the colour of the Senegal race as red-yellow, the mane poorly
developed. Of the four male West African specimens in the British Museum two,
from the distributional and environmental point of view, might be supposed to be
Senegal lions, those from Gambia and Upper Volta. Neither has any sign of a mane
or elbow tufts. Sir Cecil Armitage, who took a good de.al of first-hand interest in these
matters, stated in the British Museum letter referred to in the previous section (page
462) that the lions of Gambia were practically maneless; and certainly a young male
from Gambia in the London Zoo in the 1930s evinced no sign of a mane. The Sierra"
Leone lion showni in the photograph facing page 144 of Stanley' & Hodgson (n.d.)
had a poor, rather short mane. The other two British Museum West African speci-
mens, both from Nigeria, might be regarded as more probably belonging to Mat-
schie's kamptzi from Canieroun. What is left of the manes of these is a very bright
giiiger colour, in places almost orange, with a few scattered black hairs; but a Nigerian
lion from Lokoja, in the same kind of vegetation on the Niger near the confluence
of the Benue, also on exhibition at the London Zoo in the 1930s, had a fairly well-
developed mane certainly not bright ginger.
The mane, therefore, is of very variable appearance and no set description of colour
or abundance is possible for West Africa. Another factor has not been mentioned. It is
sometimes asserted that manes arc apt to get worn out, or torn out, by constantly
thrusting through dense undergrowth, and that the full potential of a lion may only
466 THE CAKNIVORES OF WEST AIRICA
bfcomc evident after some tune in captivity. Tliere is some possibility ot trutli in tins
as concerns lions winch habitually live amongst dense thorn bushes; but it counts for
little or nothuig for the great majontv that spend their lives in sparse open-woodlands.
The heads ot both lion and lioness are basically of the same shape, masked, however,
m the former by the mane. It is rather rounded and broad in general feline fashion but
the massive muzzle not so shortened as in the most typical cats. The ears are of fan'
size, set well apart on the sides of the head, with a marginal bursa, the apical part very
rounded, almost semicircular in outline. The front face of the pinna and the car opening
are well protected by dense, long, stiff hairs. On the posterior side there is an intense
black patch occupying the middle part, a little variable in shape but in general with
concave upper and lower margins resulting in a band that is broad at the ear rims and
narrow waisted in the middle, leaving both basal and apical areas of the normal coat
colour. The pinnae have considerable freedom of movement, able to stand erect, he
flat or by twisting to present the posterior black marks to view from the front as a
threat.
The nose is broad and flat; the rhinarium very narrow, having no dorsal fice what-
soever but continuing laterally backwards around the somewhat comma-shaped,
posteriorly long, slit-like nostrils. From the rhinarium a bare vertical median stripe
parts the heavy^ upper lips, which are white or whitish and plentifully furnished with
strong vibrissae, the majority of them set in about three or four parallel rows. The lower
lips and chin are also white or whitish; and there is a conspicuous pale line beneath
each eye and, less obvious, often a smaller similar mark at the inner corner. The
eyes themselves arc fliirly large with yellowish-amber irises and pupils that remain
round on contraction. In the adult the eyes have for the most part, except at the
approach to a kill, a rather tired or sombre look and are often held half closed. This is
quite diflferent from other African felines, say the alert, perky look of the serval or
penetratingly fierce stare of the caracal. There are well-developed supercihary tufts of
vibrissae. The legs are stout and extremely muscular, the paws massive and armed
with very powerful retractile claws. The tail is, in its colour and close-lying short hair,
exactly similar to the body. It is very flexible, about two-fifths to a half of the head
& body in length, of circular section but markedly tapering from root to tip, the
latter cariymg a small tuft of long black hairs. Hidden amongst this last is an elongated
patch of hard, calloused skin, sometimes popularly referred to as the "thorn".
Yoimg lions are spotted all over, the spots being rather of the "rosette" form charac-
teristic of leopards but far more obscure, of paler colour and more closely approxi-
mated than is general in that species. There is some tendency for them to be disposed
in transverse lines. In all but extremely exceptional cases these markings disappear
entirely from the back of the adult animal but may sometimes remain fairlv clearly
on the lower flanks. The belly, legs and feet may more frequently remain spotted
throughout life. In colour, the luiderside, chest and insides of the limbs are whitish or
at least very pale buff.
Skull (figs. 63 and 64). This is very similar in appearance to that of the leopard except
that It is almost twice its size. It is yet more solidly built, with massive zygomatic
arches .md a large sagittal crest for the attachment of powerful muscles. This crest
PANTHERA LEO 467
exists also in the female though it is there a little lower; and, apart from a slight dis-
parity of size, there is not that clear difference of build between the sexes that exists
m the leopard. But proper comparison in this respect is not so easy as it might be since
a very high percentage of British Museum African specimens were not scxed by their
collectors and relatively few authentic adult female skulls are available. Only a single
West African skull, a male, exists.
The braincasc, as in the leopard, is very narrow; and, in fact, appears to be in pro-
portion even narrower. Both are much more slender than in the rest of the Felinac ; in the
only West African hoii skull available the braincase measures some 34 per cent of the
condylobasal length ; m members of the genera Fclis and Aciiwnyx it lies between 42
and 54 per cent. The skull profile is not so curved as in the leopard, falling away much
more gradually both anteriorly and posteriorly from a slightly sunken frontal region
between the postorbital processes. These latter are short, blunt and distantly separated
from the large, sharp jugal processes leaving a widely open orbital ring. The rostrum
is slightly longer than in the leopard. The upper branch of the premaxilla extends,
very narrowly, for a short distance between the nasal and maxilla; the anterior nares
are large and open. The palate is broad in comparison with the majority of Carnivora
but not so broad in relation to the condylobasal length as in the other two feline
genera. The postdental palate is much wider than long. The mcsopterygoid fossa is
wide and deep, the pterygoid processes basally large but distally slender, verj' long
and hooked towards the rear. The bullae are not large in comparison vnth the overall
skull size; but the flanking mastoids have very well-developed subpyramidal pro-
cesses; and each paroccipital process comprises a broad flat portion that closely invests
the posterior face of the bulla and a thick downwardly projecting finger. The lower
jaw is massive, the rami deeply excavated posteriorly, the angular processes strong and
incurved.
The incisors of both jaws form sohdly compact transverse rows; the outer ones of
the upper jaw much larger than the inner ones and subcaniniform in shape. The canines
themselves arc extremely large, about 60 mm tall. The cheekteeth are set in a straight
row; p'^ is lackuig; p^ is much smaller than the rest and is little, if at all, larger than the
inner incisors; p^ has one cusp anterior and two posterior to the main triangular cusp;
p*, the carnassial, is very powerful, the blade trifid, but there is no cusp above the
antero-internal root. The solitarv' upper molar, set transversely in the jaw, is small
and with limited function, bearing against the posterior face of the lower carnassial.
In the mandible the canines are very large but not quite so tall as those of the upper
jaw. There are very long diastemas between them and the anterior premolars (pa),
and when the jaw closes a very large postcanine gap results. The two premolars are
each triftd; the carnassial much smaller than that of the upper jaw.
Hollister (1918) found that the "skulls and teeth of females vary much more than
do those of males. The range ot variation in size of the teeth in honesses from one
locality is startling". Much the same applied to bullae. J. A. Allen (1924) stated that
the upper carnassial is relatively shorter and broader in females than males and more
variable in both size and form. He also found the upper canines to be extremely variable,
and the vestigial upper molar the most variable of all.
468
THE CARNIVORIS n] VVI.ST AriUCA
It is pcrliaps worth icccirclmg licic tli.it in .in carlu'r paper Ilollistcr (1917) was
emphatic that hons hrcd 111 captivity were useless for taxonomic purposes as, through
the absence of normal muscular use called for m the wild, the skulls display very pro-
nounced differences in development and consequent measurements and proportions.
Habits. The lion has been more written about, more photographed, more talked
of than any other animal; and therefore a good deal is known about the species. The
literature is indeed vast, some of it scientific, the greater part of it anecdotal, not all
of it wholly reliable. The most complete, recent, survey of this much discussed animal
from the earliest times to the present day is that of Guggisberg (1963) who has brought
Fig. 63. I'aiiiluui Ico: skuli, H.M. No. 21. 7.23.1,
^ ; l,itcr.il view
together a very detailed account from his own wide firsthand experience and an
extremely lengthy reading list. The notes which follow arc necessarily by comparison
brief and therefore often of a general nature m a field where generalizations arc risky.
In tlie introduction at the beginning of this present work the reader was cautioned that
all mamm.ils are individirals with recognisable characters of their own; the lion is
perhaps more of an individtiahst than any other. Each animal has clear personal idio-
syncrasies; prides may differ from one another in their group behaviour; and in different
parts of the continent whole populations may exhibit diverse preferences m feeding or
peculiarities of hunting methods. ^X/^lat we know of the lion comes almost com-
PANTHERA LEO
469
Fig. 64. Pmithera leo: skull, B.M. No. 21. 7.23. 1, ;?, x f ; palatal & dorsal views
470 TIIF, CAUNIVCIRES OF WF.ST AFRICA
plctcly from observations made in castciii or southern Africa and may therefore
be foimd to be not always equally true ot West Africa. Some differences certainly
exist. Howell (196.S) writing of the lions of Borgu in western Nigeria characterizes
them as "lethargic, unaggressive" and says that the local people do not fear them and
allow them to reside permanently near the villages, and if necessary rouse them when
they are sleeping to move them from the path. They even chase them from kills.
Anyone acquainted with lions m East Africa would agree at once that this is certainly
a marked difference m character; and, indeed, it would be foolish to assume that
even in West Africa such docility was common. In dealing with habits and behaviour,
therefore, one can only speak in generalities.
Lions are, for the greater part, nocturnally active; that is to say they most conunonly
do their limiting and killing by night; but this does not prevent their pursuing these
activities in daylight if circumstances are suitable. Lions, indeed, may be commonly
observed at any hour of the day unpurposefuUy on the move or cleaning themselves,
or playing or climbing into the lower branches of trees, or reclining along them
except in localities where they are under constant harassment, when they become
secretive and entirely nocturnal. Odierwise, except when hunting, they take no
particular pains to conceal themselves, even lying about m the open under the shade
of a tree or at most half hidden in grass or other low vegetation. This does not mean
that they are readily seen; for they generally match such cover extremely well, a lion's
mane being often ditiicult to distinguish at any distance from a dry grass tussock.
Solitary lions exist, both male and female; but for the most part the species is dis-
tinctly social, living and hunting in "prides". The lion is, in fict, the only cat that
regularly does so in companies of any size, the cheetah being limited mostly to pairs
or rarely more than three. Since the pride is the basic unit in the life and behaviour of
the lion it is as well to examine it in some detail. It may consist of four or five indivi-
duals but numbers of up to a dozen not luicommonly exist; and in parts of East Africa
where game is plentiful prides of 30 or even more are known. Howell (196S) says that
in western Nigeria prides of up to eight are to be seen. There is, of course, an optimum
size for a pride; and this is dictated by the size of the prey most commonly available
and the amount of scavenger competition. A solitary lion is at a disadvantage in two
ways: the size of the prey that can be tackled single-handed, and the fact that when it
leaves its kill in order to drink, as lions always must, there is no one to guard the car-
cass from jackals, hyaenas, vultures or other hungry raiders. Co-operation in a pride
not only enables larger victims to be brought down and killed but ensures also a
greater success rate as well as subsequent protection of the meal while some of its
members absent themselves to drink, rest or defaecatc. If large ungulates such as zebras
are regularlv available then a single kill can satisfy a fair number of lions and the
pride can be numerous; but if as in West Africa, such meaty prey-species are not
available and recourse must most commonly be had to gazelle or kob then a single
victim can satisfy only a smaller pride, or more than one kill must be made.
The pride is not, as often popidarly supposed, made up of a single lion lording it
ovTr his harem of females. It may be all male, all female or a mixture of the two. All-
male prides are the most unusual and are generally rather loose associations of 2, 3 or 4
PANTHERA LEO 471
lions which prefer to consort with one another but often break up, at least temporarily,
to join other prides or lionesses looking for mates. The most stable pride is one that is
wholly or basically an association of adult lionesses, some perhaps with families, some
without. They assist one another not only in hunting but in protecting and caring for
the yoiuig. Such prides may be joined for short periods by one or two males, especially
if one or more of the honesses comes into season. The purely female pride is thus turned
into a mixed one; but such prides may also simply be the natural outcome of the
male and female cubs of one or more mothers growing up and remaining together.
On the whole, lionesses are far more static than hons, which arc more given to deserting
the family or pride and joining others. Estes (1967) cites the case of one male that
simultaneously dominated two prides. Though one lion may be hierarchically dominant
there may be more than one adult male in a pride. Amongst the females themselves
one always assumes the dominant role. When one dispassionately analyses the hon
organization it becomes apparent that though it is the magnificent bearing and im-
pressive roar of the male that have captured popular imagination it is in fact the lioness
that is the backbone of the society. The pride is essentially a femmmc unit; and, as will
be seen later, it is mostly the honesses that initiate and lead the hunt — though after the
kill they may have to stand back while the males satisfy themselves; and it is the
females that bring up and almost luiaided train the future generations, which, later,
often keep in touch with their mothers for several years. Apart from short bursts of
intense activity when capturing and killing prey all hons are by nature rather indolent,
displaying nothing of the inquisitive fussiness of the mongooses, the ceaseless play-
fulness of the otters, the vigorous hmitmg of the dogs, the earnest journeyings of the
ratel or civet, or even much of purposeful prowling of the leopard. For the most
part they prefer to he about, doze or watch the world go by; and in this the lion itself
excels the Honess.
Li the normal course of events, that is to say with nocturnal, not dayhght, hunting
in view, the pride rouses itself from its lethargy in the late afternoon, the members
walking round greeting one another by head rubbing (Estes, 1967). The cubs, and in a
more restrained fashion the honesses, but not the mature males, engage in good-
hiuiioured play, chasing one another, lying in ambush, springing out and tumbling
one another over. Eventually, as the day draws to its close, the lionesses sit up and begin
to stare at neighbouring herds of game with intensity as if weighing up the possibilities
of a Idll. They may sometimes chmb into low trees to secure a better view. Eyesight,
thus, plays a prominent part in the business of hunting, and the lion's eyes are, in fact,
exceptionally large; but the sense of smell, though not so acute as m the canids, is also
highly important and the wind is often tested with the nostrils for any message it may
carry. It is the dominant lioness that takes the lead, decides if and when to move off"
and in what direction.
Li popular imagination the lion always obtains its meal in one way: it stalks its prey
and with a mighty roar leaps upon its back, clawing and biting the victim to death.
Such a picture is only partly true. Lions, bcuig lazy, will readily scavenge the kills of
others if they get the chance, driving competitors away by their superior strength.
Indeed, reversing the generally held notion, they not uncommonly let spotted hyaenas
472 THn CARNIVOKHS OP WEST AFRICA
do the killing and then come m and lob them. To this end they arc great observers of
vultures in the sky — though if they Jiave to travel any distance there may be little left
of a carcass revealed m this way by the time they reach the site. Or, in the course of their
wanderings they may come across a welcome meal by accident, a deserted fawn in the
grass, a resting oribi taken by surprise, or a bustard suddenly flushed — to be immediately
struck down with a sideways swipe of the paw. At times, instead of the usual hunt a
warthog or aardvark may actually be dug out ot its burrow. This is unusual for a felid;
and It may entail considerable effort, Schaller (1972a) recording a case in which about
3 metres of tunnel were thus laid open. The same author twice saw a lioness pull a
swimming gazelle from a river.
But the most usual forage certainly does entail stalkuig, though the climax to this,
that IS to say the kill, may be achieved in two diflrrent ways: a very close approach
succeeded by a sudden rush; or a more distant approach followed by patient waiting
111 ambush for the prey itself to come near enough to be successfully attacked. Taking
the prey almost completely unawares is an essential element of the lion's technique since
the species is normally incapable of running far or for very long as fist as most ungulates
can go once they have got into their stride. Hence the surprise spring is the lion's chief
aim; or, as second best if it cannot get itself close enough for this, a vei'y short run of
not more than 50 to 100 metres at high speed — about 60 km/h — before the culminating
leap. During such a run the body is held low, the tail stiffly erect. If the attack does not
succeed, die lion never, or rarely, makes a second attempt since by that time an antelope
would be well on its way to escape. However, Guggisberg (1963: los) records an
luuisual chase which he witnessed covering about 500 metres.
The stalk, therefore, is vital. It is carried out 111 complete silence except possibly
for an occasional low "liiih" of suppressed tension, accompanied by a nervous ffick ot
the tail, especially as the approach becomes nearer and the excitement mounts. The
head and body are held low to the ground and movement is slow and gradual. The
utmost use is made of the slightest cover; for, despite its bulk the lion is, when it wishes,
a past-master at the art of merging into its background. It crouches motionless luitil it
sees its chance of advancing furriier. With eyes never lifted from its p>rospective victim
it waits until its prey's head is down in the grass, or until m the course of grazing the
animal turns away and is badly placed to detect any movement as its attacker proceeds
to its next cover. Should the prey itself get behind cover the stalking hon may take full
advantage of this and, m no danger of being observed, carry out its next advance at a
full trot. A stalk, therefore, may be a fiirly continuous action if the cover is favourable
and the prey not very alert; or it may resolve itself into a long-drawn-out affair piunctua-
ted by lengthy pauses whilst the lion with infinite patience motionless awaits its
opportunities to advance. Approach to the prey is rarely made from the front since
the risk of being observed, even with a cautious use of cover, is too great; but Schaller
(1972a) did once see a lioness run direct!}' towards some gazelles and capture one,
but this direct onslaught was largely obscured by the lie of the land. In considering its
attack the lion must, however, take the wind into account and with this fictor in mind
manoeuvre itself only so much to the rear of its intended prey as it can safely do without
being in danger of alarming them by its scent. However, lions, like dogs, sometimes
PANTHERA LEO 473
roll on their backs in duiig or other odoriferous matter on the ground, and the only
satisfactory explanation of such a habit would seem to be the instinct to disguise their
own smell and thus render approach to their prey easier.
Lone lions must, of course, hunt for themselves; but where a pride is concerned it is
usually believed that hunting is a co-operative affair. Nevertheless, this is not a matter
upon which it is easy to be completely certain or about which there is unquestioned
agreement. Where it appears that co-operation has taken place has there really been a
common and well-iuiderstood plan of attack by several hons or has one, possibly
more active and dominant, made a kill, the others watching and ready to take advantage
of this animal's superior skill and energy? In a field study of the lion Estcs (1967: 27)
never witnessed co-operative huntmg, though he agreed that it might occur and thought
that perhaps the black patch on the back of the ears might assist visual contact between
the members of a hunting parry'. Kruuk & Turner (1967) considered it at least doubtful
that lions co-operate in hunting and thought it probable that each hon went its own
way but took advantage of any situation arising from the action of its companions.
A group might all stalk at the same time, keeping some distance apart; but this is not
the same as a dehberately concerted plan of attack. There are, however, a good many
accounts of hunts in which these animals, in Guggisberg's words (1963 : 109) "show a
high degree of co-operation and act with amazing strategy". Tliis co-operarion usually
takes the form of stealthy approach by a group in open formation to a herd of grazing
luigulates which are, at the correct moment dehberately alarmed from a given quarter
by some of the aggressors and driven into the jaws of one or more strategically placed,
concealed members of the pride who, as the fleeing animals pass, spring out of ambush
upon an misuspecting prey and effect an easy kill or kills. The rest of the pride then
close in at a trot to enjoy a meal. Some accounts add the giving of a vocal signal when
the whole ambush has been set up and all participants are in position.
Such a sequence of events may be deliberate. On the other hand, if the members
of a large pride are sufficiently spread out on the hunt, slinking along under cover,
each acting primarily for itself, it is virtually certain that at some point the prey will
take alarm and there is a very good chance that the ensuing frightened stampede will
inevitably pass within striking distance of some of the widely dispersed lions. Appear-
ances are not always what they seem, and it is sometimes difficult to arrive at a correct
interpretation of animal behaviour. Nevertheless, Schaller (1972a) is in no doubt that
co-operation takes place and, in fact, brings 30 per cent success 111 the kill as contrasted
with only 17 to 19 per cent for single stalking. Indeed, where really large or powerful
prey is tackled, as it sometimes is, concerted approach and kill by tAvo or more hons is
generally essential.
One thing, however, seems certain ; that in the vast majority of cases the honesses
take the most active part in hunting and killing. In fact, Schaller (1972a) records that
in 71 hiuits observed by him a male took the initiative in only two cases. This may be
because it is tacitly recognised as faUing to the female's duty to feed first its young
and then later the adolescent or grown male; or that she naturally has more skill;
or, being less cumbersome, is more agile and energetic; or, the common explanation,
that the males are plainly lazy. Male prides must, of course, kill for themselves; but
474 THE CARNIVORES OF WEST AFRICA
where a pride consists of botli sexes tlie males, or at least those that are fidly adiilt,
follow the females well to the rear of a hunt, only running up when tlie kill has been
made, often securing a major part of it. This in Schaller's view (1972a) has its advan-
tages for the pride; for not only are the lionesses less conspicuous in the hunt but,
in addition, large males in the rear may well discourage spotted hyaenas from making
attempts to snatch the young while the attention of the females is concentrated else-
where.
The aim of the hon m the majority of stalks is to get rather closer, if possible, to its
intended prey than about 100 metres so that the ensuing rush from cover takes too
brief a time for the v-ictim to realize its danger and accelerate to its top speed. At
60 km an hour a lion would cover 100 metres in 6 seconds. It prefers, of course, to get so
close that a single standing spring is sufficient to carry it onto the victim's back. In
places, however, the terrain is so open that really close approach is not possible and the
only hope of success lies in quietly waiting in ambush until grazing brings the herd
within attacking distance. A remarkable degree of patience is sometimes called for —
and exercised, as both Estes (1967) and Schaller (1972a) testify, the latter recording
that a lion may wait as long as 9 hours. This, of course, can be veiy considerably shor-
tened if the lion is not sohtary and if, as asserted by many, concerted hiuning takes place
and the prey is dehberatcly driven into a tacitly understood ambush.
This raises the interesting matter of the reaction of prey to predators. Many observers
have noted how quite unconcerned a herd of antelopes can be m the presence of
potentially dangerous carnivores; and this applies as much to lions as to others. Ante-
lopes mstuictively know when lions are well-fed and not interested in them, or when
they mean business. In the first case, while keeping a wary eye on them, they will go
on grazing even though the hons may be only 100 mefes away; in the second case
they take alarm as soon as there is any movement and further lengthen the distance,
though not a great deal, between themselves and their possible attackers — only far
enough, indeed, to be able to keep the lions still in view (Krumbiegel, 1953). This is
on the principle that what can be seen can be avoided; it is the surprise attack of the
hidden foe that is to be feared. It has many times been asserted that lions roar to frighten
antelopes and stampede them towards an ambush. Careful observation indicates that
this is not true. Lions do not roar as a preface to a kill but afterwards, mostly as a
means of announcing territory or of communication within the species itself Estes
(1967) found that game took no notice at all of lions roaring and played tape recordings
40 metres away from a herd of antelopes without exciting the least alarm.
Lions arc widely regarded as feeding almost exclusively on antelopes or zebras;
but though in certain parts of Africa this kind of prey may be the commonest diey do,
m fact, indulge in a very much wider range of diet, from the very large to the un-
expectedly small. It is remarkable, too, that preferences clearly vary with locality, if not
with individual prides. Goodwin (1953), writing of tlie Serengeti (Tanzani.i) reckoned
that zebra was the favourite kill; Estes (1967) found that 111 the Ngorongoro crater,
not far away, adult wildebeest came first with zebra second, the two between them
accounting for over 90 per cent of die kills. On the other hand, Mitchell it a\. (1965)
show that in the Kafue National Park (Zambia) the lion's fivourite prey was easily
PANTHERA LEO 47S
buffalo, 130 of them being taken as compared with 67 hartebcest, 30 zebra and 25
wildebeest. Choice must, of course, depend to some extent upon availabihty and in
West Africa is very much more limited. Howell (1968) thinks that in westenr Nigeria
BufFon's kob forms the main source of food; but further north in the region, in the
drier zones, gazelle most hkcly play a more important role; and probably the ubiquitous
warthog forms one of the commonest meals throughout the area. Other ungulate
species which must with greater or lesser frequency fall prey to Hons in West Africa
are hartebcest, both western and Senegal, defassa waterbuck, rccdbuck, bushbuck,
oribi, duiker, buffalo and roan. The young of the larger species, Lord Derby's eland,
giraffes, now rare, and hippo may also be taken as opportunity offers; but hons are
very wary of attacking the adults of these, and usually fnid it too dangerous to attempt
baby elephants. Nevertheless, anything is possible; and lions incontrovertibly sometimes
co-operate in the fuial bringing to earth and killing of large prey that proves too strong
for a single attacker. Vivid accoiuits have been given of fearful tussles between two
or more lions and large victims, sometunes lasting a couple of hours. Schaller (1972a)
described how five lions acted together in capturing, bringing down, turning over and
kiUing a buffalo, though this was not so much a struggle as the co-operative fmishing
off of an already exliausted animal.
The species is not above stealing its meals from other animals, and one of the most
unexpected surprises revealed by modern field research is how frequently lions feed
from the kills of hyaenas, not, as usually beheved, the other way round (Kruuk, various
papers). The intcr-rclationship of the two species in the matter of killing and feeding
is not always as simple and straightforward as it might from this appear — see page 368.
Lions take a variety of foods besides the flesh of the ungulates; but these foods are
scarcely deliberately hiuited as antelopes are but are, save in exceptional cases, come
across by accident. Whether a hon is interested in these smaller items depends on its
degree of hunger, its lack of success in pursuit of better victims, its state of health
and consequent abihry or inability through sickness or age to deal with larger prey.
Aardvarks, giant pangolins and crested porcupines are often taken, the last sometimes,
as evidenced by residual quills, the cause of painful and incapacitating woimds in body
or paws. Occasionally a grass monkey may fall as chance prey to a hon, but these
animals are normally too wary and agOe to be caught. Much smaller things are known
to be taken from time to time: hares, gromid squirrels, fat gcrbils and other rodents;
lizards, tortoises, pythons and lesser snakes; and even insects such as locusts or termites
when they swarm. Terrestrial birds are often welcome, though the only one that forms
a really satisfying meal, the ostrich, is now too rare to be of much account. And hons,
like leopards, sometimes fish, capturing catfish, or others, from lakes or rivers with a
sweep of the paw. Evidence of fruit eating is not very clear; but it has been said that
they sometimes pick up fallen plums and berries and will eat groundnuts or even rotten
wood; and that, like dogs, they eat grass to regulate their bowels. Lions are, indeed,
not so exclusively noble in their choice of food as is often supposed; venison may be
their common diet, but they sometimes take it when it is exceedingly rotten; they are
not above cannibalism should one of the pride be killed; and, if pushed for a meal,
will eat plain garbage from a village refuse heap.
476 THE CARNIVORES OF WEST AFRICA
Man-cating is a wcU-ostablishcd fact. It is usually, though not invariably, carried
out by old or sick lions. The reason for this is simple. Man, though equipped with a
superior brain, is in comparison with the rest of the animal kingdom very inadequately
provided in the matter of speed, strength, chmbing capacirv,', sensitivity to smell and
hearing, and other qualities vital to survival in the wild and is thus, unless he makes
full use ot his mental superiorit)', exceptionally poorly furnished to meet the challenge
of ciuming and determined carnivores lusting after his flesh. He is, in fact, when
unarmed a singularly easy kill, well within the capacity of a sick or ageing lion. More-
over, lions are by nature indolent; and should by chance even healthy specimens
discover how almost effortless a man kill is, they may well elect to continue exclusively
widi the same easily-obtained diet. Man-eaters have, indeed, been known to carry
on their destruction week after week, causing the death of dozens of human-beings ;
and there have been several cases where villages or districts have had to be abandoned
in the face of a persistent and relentless killer. Development projects opening up hitherto
unpopulated areas have sometimes been brought to the verge of failure.
Next easiest to man himself comes domestic stock, which, often designedly kept in
confinement, is thus unable to escape and is, in any case, ill-acquainted with the struggle
for existence in the outside world. In West Africa the cattle, sheep and goats of nomadic
herdsmen are particularly vulnerable being, in this case, quite unprotected by any
fencing; but though from time to time losses occur (Howell, 1968) the lion population
is m most areas so limited that predation does not constitute the constant and serious
threat that it does in odier parts of the continent. Even where dense and high fences of
thorn trees have been built around more permanent farmsteads lions have commonly
been known to leap or scramble over them and find their way out again burdened
with a heavy carcass.
To sum up, idiosyncratic choice may possibly enter slightly into the question, but
what lions live on, m fact, depends primarily on availability, and secondly on their
own degree of skill. The former is basically determined by many ecological factors
and may vary with the seasons; the latter depends on age, experience, inherent abihty
and the state of health. As Schallcr (1972a) has pointed out the prey lies most commonly
111 the middle range of weight, small animals being rarely worth the effort, while
really large species may well be too strong to tackle successfully. The latter difficulty
may on occasions, but rarely, be overcome by co-operation m attack. There are numer-
ous well-authenticated accounts of two or more lions entering into vei-y prolonged
struggles with adult elephants or, more commonly, hippos; but even then they may
not succeed. Goodwin (1953) mentions the case of three lions attacking a hippo but
being compelled to let go when their intended victim dragged them into the water.
Kruuk & Turner (1967) thought that lone lionesses tended to kill smaller game than
lone lions.
The methods by which lions actually bring about the death ot their victims necessarily
vary with circumstances; and there is, moreover, some evidence tliat there may be
different fashions in different areas depending partly on the type ot prey most com-
monly to be tackled in a district, and possibly partly on a continuing local tradition
handed down through the instructional influence of pride mothers. Small prey, of the
PANTHERA LEO 477
nature of the slowei-moving mammals or grouiid-haunting birds, arc struck and
stunned or perhaps killed outright with a single blow of the paw followed if necessary
with a quick bite. Swift-rimnnig ungulates have to be dealt with in quite another way;
and even here it is obvious that there must be differences since an antelope of the size
of a rccdbuck or oribi must collapse at once luider the sheer weight of its attacker
whereas an animal of the size and strength of a hartebcest or watcrbuck is not so
immediately affected by bulk alone. The essential fust move in a kill is to make effective
contact with the victim, and this the lion brings about by a powerful leap, either standing
from ambush or as the chmax of a short approach run. This, of course, is quite different
from the cheetah's method (see page 502). Li dealing with the larger ungulates the
hon generally aims to land not squarely on its victim's back but a Uttle sideways upon
the shoulder and flank, the hind-claws deeply embedded in the latter, one fore paw
clutching the shoulder or neck on the far side of the body, the second one the chest
or neck or, often, seizmg the muzzle and twisting the head round. In the case of a run-
ning chase this may cause the prey to stumble and fall, and with the speed at which the
crash takes place, the angle of the head and the extremely heavy weight on the ante-
lope's back may well result in the latter breaking its neck. In the case of a standing leap
on to a more or less stationary prey, or if, as the result of these tactics, there is no
immediate collapse, the lion sinks its teeth into the neck vertebrae and spinal cord.
Once the prey has come to the ground the lion, instead of the neck bite, seizes the
throat of its victim and brings about death by suffocation in the manner of a cheetah.
Such is the standard pattern of cffectmg a kill; but there are, as always, exceptional
happenings. Without becoming anecdotal it is not possible to detail these but the
following briefly summarizes some of the commoner or more interesting observations
that have been made. With medium-sized prey which draws sufficiently near to an
ambush lions sometimes do not trouble, or fmd it necessary, to spring, merely rising
onto their hind legs, seizing the animal with the forcpaws and clawing it to the ground
or giving it a bite in the cervical vetebrae. Hamstringing, that is to say severmg the
vital tendon of the hindleg, is a well-authenticated method of incapacitating such large
and powerful prey as a fully-grown buffalo or eland, or even immature elephants.
This may exceptionally be effected by a spring from ambush of a single hon but more
usually calls for the co-operation of two or more attackers, some to occupy the head
end of the prospective victim whilst another fastens its teeth into the vital place above
the hock with a severing bite.
Possibly the most interesting variant in kilhng is recorded by Eloff (1964). This is
seemingly employed only by the lions of the Kalahari (Botswana) but is worth a glance
here with a view to stimulating careful observation in West Africa. In this subdescrtic
region of southern Africa the lions leap not upon the fore but upon the huid quarters
and, as dissection has shown, by a combination of their own oppressive weight and an
upward jerk of the victim's haunches break its back at its weakest point, that is between
the last lumbar and first sacral vertebrae, snapping the spinal cord. Death is finally
brought about in the usual fashion by a bite in the throat. This method is chiefly
used with gemsbok, but horses, donkeys, cattle, blue wildebeest and eland have been
observed killed in a similar way.
478 Tlin CARNIVORES OF WEST AEUICA
Rc.il co-opcr.uion iii killing, it not in liuntmg, certainly often takes place. The
incident observed by Schallcr (1972a) and mentioned earlier (page 475) may be cited
as an example. In this, five lions worked together in bringing about the death of a bull
buffalo that had already been badly mauled by another pride and was in no condition
to put up any resistance. Nevertheless, the moves used by the lions formed a deliberate
co-operative sequence. The first grabbed the victim by the rump; a second placed a
paw over the back and bit the shoulder, bringing the buffalo to its knees. Two lions
then pulled him onto his side, and, by manipulating a leg, turned him fully onto his
back: a third bit the victim m the throat, whilst the fourth held the nose and mouth
closed. The fifth lion did nothing very much. The whole affair took about 25 minutes;
death by suffocation resulting in about 10 minutes.
Kills are sometimes multiple, a pride frequently killing more than one of the lesser
antelopes at a time. Kruuk & Turner (1967) mention a lioness which killed four gazelles
in one morning, two of them at a single rush. Achievement of a kill is by no means
always so easy and there are accounts of prolonged struggles with powerful prey which
may last as long as an hour or two during which the victim is lacerated by tooth and
claw all over its body before finally succumbing, exhausted though not actually
successfully bitten in a lethal spot. But an attack may go awry much earlier than this
and never achieve a kill. If an immediate kill is not made it ma)-, indeed, result in the
death of the assailant from a forceful kick in the skull or the penetration of the body by
a powerful sweep of a horn. Even porcupine quills have been known to be responsible
for the death of a lion. At best, lions arc far from being infallible killers. It has already
been pointed out above how Schallcr (19723) reckoned that the success of stalks might
be as low as 17 to 19 per cent for a single lion, increased to only 30 per cent by co-
operation. The success rate for a shigle lion running at its prey he found to be as little
as only 8 per cent. From the prey's point of view diere is obviously more safety in
standing clear of cover than grazing on the edge of the herd where an approach by an
aggressor is far more likely to meet with success.
After a successful kill the lion or lions may be content to make a meal at the site in
full view in the open or may drag the carcass to the protection of the hght cover of a
tangle of shrubs or of tall grass. This in the case of large and heavy prey may call for
the exercise of tremendous muscular power. The first step in feeding may be to lick
up any blood that is available; the belly is then ripped open, the intestines clawed out
and sometimes, but by no means always, buried or at least covered by having earth
scraped (wer them. Sometimes they are eaten; but always the tit-bits first consumed
seem to be the liver, kidneys, spleen, lungs and heart. These disposed of, a start is made
on the haunch, flank or breast. If there are any soft bones, such as ribs, they and the skin
are eaten together with the flesh; but strong leg bones and the like are left. Behaviour
to others during feeding varies very much since all lions possess difTercnt temperaments.
Like odier animals they may utter protective snarls while feeding and strike out at
other lions which they may consider to threaten their own particular portion of the
meal, or at hyaenas or jackals if these are foolhardy enough to attempt a theft. But
while a number of lions or lionesses may feed together oft a corpse — Goodwin (1953)
observed six males to share one without quarrelling — it has been noticed that the fem-
PANTHERA LEO 479
ales, although they have done the killing, may have to wait until the pride male has
satisfied himself (Estes, 1967). The attitude adopted towards the young seems to vary
considerably; but it has been said that lions are more tolerant towards them than are
the lionesses, which may drive ofFhungry cubs and keep them waiting (Schaller, 1972a).
Lions, provided there is abiuidance, tend to gorge themselves at a meal until they
arc almost immobilized; their bellies hang low and heavily swing firom side to side as
their owners walk. At such tmies neighbouring antelopes become tully aware that they
themselves then stand in no danger whatsoever. If the kill has been a small one it
may be entirely consumed at a single sitting by the lion or pride responsible; but if it
is larger and not fully disposed of the remains are often carefully guarded and used
for a second, or even a third, meal. This is more easily achieved when a pride is con-
cerned and turns can be taken to sleep, drink or dcfaecatc; but even a single lion may
stand guard over its kill for a long time. There are many thieves ready to pounce
upon a deserted corpse, other felines, hyaenas, jackals and vultures. But a guard, or at
least a successful guard, is not always kept. A single lion, having satisfied his immediate
wants, has no recourse but to stand guard itself; and this is not always continuously
possible for it must desert the carcass at least temporarily to drink or to dcfaecatc.
Further, a single lion may be driven oft by a determined pack of hungry hyaenas,
especially when he himself is well-fed and consequently lethargic and less aggressive.
The problem of the lone animal may sometimes be increased by its own action;
as already mentioned, Kruuk & Turner (1967) observed a solitary lioness to kill four
gazelles in one morning, two of them at a single rush; but she ate only one. These
same authors state that lions may stay for days with a kill, keeping hyaenas at bay and
utilizing it to its utmost. The lone hon's problem is somewhat reduced by the fact of
pretty rapid digestion and the ability, therefore, to make a second meal after no great
interval.
Water is a very essential accompaniment to the lion's meal, and always immedia-
tely after feeding, or sometimes even in the middle of a long and heavy feast, these
animals visit a stream or pool and drink deeply. This, when a pride is concerned, they
do singly, not all together, so that there is always someone to guard the remains of the
kOl against marauders. Lions often drink at other times too, perhaps in the evening
before setting out to hunt, or, more regularly, at the end of the night before retiring
to rest during the day, irrespective of whether a meal has recently been taken or not.
After feeding, lions clean themselves up from blood left on the fur by hcking, cat
fashion. One lioness may do tliis to another lioness's face where it is more difficult
for the animal's own tongue to reach; but with a lion the encircling mane makes
attempts at cleaning up more difficult. Some cleaning of the fice and coat is also carried
out by rubbing with paws or on the ground, or rolling on the back.
How destructive are lions; how often do they feed; and how much do they eat?
These arc all questions wliich have only recently begun to be seriously studied; and
the answers arc still not very clear. It has already been pointed out above that Hons
can scarcely be regarded as highly efficient hunters in view of the fact that their success
rate, even acting in consort as a pride, was found by Schaller (1972a) to be no greater
than 30 per cent. They are, indeed, at least in East Africa, dependent to a vcr)' great
4^0 THE CARNIVORES OE WEST AFRICA
extent on tlic energy and skill oi otlier predators, notably the spotted hyaena, as Kruuk
and others have revealed (see page 475). The situation is far from straightforward:
lions may steal their meals from hyaenas, but the latter may perhaps sometimes make
ruse of the lions to effect a kill (Estcs, 1967). Careful scientific observation is only just
beginning to replace hearsay and tradition, and we are merely on the fringe of apprecia-
ting the complexities of predation m Africa. One of the most unexpected discoveries
is the extent of the reversal of the long-held belief of the roles of the lion and spotted
hyaena as noble killer and despicable sncak-thicf: Schallcr (1972b) found that of 63
carcases on which hons were observed feeding no less dian Si per cent had been killed
by hyaenas. This was for the Ngorongoro Crater (Tanzania) ; it is by no means certain
that a comparable situation apphes 111 western Africa, where the conditions of vegeta-
tion, abundance of animal life in general, and the concentration of ungidate herds in
particular are almost everywhere in that region vastly removed from those existing
m the east.
These are questions demandnig local research; as, too, with Schaller's findings
(1972c) on the differing effects of predation in general on the limitation of species:
namely, that in Ngorongoro it was of no consequence for buffalo, nor an important
factor for wildebeest, but has considerable effect on zebra and gazelle populations.
These are matters which, with the greater interest now taken in the preservation of
wild life, including increasing populations of lions, call for closer investigation and
imderstandmg m West Atlica. It has been pointed out carher in diis work that pre-
dators are very quick to perceive any weakness in prospective prey, due to sickness
or injur)', and often select such subjects for their victims as easier to secure than animals
in full vigour. Thus, while hons and other predators cause some destruction amongst
the ungulates they not only preserve a reasonable balance of nature in a given locahty
and help to prevent overgrazing but, m fict, as Schaller (1972c) points out may very
well actually benefit the prey species by weeding out the sick, old and generally less
capable indviduals, keeping the herds healthy and alert.
To get down to figures, several esrimates of how much lions cat m the course of a
day or of a year have been made but vary widely; but some sort of an average can be
arrived at as at least a better guide than a pure guess. The smallness of the amount of
annual destruction attributable to a lion will probably come as something of a surprise
to those who have given little riiought to the matter. Figures derived from animals
kept in captivity are no veiy good guide since such conditions call for the use of
relatively little energy. In these circumstances they may consume something in the
nature of 4 to 6 kg of meat a day. In the wild, however, a Hon may need 6 per cent of
its own weight a day (Krumbiegel, 1953), that is to say for a large lion something like
10 to 12 kg. But the matter is not arranged quite like this, for wild lions more often
than not feed somewhat irregularly, gorging themselves for one, two or three days,
then going for as long without killing or eating. At such a feast a healthy fullgrown
lion may get through as much as 20 to 30 kg — and then not very long after come back
for more. Perhaps a better way of looking at the question is to estimate how much
destruction a lion may cause amongst the ungulates in a year. No exact eiunnerations
over long periods have been possible; but a number of authors from observations
PANTHERA LEO 481
made iii eastern and soutliL-rn Africa have given fairly closely reasoned estimates
based on the short-term behaviour of a pride. These in sum boil dowii to the likelihood
that a single lion is responsible for the death of something between a dozen and a score
of prey animals in a year. This, of course, is in a way fairly meaningless since the
victims vary considerably in bulk from, say, a large bull buffalo or eland to a gazelle;
but it is intended to represent animals of average size from the mixture of prey upon
which lions commonly feed. Surprisingly low as it is it would be still smaller were only
large prey such as wildebeest or zebra continuously concerned. On the other hand this
mean figure for animals destroyed annually must almost certamly be somewhat higher
for West Africa since large prey species are there relatively uncommon, and much
greater dependence must rest upon the smaller sorts of antelope, warthog, and probably
a good deal of much lesser fry. Solitary lions, because of the difficulty with which
they are faced m guarding their kills, almost certainly feed less efficiently than a pride
and are forced into killing at a higher rate. But what they themselves do not consume
of their kills goes to satisfy other predators, so that lu sum the total of destruction of
prey species works out at much the same. It is said that man-eating alters a lion's whole
attitude towards guarding its kills, and that it never returns to the remains of a human
corpse, or for that matter to any other once it has taken up this habit (Goodwin,
1953).
Lions are large and powerfid enough not to fear attack in the ordinary course of
events from any single animal other than man. Nevertheless, they do on occasion fmd
themselves faced with dangerous opponents and are sometimes killed by them. Their
commonest enemies in the competition for food are spotted hyaenas, though in this
struggle the lion stands m no actual danger of its hfe. The conflict is one which turns
in favour of one side or the other according to circumstances. It has already been
mentioned above that a determined and angry pack of hyaenas may scare a lioness off
her kill — or sometimes more than one lioness; but Estes (1967) once wimessed how a
fully-grown male lion could restore the situation and drive the hyaenas from their
stolen meal and allow the cowed lionesses to return — and, indeed, even permit them to
eat their fdl before he himself fed. Yet the conflict may start earUer than this especially
where large prey species are concerned. Spotted hyaenas may closely trail a hunting
lioness, carefully following her at a distance as she, her attention concentrated on her
prospective victim, stalks her prey unaware of their interest in her. Should her ultimate
attack prove immediately successful in bringing her prey crashing to the ground the
hyaenas rush in and, if she is really sohtary, drive her from her kill; but if she leaps
upon the prey's flank, grasping its head, and the victim does not faU at once but is
strong enough to continue galloping over some distance the hyaenas may close in,
tearing at its hindquarters until the honess becomes so disconcerted at this unlooked-for
turn of events that she lets go and jumps clear (Estes, 1967). The prospective victim
would then be followed by the hyaenas to which, partially disabled and exhausted
as it must surely be, it faUs an easy victim. Thus hyaenas may at times make use of
the superior strength of the lion; but the reverse side of the picture, detailed earlier
herein, must not be forgotten, where the lion, apparently far more frequently, makes
use of the energy and hunting skill of hyaenas.
482 Tin; CARNIVORES OF WEST AFRICA
Apart from the cvcr-prcsciit competition ot hyaenas, lions may in their search for
food find themselves in ditiicukies from other sources. It attack on some of their more
powerful prey — the larger antelopes, buffalo or zebra — is not executed with sufficient
surprise or skill, as may happen with young and relatively inexperienced lions, the
aggressor may fuid itself impaled by a horn im- badly injured or even with its skull
fatally fractured by a powerful kick. Baby elephants have been attacked by lions,
sometimes successfully, but sometimes with dire results for the attackers from an
infuriated cow coming to her offspring's defence. Estes (1967) records a case of a
lioness that made an attempt upon a rhino calf and was killed by the mother. Less
spectacular than such battles but no less fatal for the lion is when a young animal
not yet in its full strength drinking at a stream is seized in the muzzle by a crocodile,
dragged into the water and drowned. Such reptilian attacks do not always succeed.
As regards its attitude towards other animals not strictly in the prey class, a lion
may on occasion attack other large carnivores. This as a rule is in response to some real
or imagined threat, possibly to its cubs; but need not always be so. Guggisberg (1963)
relates how a family of ten lions came across an old sleeping cheetah and killed him.
Edmond-Blanc (1957) cites the case of a lioness, probably m defence of her newly-born
young, attacking a fully-grown leopard; and others have noted much the same sort
of thing. Their attitude towards their own kind is mostly remarkably tolerant, but from
time to time lions and lionesses have been known to kill each other, mostly in cases
of trespass (Guggisberg, 1963; Schenkel, 1966).
This raises the question of territory. Despite the amount of observation that has been
devoted to lions nothing very definite lias so fir emerged on this matter in any part
of Africa. The tact is that, apart from the difficulty of checking such ill-defined boun-
daries as may be involved and of estimating large areas, the matter is not entirely straight-
forward in this species, where fictors of sex and varying social habits play important
parts. Guggisberg (1963 : T41S) expressed the view that there is a fundamental difierence
between the territories of female and male lions; and this is so since lionesses are
appreciably more sedentary than lions, which arc often frankly nomadic, ranging
over relatively large areas. But there are other complexities. There is, for example, the
difference in requirement between a single lion, a single lioness, a lioness with cubs,
a pride of lionesses, a mixed pride, large or small. There is also the very basic factor
ot the density of prey species in a given area; quite obviously where, in parts of East
Africa, there are abundant herds of game there is no need for lions to concern them-
selves with large areas such as would have to be hunted over in, say, much of West
Africa where prey may be reckoned in scores rather than in hundreds or even thousands.
It is clear that estimates of territorial size made in Tanzania can have little application
to the region covered by this present work. Further, where the density of lion popula-
tion is itself high there is a need for the limitation of hunting territory and the fairly
strict observance of boundaries; in West Africa, where by comparison lions are few
and far between, a pride may seek for food quite undeterred over very wide and pos-
sibly undefined areas. Such a situation may change in the course of years as both game
and predators become more abundant and concentrated in the restricted areas of
national parks.
PANTHERA LEO 483
However, for what they arc worth as some rough indication of the sort of position
found to exist in other parts of the continent, these figures may be quoted: Krumbiegel
(1953) recorded that in the Kruger National Park (South Africa) a lion needed about
15 sq km of territory; Schenkel (1966) estimated the home range in Nairobi National
Park as being from 25 to 50 sq km; and Schaller (1972a), for Ngorongoro, Tanzania,
found that a pride of lionesses might have a territory of anything between 20 and 400
sq km, and that nomads might roam over as much as 4000 sq km. As this last author
points out, one of the great benefits of a fixed and not too large territory is the acquisi-
tion by all members of the pride of an intimate knowledge of its topography, and this
gives advantage in hunting not only m knowing the favoured grazing grounds, drinking
points and probable movement of herds but also in becoming closely acquainted with
every scrap of cover, major or minor, which the lie of the land and vegetation offer.
Where territories exist their boundaries must be demarcated and guarded. Scent-
marking by urination is the method used by lions. Schenkel (1966) noticed that this
was effected 111 two ways; by squirting urine upwards into the branches of bushes
where it would remain evident at head height, emphatically registering ownership
at that particular point of the boundary; or by squirting it downwards onto the ground
and trampling in it so that the paws subsequently leave a trail of ownership along the
path taken. A less permanent but more impressive way of declaring occupancy is for
the pride leaders to stand on some slight eminence and roar; and this, indeed, is
probably the main purpose of the lion's famous deep-throated, far-reaching roar —
not to annoimce that the animal is on the hunt or to frighten game, as often said;
for this, by alerting its prey would, in fact, deprive the lion of the most essential clement
of its attack, surprise.
Territories are not always strictly respected, and may sometimes even overlap
(Schaller, 1972a); but intruders are frightened off if possible by aggressive behaviour,
though as a last resort recourse may be had to fighting and possibly killing. Awareness
of rightful possession gives to the owner a feeling of confidence often obviously
cxliibited in its bearing; and the reverse is true of intruders, which arc apt to behave in
a guilty, morally inferior manner which as often as not renders their expulsion without
actually fighting easy; or, should it come to combat, victory over them more readily
achieved. In this connexion Schenkel (1966) observed that while lions may not avoid
territories belonging to other prides, if they do trespass they become wary to an
unwonted degree. He once observed two lionesses and a half-grown male to make a
kill within the territory of another pride; but they made no attempt to start eating
until they had carefully scrutinized the surroundings to make sure that the real owners
were not about in the neighbourhood. When lions acquire the wandering habit,
mostly males, they often travel very far indeed, though, in fact, little positive research
has been done into this. But they appear to be received into other prides mostly without
much question; and it would seem that this must depend upon there being females
in season ready to welcome them as mates. There are also, though much more rarely,
nomadic females; but these make poor mothers.
It has been said above that the main purpose of the lion's roar is to assert ownership
of a territory; but there is, in fact, some disagreement over the use of this the best
484 THE CARNMVORKS OF WIIST AIHICA
known ot thi- lion's vocal iz.uions. Goodwin (i9:J3), for example, wrote that deep
guttural roaring starts at sunset and continues up to the kill; others have told of roaring
continunig throughout the night; and some tif its being used after a kill as a paean of
triumph — or more probably to announce success to the rest of the pride and gather
them to the meal. Wliile roaring may well be carried out in the early evening to
warn other lions away from a hunting territory its use as a prehminai-y to the hunt
itself seems improbable since it would serve to alert all game within a considerable
distance. The roar, which is performed with the head lowered so that the sound is
deflected upwards off the ground, may in fact carry a mile or even two according to
weather conditions and topograpliy. Other vocalizations are a grunt uttered when
charging: and a similar sort ot repeated grunt in five or six stages of crescendo under
the influence ot mild excitement, sometimes ending in a sigh due to the eventual intake
of breath, hi defence of itself, its young or its food the lion utters deep warning growls
which, if they prove mettective, may be increased to a semi-roar. The teeth are bared,
the lips being drawn back, and the ears laid flat and turned so that, m the female, the
black marks on their backs are exposed, hi a previous section it has been explained that
hons arc enabled to roar loudly because of the specialized elastic bone structure in the
throat permitting a 50 per cent expansion of the vocal chords. This structure inhibits
the rapidly vibrating piurr of the smaller cats, though a throaty rumbling can be
achieved. In areas m wliich they sense themselves m considerable danger from per-
sistent hunting by human beings — as for example in the case of man-eaters — lions
cease to draw attention to themselves by roaring.
When the lion goes about affairs of an unimportant nature it does so at an easy,
sedate walk; when in more of a hurry it breaks into a relatively rapid trot, which
it can if necessary maintain over some distance. Both the easy walk and the business-
like trot may, as well as the slow, painstaking slink, be used in approaching prospective
prey; the trot being employed mostly in the early stages when at a distance, but also,
more rarely, at closer quarters when the cover is so favourable that the intended victim
is 111 no position to detect rapid movement. The lion's most purposeful gait is the
gallop. This is carried out in the usual feline fashion of both fore and hmdfeet used
approximately as pairs, alternating in their contact with the ground. It may reach a
speed of some 55 to 60 km an hour but normally cannot be maintained for much more
than about 100 metres, that is to say for 6 or 7 seconds. As the speed of most of its
quarry is of the order ot 65 km an hour the necessin,' of surprise and of accelerating to
maximum speed before the prey has had time to react to attack is obvious.
The fmal approach to the victim is a spring, a movement calling for the lion's
extreme muscular power, executed with the claws extended, as with an athlete's nailed
shoes, leaving their marks obviously in the soil at the point of take oft. This, of course,
is a running jump: as a standing jump it is said that this species can cover 12 metres
in lengdi and reach a vertical landing place nearly 4 metres high (Guggisberg, 1963).
There is good evidence that in cases of necessity, such as withdrawing with killed prey
from a fenced enclosure, a lion can leap a couple of metres, possibly m two stages,
carrying a corpse weighing perhaps 300 kg, the body, at least sometimes, supported
across its back.
PANTHERA LEO 485
The spoor of a lion is, of course, larger than that of any other African carnivore but,
strangely in view of the great discrepancy in body size, not as much bigger than that
of a leopard as might be expected. The feet are, indeed, short, broad, compact and
difficult to spread; and m the forefeet the webs reach almost to the distal ends of the
digital pads, but m the hmdfect are less extensive. The spoor of the sexes can be told
apart because in lionesses the fore and hindfeet are subequal in size, whereas in the
males the forefeet are bigger than the hind. In a large lion the mark left by a forcpaw
may measure some 120 mm across and about no from front to rear; hindpaws and
females are about 10 mm less. The middle subdigital pads show in the footprint as
long narrow ovals, 45 X 20 mm, the two outside ones shorter and broader, 35 X 25
mm; the central pad is roughly triangular with very blunt roimded corners, and a
hind-margin that is often concave. It measures in the male roughly 80 X 60 mm. In
soft ground the print left by the spreadmg foot of a spotted hyaena measures not a
great deal less but differs markedly m exhibiting long claw marks, absent from the lion
except at the point of take off for a leap.
Lion cubs are tolerably good chmbers, starting at a fairly early age on trees with
sloping trunks in the shade of which their parents may be resting. While it is a common
thing for adult lionesses to climb to, and lie along, the lower branches of small trees
it is often held that they never climb higher, being unwilling, possibly, to trust their
heavy weight to the smaller upper branches. However, in this as in other matters of
behaviour lions differ individually; and while some, doubtless, make no attempt to
climb, others have been seen relatively high up, as much as 10 metres above ground.
This may be sometimes apparently only to survey the coiuitryside for game, cheetah
fashion ; but cases arc recorded where a lion has climbed high in order to rob a leopard's
larder, Schaller (1972b) having witnessed no less than three such instances. Lions not
infrequently stand upright on their hindlegs at the base of a tree, stretching their fore-
legs up the bole, scratching at the bark in the action commonly known as "sharpening
its claws" — though this seems more likely to blunt the fine points than the reverse.
Lions are also competent swimmers, taking readily to water if necessary and crossing
rivers 30 metres wide without difficulty.
Lions may occasionally be monogamous but are much more commonly polygamous.
A hon may remam faithful to a lioness, or to the lionesses of a pride, for some years;
others are given to wandering far afield seeking new mates, or possibly to estabhsh
themselves for the first time as eligible partners. Visiting lions may sometimes become
involved in scuffles, which occasionally may develop into something more serious
resulting in the death of one of the contestants; but within the pride there is httle
or no interference by the others with the mating of the dominant male. Despite various
general claims to the contrary, no favoured period of the year for mating has been
clearly estabhshed. Lionesses can come into season about every three months, or some-
what less, and remain on heat for some four days, the last of which appears to be the
critical one for conception. During this time they emit a powerful odour and announce
their readiness for coupling far and wide by spraying their urine over shrubs, grass
tufts and the like. Such advertisement may not be of much moment within a mixed
pride but is important in prides consisting of females alone or accompanied only by
GG
486 THE CARNIVORES Of WEST AFRICA
their last Utters. Apart trom any preliminary dispute there may be for her possession
there is little display or noise; the chosen male follows his prospective partner about
closely for some time, nose to tail, and occasionally licks her. Eventually she lies down
on her belly in the crouching position, the lion straddles her with his forelegs and
effects penetration from behind. This may take place m a secluded spot or in full view
of the pride; and it may happen by day or by night. The act is repeated three or four
times ill fairly rapid succession at intervals of from lo to 20 minutes. According to
Dadic (1947) the lion bites the lioness in tlie back of the neck only at the moment of
ejaculation. He utters occasional growls.
The period ot gestation is generally between 105 and loS days, but as long as tl2
days has been recorded. This discrepancy might be accounted for by the observation
having possibly been reckoned from a mating on the first day of a lioness's four-day
season whereas fertilization takes place only on the last day. As her tune approaches
the prospective mother seeks out a secluded spot a little removed from her pride
amongst dense undergrowth or rocks should these be available. Here she gives birth
usually to between 2 and 5 cubs, 3 being perhaps the commonest number. According
to Krumbiegel (1952) as many as 9 m a litter have been known. K4itcliell ct al. (1965)
foimd the sex ratio to be i male to 1-73 females.
Schenkel (1966) furnishes details of the early days and development of wild litters
111 Nairobi National Park: and Hopkins (1968) a seemingly unique account for West
Africa of hand-raising orphaned cubs 111 Nigeria. There are numerous other less detailed
held observations from East and South Africa; and records of birth and development
in zoos, though the latter may not, of course, necessarily be typical of what takes place
amongst wild populations. The cubs may apparently be born either with their eyes
closed or with them open (Guggisberg, 1963); but as information regarding this
obviously emanates almost entirely from zoos the latter condition may possibly be the
outcome of unnatural circumstances. According to Goodwin (1953) in the Serengeti
the eyes are closed at birth and do not open luuil the 6th to 9th day. When they do
open they are blue and cloudy and obviously not fully functional for some time. The
length of the new-born cub is of the order of 300 mm; its weight between i-o and
1-7) kg. The head is rounded, the ears small, the tail relatively shorter than in the adult,
the coat sott and usually heavily dark-spotted; but cases are known wlicre the spots
have been almost completely lacking.
Hopkins (1968) furnishes details of early development as exhibited by a wild West
African lion cub hand-reared in Nigeria. In this animal eruption of teeth took place
111 the mandible a few days before the corresponding ones in the upper jaw. At 3 weeks
4 incisors were visible top and bottom. At this age, too, the cub started to drink water
and was steady on its legs but could not walk fir, though at 6 weeks it could run. The
lower canines appeared in the 4th week; and at about the same time the eyes started to
lose their blue hazy look and become brown with a black pupil, the whole process of
clearing taking about a couple of weeks. The voice at this stage was a staccato mew.
At 8 weeks the first, lower, premolar developed and the cub evinced an interest in
solid food; at 12 weeks it tore at chiniks of meat, though the carnassials did not, in fact,
appear until the 15th week. The black end to the tail also developed rapidly at this
PANTHERA LEO 487
time; but a photo demonstrates that there is nothing of the adult contrast between long
tuft and close-haired shaft but that the whole structure is still woolly throughout.
Accordmg to Hopkms the growth curve was fairly regular up to a weight of 1 8 kg at
19 weeks. At the age of 4 months the cub could run fast and leap.
So much for physical development in circumstances of careful hand-rearing ; some
data regarding social and behavioural advancement in the wild can be assembled from
the observations of various authors. A few days after giving birth the mother, miless
she is a lone female, rejoins her pride for a period or periods every day; but the cubs
remain deeply hidden and the other Hons make no attempt to go near them. She is,
of course, compelled to leave them from the necessity of feeding herself, which she
may do by taking part in pride activities or, if she is solitary, himting for herself.
Such periods of absence are times of danger for the helpless cubs; for no matter how
well hidden they may be their scent may give them away to other maraudmg carni-
vores, perhaps especially hyaenas. This period during which the cubs are daily, or
nightly, at high risk through the enforced absences of their mother may last for some
weeks. It may be added here that while most lionesses are excellent mothers there are
unquestionably some that are very bad and deliberately desert their young.
At the end of three or four weeks, when the infant lions have grown sufficiently
strong to be able to get about, the mother on returning from her excursions to the
hiding place summons her cubs with a gentle roar, and they hasten to welcome her,
scrambling out of the nest uttering high-pitched answering squeaks, and when they
reach her rubbing their heads against her muzzle and legs in a greeting ceremony
(Schenkel, 1966). She in her turn hcks them and, in the early days, permits them to
suck, sometimes lying actually on her back to allow them to do this; but at a later
stage of advancement she may present them with some small animal she has brought.
They tlien play together in the open, the cubs amongst themselves with mock-fightmg,
tumbhng each other over or wrestling; or with their mother by clambering over her
or playing with her tail, which she flicks from side to side, wliile she responds with
soft taps of her paws or pokes with her nose. When the yoiuig are feeding from the
breast they mould it with their paws in a manner commonly seen in domestic cats
and dogs.
According to Schenkel the cubs are introduced to the pride when they are about
10 weeks old, and they arc then treated by the other lions with tolerance and apparent
affection^thougli response to their playful advances may be colder on the part of
adult lions than by the lionesses and younger members. Thenceforward, during their
mother's absence they may be guarded for some of the time by other females; and
such care may extend also to suckling them by such as are in milk. Indeed, cubs from
two or more different litters, and perhaps of quite different ages, may feed side by
side off a single lioness. The amount of communal care is, indeed, remarkable and at
this stage is almost completely devoid of jealous squabbling. However, the mother
always returns her cubs to their own hide-out, the original birth place being, according
to Schenkel (1966) retained as this for some two or three months, the location of the
retreat thereafter being changed every four to six days. This last has the advantage not
ordy of putting possible predators off the track but, probably more importantly, also of
488 Tlir CARNIVORKS OF WEST AFRICA
gr.idiully widening the young lions" knowledge of the topography of" their pride's
territory. Two or three weeks after their introduction to the pride, when they have
become well acquainted with the others as well as stronger from rough-and-tumble
play with yoiuig adults, they accompany their elders on hunting expeditions and
thus, from the early age of about three months, start to observe essential techniques
and acquire a wider knowledge still of the terrain. At a kill they are almost always
allowed to be amongst the first to feed; but the adoption of a meat diet does not mean
the cessation of breast feeding, and it is no unusual thing for young lions to make a
heavy meal at a kill and then persuade their mother to give suck. Weaning starts at
about 6 months; but cubs commonly go on attempting to suck up to perhaps lo
months, though the milk may have dried up some time before this. Play at this stage,
m conjunction with the other members of the pride, becomes more purposeful, losing
its mere kittenish abandon for actions essential to success in later life: crouching both
to hide ar.d to spring, sudden pounces upon the back, aimed sweeps of the forepaws,
and biting m the neck.
Development is, of course, continuous; but from about the loth month important
changes are observable. At this time the juvenile pattern of spots, save in very excep-
tional cases, has largely vanished from the back and flanks; and when the cubs are a
year old the milk dentition is replaced and the whole body begins to throw off its
childish look and take on the imdoubted appearance of a yomig adult. Up to this
period, although there has been some freedom to come and go amongst the pride
and to take part in its activities, this has nevertheless for the most part been very much
under the eye and protection of the mother. From now on, though she still takes them
with her and tries to inculcate upon them the correct techniques of hunting and killing,
they gain ever more and more self-confidencc in this and other matters, becoming
fully independent individuals at about iS months. The lioness may then reniate, or she
may have already done so and be more concerned about the birth of a new litter than
with her existing one. Two htters in one year have been recorded. However this
may be, at the same time as the youngsters gain independence the attitude of the
pride changes towards them; and from being favoured members of the society they
now receive no further special treatment but have to learn their place and show due
respect to their elders. These, instead of allowing them privileged access to a kill, may
as often as not drive them away, compelling them to take a low place m the feeding
order, and even at times to go hungry. It is without question a very trying period for
adolescent lions; one, indeed, which has sometimes fatal results. There is the danger
of loss of strength and vitality through undcr-nourishment at a tune of still very
active growth and large appetites; and of injury arising from lack of expertise in
hunting, as well as from the fierce repulsion by senior members of the pride themselves.
One observer has drawn attention to the number of young one-eyed lions probably
the result of angry swipes by impatient adidts at this stage.
The difficulty of establishing themselves fully into the pride is often the reason
for young lions leaving it and striking out on their own; but their relative inexperience
in himting, apart from risk, may bring such low success that they face feeding difficul-
ties as great as those they have left behind. It is not until they are about five or even
PANTHERA LEO 489
six years old that lions attain their prime, though sexual maturity may be reached at
four. In the males the mane, which may show its first signs of growth at about i8
months, attains a fair size at 3 years but goes on increasing in luxuriance for twice this
time. Thereafter it may darken in colour from the increase in the number of black
hairs in it until at length these may dominate — though of this eventual blackening
nothing is recorded for West Africa. The sex link of this structure is nicely illustrated
by the fact that a castrated lion lost its mane. In captivity lions have been knovm to
live for almost 30 years, though this is an exceptional figure. Of their life-span in the
wild nothing is known but almost certainly never reaches anything like that under
sheltered conditions.
At one time, not so very long ago, the successful breeding of lions in zoos was
regarded as something unusual and a matter for congratulation. In recent years, how-
ever, with a better understanding of basic requirements it has been found that they
are capable of breeding very freely in captivity; so much so, in fact, that their poten-
tiahty in this respect has sometimes become a source of embarrassment. Litters may be
of large size, 5, 7, 9; and the care with which the cubs are treated and sheltered from
harm ensures a survival rate far in excess of what must prevail in nature. There is no
doubt that under natural conditions there is, despite the watchfulness of the majority
of mothers, a tolerably high rate of mortality from predators, accident and disease in
cubs and young adolescents up to the age of two years or more. There may, indeed,
be as httle as a 50-50 average chance of survival to bccoining a parent; but the death-
rate is doubtless higher as regards large litters than where there are only one or two cubs.
The weights of 7 cubs born at Arnhem Zoo in 1964 at the end of 6 weeks were: 2 males
5-5 to 6-3 kg; 5 females 3-5 to 4-8 kg (van Hoof, 1965). In zoos, crosses of lions wdth
other big cats have sometimes been successfully made. Pocock (1908a) records a lion-
leopard-jaguar cross; and one between a leopard and a lioness, resulting in two cubs,
took place m Japan in 1959. No such thing as this is ever likely in nature.
Taxonomy. The generic status of the hon has already been gone into earher in
this work (page 437 ct scq.), and it remains here to consider the question of specific and
subspecific division. It was at one time believed that Asiatic and African lions represen-
ted different species; and at times it has been held that within Africa itself differences
were such as to merit yet further distinction at tliis level. Such opinions are no longer
current, and it is generally appreciated that throughout the major duration of their
history there has been little practical isolation of the Asian from the African lions
(Mazak, 1964b); and that variations of colour or size have certainly neither the degree
nor constancy necessary to warrant differentiation at specific level.
The position at subspecific level is rather more in dispute. G. M. Allen (1939)
accepted 10 African races; but it is now recognised that the grounds on which at least
some of these were founded are very shaky. The characters used in diagnosis have
been colour of both body and mane, together with size, particularly of skull and
teeth; and most modern authors and field observers are emphatic that all these are
highly inconstant. Pocock (1945) writing of the few remaining Asiatic hons, now
limited to north-west Lidia, observed that the variation "witliin the confined hmits of
the Gir forest covers that of the lions of the whole of Central Africa which have been
490 THE CARNIVORES OF WEST AFRICA
assigned to several subspecies largely on differences of colour". As regards morphology,
J. A. Allen (1924) found the teeth of African lions extremely variable in size and form
m both sexes, there being a variation in length and breadth of from 1 1 to 15-7 per cent.
"These statistics indicate that cranial and dental characters are not so stable a basis for
the discrimination of regional forms as has been often assumed . . .".
There is abundant other testimony to the high variability of lions in single localities;
and the present author is not convinced that it is possible to draw useful taxonomic
distinctions between the lions of different areas. Roberts (1951) thought that because
of the hon's habit of wandering local forms do not become stabilized; but if large areas
were taken into consideration recognisable differences do exist. Unless the provenance
is known it seems virtually impossible to assign a specimen certainly to any particular
race. Nevertheless, it can be argued that mean differences exist and have their taxonomic
value. In spite of his findings quoted above regarding the inconstancy of commonly
used morphological characters J. A. Allen (1924) aimed to differentiate the lions of
different localities by their mean measurements. To the present writer this does not
appear very comancing; but it is fair to add, however, that Allen took a widely different
view and, writing of the four geographic races he sought to establish, asserted that the
r\vo extremes of them were "not only widely separated geographically and environ-
mentally, but appreciably in size and so strikingly in coloration that if one had to deal
with them separately, or without the connecting intermediates, it might seem reasonable
to consider them as specifically separable".
Be that as it may, the amount of material available from West Africa is so limited
and so poor that it is impossible to draw from it reasonable conclusions as to the
existence of valid races. Of the two that are commonly held to occur in the region
covered by this present work the following may be said. Exactly what Siiui^ahnsis is
and whether there is any valid difference between it and what Linnaeus described as
ho it is not really possible to determine; and, apart from reputed colour distinction —
notoriously inconstant, and in this case founded on a single skm — Matschic's diagnosis
o{ka}npt:i depends on the comparison of complex and rather artificial ratios in cramal
and dental measurements that have no real taxonomic worth. The five West African
study specimens available in the British Museum exhibit a fairly wide range of colour,
the one constant factor being the bright orangey colour in the manes of two of them.
Jeannin (1936) with wide field experience found himself unable to distinguish races
and asserted that he had come across lions in Cameroun that corresponded to the four
reputed subspecies scne<:;alaisis, kviiptzi, somaliaisis and imusdica. Obviously the position
is such that, at any rate in the opinion of the present author, any attempt to distinguish
West African lions more fmely than as Fills Ico and relate them to reputed subspecies
would be to claim an accuracy and a knowledge not warranted by the existing studv
material and unrealistic racial diagnoses.
However, it may be of interest to record here the appearance of the specunens which
at present exist in the British Museum.
PANTHERA LEO
4,91
B.M. No. 21. 10.7. 1, o- Tambana (Gambia). Sudan or Guinea woodland. This is
basically of a brownish hue but is considerably darkened by heavy speckling with
black-tippcd hairs. There is no mane or elbow tufts; and no reversal of fur on the
back. The skin is m poor condition with large gunshot wounds and slit-hke scars.
This has no study skull; the measurements which arc given below arc those of
a young adult from Tambana (Gambia).
B.M. No. 30.4.7.1, 0. Upper Volta. Sudan woodland. This is very greyish-sandy —
a much greyer skin than any of the others. There is no sign of a mane, and no
elbow tufts ; the dorsal fur is reversed.
B.M. No. 46.387, (J. Tamale (Ghana). Doka woodland. This has a greyish-sandy
colour. There is a not very abundant mane of a bright orangey-ginger colour;
no elbow tufts; dorsal hair reversed.
B.M. No. 48.762, S. Ogoja Province (eastern Nigeria). ? Gumea woodland. This agrees
fairly well in pelage and mane colour with the previous (Ghana) specimen, but
only a mere remnant of the mane, on the side of the face, exists. There are no
elbow tufts; the whole specimen in very poor condition.
B.M. No. 71.1754, $. Nigeria. Vegetation unknown. This is of a sandy colour, a shade
paler, having slightly less reddish tmge than the two previous specimens (Ghana
and Ogoja); perhaps faded since, though in fair condition, it is mounted as a rug.
Skull in the mount.
Table 30: Numerical data for Panthera leo
Gambia, //}'.\' are still apparent from this once regal pastime. It
w as found that cheetahs taken as kittens turned into poor hunters and that real success
could be obtained only by their capture as adults. This is because the young, though
boni with some inherent predatory instinct, have to learn from their mothers the finer
arts of hunting and killing (Eaton, 1970c). This they do partly by continual observation
over a long childhood lasting about a year, and partly by practice deliberately given
to them by their parent. Speed for the chase is inborn; and, as demonstrated by their
behaviour in play, so is the striking down action of the paw, which Encke (i960) first
saw exhibited in cubs of i r to 12 weeks old. But the cautious stalking and the rest of
the sequence up to the seizing of die throat is learned by watching the mother in action
and eventually copying. Kruuk &' Turner (1967) observed a female to put a still living
fawir before her young and allow them to chase it; and Eaton (1970c) records what
appears to be a deliberate segregation of a warthog sow from her litter by a mother
cheetah whilst her own cubs practised chasing the young pigs. It is probably not until
they are about 6 months old that cheetah cubs are allowed to accompany their mother
on a foraging expedition, and certainly not to take any active part until later. They
probably attain full skill at the age of about a year.
The second point that emerges from the ancient Asiatic pastime is that adult cheetahs
are docile, qiute amenable to captivity and handling by humans, lacking the fierce
offensive resentment cxliibited by the vast majority of carnivores taken after the initial
stages of babyhood. While it is true that a cheetah driven into a corner in ultimate
defence of its life may, like any other animal, turn on its opponent it does so in no very
alarmingly fierce way; and, in fact, aggressive behaviour plays exceptionally litde part
in its life. It has already been mentioned that when two groups of cheetahs meet in the
field there is no challenge or fighting, though Stevenson-Hamilton (1947) does record
two cases of fital combat between males. Eaton (1970b) found that even when a young
male is taking over leadership of a group it is done without contest or aggression. It
was noted above that cheetahs put up very little defence of their kills, being easily
driven oR by lions or spotted hyaenas. They are, in fact, sometimes killed by lions
(Hardy, 1959); and, though there appears to be no definite record, it is not unlikely that
the young are snatched by hyaenas even tuider the eyes of the mother. In most animals,
and especially carnivores, the female is bold and determined in defence of her young;
but Aiisell (1963) records a case of a cheetah cub being taken by a man, the mother
advancing threateningly to within a few paces but making no attack even though the
male parent was also present in support. Cheetahs have, indeed, rarely been knowm
deliberately to attack man as lions, leopards and other large cats sometimes do. They
are almost without exception docile in captivity; but the exact nature of dieir relation-
ship with man depends on the degree of association. Where the latter is close and
domestic they can pass beyond the imtial phase of somewhat indifferent tolerance to
ACINONYX 507
friendliness and even sonic kind of affection (Florio & Spinelli, 1967 and 1968). When
brought up with them they will play with dogs.
Can the cheetah climb? It has very often been asserted that because of its blunt
dog-like claws and long rather stiff legs it cannot. The answer depends entirely on the
defniition of climbing, for this problem is solved in different ways. Many animals, for
example squirrels, the domestic and most wild cats, can with complete ease ascend a
tree trunk vertically by anchoring their feet in the bark at each step with their sharp
curved claws. In this sense the cheetah is unable to cimib, at least not once past the
young kitten stage; but given a tree with a sloping trunk and conveniently placed
branches it can, and does, climb or at any rate get fairly high up by an initial leap onto
the stem, a walk up it, a further leap onto a branch, and so on. Trees are, in fact, not
infrequently utilized as observation posts for game, affording a more distant view than
can be gained from hillocks, termite mounds or other similar and often used vantage
pouits. It would seem, too, that look-out trees are territorially marked; for Hanstrom
(1949) observed a cheetah to defiecate or urinate on a branch, to be followed by a
similar action on the same spot by its companion. Cheetahs have been said to "sharpen
their claws" on the boles of trees; but this is most likely nothing more than a muscle-
stretching action, often seen in cats and dogs.
Intense visual concentration on potential prey is one of the cheetah's leading charac-
teristics, both durnig its final cautious approach and, earher, as it makes its initial, more
removed, survey of the possibilities, sizing up relevant factors — wind, distance,
intervening cover, vigilance of the herd, indications of weaker members, and so forth —
before deciding whether a hunt would hold out prospects of success. This observation
may be carried out from the branches of a tree, from a hillside, a hillock or even a
termite mound, or, not infrequently in suitable terrain, merely wliile lying hidden ui
the grass. The cheetah, in fact, spends a good deal of the day lying concealed, watching.
To what sort of shelter it retires at night is not so clear; whether merely curled up in a
"form" in the grass, or in some more secluded spot as a hole in the ground or amongst
rocks is not recorded. And whether a group habitually frequents the same spot for
several nights or is wholly nomadic has never been ascertained.
Nor have actual breeding places been often observed. These are sometimes, and
perhaps generally, deserted terrestrial burrows, though in suitably rocky country natural
cavities between boulders are doubtless used. Nothing has been recorded of breeding
habits in the wild; and strangely, ni view of this animars placid nature and semi-
domesticity, it has proved to be an exceptionally difficult species to get to breed
successfully in captivity. There has been either a refusal to mate or, if coupling has taken
place, an almost total mortality' amongst cubs. Pournelle (1964), for instance, records
that of 4 litters of 3, 4, 3 and 2 only 2 cubs survived. In the last decade or so, however,
more success has been met with and more of this important matter is now known
thanks to the accounts given by W. D. Thomas (1965), Florio & Spinelli (1967 and
1968), and Manton (1970).
The female comes into season repeatedly at intervals of from 7 to 10 days, each
period lasting about 15 days. In the pre-mating phase th.re is close association between
the sexes; but tlie considerable interest that the male then exhibits sinks at once into
508 THE CARNIVORES OF WEST AFRICA
iiidiftcrcncc as soon as oc'strus comes to an end with conception. Copulation has not
been much observed but appears to be repeated during the oestrus period (W. D.
Thomas, 1965). In the cheetah's natural surroundings it probably takes place after dark.
Gestation lasts 91 to 95 days. There may be from i to 4 cubs in a litter in captivir^', 2 or
3 being the most usual numbers; but in the wild, litter size appears to go up to 8, 4
not being uncommon. At birth each cub weighs some 250 to 300 grammes; but increase
is rapid, and Encke (i960) found his to weigh 370 grannnes on the 3rd day; and these
reached a weight of 8J kg at 5^ months. They can start to crawl at the age of about 2
or 3 days; stand up at 10 days; and walk at about 16 days. The date of the opening of the
eyes seems to be widely variable. Florio &; Spinclli (196S) record one litter in which
the eyes started to open only on the lorh day, were not fully open until the 15th day,
and started to focus properly about the 28th day; but in a previous cub from the same
female (1967) the eyes were open on the 4th day. Manton (1970) tells of a litter of 3 cubs
ill which the eyes were open on what appears to have been the 6th or 7th day; while
both Pournelle (1964) and Encke (1960) found the eyes to open after S days. The latter
author noticed that the first pupil reflex became obvious on the 20th day.
The first teeth erupt at about 3 weeks; and meat eating normally starts probably
when the cubs are about a month old. Nevertheless, one cub ate meat regurgitated by
the mother on the i8th day; but though by the time it was 4 months old it was regularly
eating meat and chicken heads it still took milk from its mother and even attempted to
do so at 6 months. In other litters weaning took place a month earlier than this. The
mat of dense grey hair which covers the dorsal aspect of cheetah cubs is moulted
gradually from behind forwards and mostly disappears at about 3 months, though it
persists rather longer over the shoulders and neck. The claws of the young are very
sharp and cat-like, especially those of the forefeet, and can be used effectively for
vertical climbuig. Sexual maturity is attained in males at the age of about 14 mondis.
In females it is somewhat earlier, the first oestrus period being exhibited at 9 or 10
months. After parturition a female can come into season again at the end of 3! to 4
months. It is thus possible for a cheetah to have nvo litters in a year; but whether this
actually takes place in nature is another matter. If it did the first litter would be only
about 7 months old when the second was born, with a further 5 to 7 months training
to undergo with the mother; but there seems to be no evidence of the existence of
parties of young cheetahs comprising two distinct age groups. There is, however,
some reason to suppose that a litter may stick together for a considerable time, with
perhaps occasional absences of a male in order to follow a female whose oestrus scent
he has picked up, and to mate. There is possibly not much actual interchange between
groups. These may consist of all males or of mixed sexes; but it seems likely that
females, once they are sexually mature, tend to be rather more solitary, going apart
to mate and thereafter bringing up their families unaided. According to S. S. Flower
(193 1) a cheetah lived in a zoo to the age of nearly 16 years; but they seldom live in
captivirj- for as much as half this.
The young soon after birth (Florio iSi Spinelli, 1968) utter a bird-like cheep or
whistle. A sinnlarly bird-like sound is made also by adults. Roosevelt & Heller (1915)
record that when they first heard this chirp, from captive cheetalis, they could not
ACINONYX 509
believe that such a sound could come from them and looked everywhere for the bird
responsible for it. Cowic (1957) says that cheetahs call to each other in the field with a
shrill whistle or squeak, more like a bird than a mammal; but according to Eaton
(1970b) they do not appear to vocaUze to attract mates, this probably being sufficiently
effected by the chemical message left in the female's urine. However, he foiuid that a
mother could give orders to her cubs when they accompanied her out foraging. A
low-pitched "ii{;hh" had the effect of making them remain in one place to await her
while she hunted; and a high-pitched "chirp" brought them to the site of the kill.
When cheetahs are content they utter a deep vibrant purr; but when angered or alarmed
they snarl and spit in the usual cat fashion. Stevenson-Hamilton (1947) describes them
as sometimes uttering a cat-hke mew.
Cheetahs frequently walk, at a rather stately pace sometimes with the head up
sometimes with it lowered and pointed forwards. This gait is used for normal travel,
and their approach to game may be initially in this manner. When proximity to their
prey calls for care they lower their bodies somewhat and move more deliberately and
circumspectly, never removing their eyes for a moment from the potential quarry,
watching with deep concentration for the least sign of alarm and for the exact moments
to rcmam stationary or cautiously to advance further. Observers, as so often, differ
remarkably. Stevenson-Hamilton (1947) describes the cheetah as crawling along
"glued flat to the ground"; Eaton (1970c) says that they never crouch; and this latter,
with the proviso that they do to some extent lower their posture and slmk, seems the
more accurate. Again, Kruuk & Turner (1967) say that the cheetah breaks into a gallop
straight from a walk; but Miuidy (1832), giving an eye-witness account of a set hunt in
India, describes the cat as approaching its prey "at a slow, crouching canter". It seems
to be a fact that this species uses the trotting gait so common in tins subfamily much
less than in the majority of felines. The gallop is a most impressive sight, though
difficult to describe succinctly. The action, together with the general movements of
the cheetah's body, have been analysed in some detail and diagrammatically depicted
by Hildebrand (1959, i960 and 1961). Omitting details, the stride may be reckoned as
starting with the feet close together under the belly, one hindfoot on the ground, the
body contracted in length. This hindleg gives a tremendous forward thrust, quickly
followed by a second from the other hindleg; the animal then flies through the air, the
body stretching out, the forelegs extending to their utmost until one forefoot strikes
the groimd, shortly followed by the second as the body continues to fly forward over
them. The hindlimbs are brought forward again, coming up once more with the fore-
limbs beneath the belly and overtaking them somewhat; and the sequence is repeated.
One such stride covers, on the average, the astonishing distance of about 7 metres.
There seems to be no record of the cheetah swimming.
Taxonomy. Arguments relating to the standing o{ julnnus as a species have been
dealt with above under the generic head; the question of subspccific division is con-
sidered here. It will be seen from the synonymy given earher that in the past a consider-
able number of proposals have been made to differentiate forms of Aciiwnyx from one
another, cliiefly with specific status. It is generally held today, and may be regarded as
virtually certain, that there is not, and never has been in recent times, more than a single
510 THE CARNIVORES OF WEST AFRICA
species 111 this genus; and therefore it these various proposed forms have any vahdity
whatsoever it can only be at racial level. Two names alone have been directly associated
with West Africa: !i(iu(;iihiisis Blainville and luckl Hilzheimer. The former is unavail-
able, having been preoccupied; but the latter is related to the same locality', Senegal,
and may reasonably be assumed to be synonymous (as in G. M. Allen, 1939). For the
rest of the suggested forms, it would seem improbable that those connected with India
or with extra-tropical southern Africa would be applicable to the region under con-
sideration in this work. This reduces consideration of proposed names to three possi-
bilities which, though extralimital, have their type localities situated 111 transcontinental
vegetation zones occurring in West Africa. These arc sociiti)iirr}n(;ii Fintzinger (Kordo-
tan), tiwgakilica Heuglin (? Bahr-cl-Abiad), and ir.Jijikri Hilzheimer (Kordofm). The
origins of these, together with hfcki Hilzheimer (Senegal), though ill-defmed or doubt-
ful, are very possibly m the Sahel woodland zone.
The standing and comparison of these forms was discussed fully by Hilzheimer
(1913). A great deal depends on the diagnosis of the earliest of them, .^octniiurriiK^ii; but
as Hilzheimer pouited out it was doubtful whether Fitzmger's very inadequate descrip-
tion of this form, differentia ting it from i,'i/fMfi(i Wagner — longer legs, darker colour,
bushier tail and scantier mane — was sufficient to make the name valid. It has, however,
been customary to retain it and to synonymise it with tiici^ahalica Heuglin, almost as
obscurely described, and sometimes (G. M. Allen, 1939) also with u\niiicri Fiilzheimcr.
In illustration of the slender basis of much of the taxonomy it is perhaps instructive to
draw attention here to that of the last of these, the diagnosis of which was not furnished
by Hilzheimer but by Wagner himself, intended purely as a supplement to Hermann's
earher inadequate description of giinaius and as a verbal complement to the painting
which Hermann had apparently intended to depict that species. Hilzheimer considered
Wagner's description to fail m both these respects having been made by him from a
stuffed skin of an animal collected by Riippell in Kordofan, differing 111 Hilzheimer's
opinion from i^uttdtus Hermann. In recent years it has been customar)' to assume that
West African animals are of the race sofiiiimrriii^ii — e.g. Rosevear, 1953, which was
based on oral hearsay from taxonomists .it that time reputed to know; and Dekeyser,
1955. which was very probably derived from that work.
The distinctions between various reputed forms o( Aciiioiiyx often hinge upon slight
differences in such characters as the ground-colour of the pelage, in the size, number
and colour of the spots, includmg those of the fice, and in the number of rings on the
tail. Without doubt differences, and sometimes marked differences, exist between
cheetahs; but the study material available is mostly insutiicient to say in what degree
such variations are due to idiosyncrasy, age, moult or other non-taxonomic factor.
Certainly the paucity of museum material makes it impossible to judge what range of
local variation may be expected in West Africa itself; but comparison of skins from
single areas in other parts of the continent make it clear that ground-colour, spot size
and shape can vary in one locality to a degree equal to that which has been thought
sufficient for the erection of new forms. The sole West African skin in London, now
almost 70 years old, differs clearly from the general run of extralimital specimens both
in its pallid ground-colour and its numerous dullish spots of small size: but it is. from its
ACINONYX
5"
mane, without doubt a young annual not yet ni the full possession of its adult coat, and
it is impossible to say into what it would eventually develop.
Wliethcr hi'cki is a valid race requires much more material than Hilzheimer's single
specimen to prove. It is in brief according to this author (1913: 290) a small, light-
coloured animal with a small number of spots, and soles with pale-coloured hairs.
However, whether size can be properly estimated from a single living animal of
unknown age; or racial colour in the wild, cither of pelage or soles, be satisfactorily
judged from one which has been kept for any length of time in a European zoo is open
to some doubt. Further, even ii soannwrriiiqii can be identified from its obscure descrip-
tion it yet remains to be demonstrated that West African specimens conform to it. The
fact is that the extent of any real knowledge of the cheetah in West Africa takes us no
further at present than the specific name. In any case it seems probable that with such a
wandering, wide-ranging species anything in the nature of a true local race is unlikely.
Measurements. The table which follows shows the only measurements available
for West Africa. Both specimens are females. The skull, though not of a particularly
young animal, is somewhat on the small side and the measurements of a fairly old
Table 31: Numerical data (or Aciiioiiyx jitbalus
Nigeria :
Yantumaki
Lake Chad
Uganda
Vegetation
Sudan
Sahel
?Sudan
Number in mean
I
I
I
Condylobasal lengtli
150-5
—
168-8
Basilar length
136-0
—
153-4
Palatilar length
58-2
—
68-7
Zygomatic breadth
116-9
—
131-0
Upper cheekteeth breadth
66-2
—
70-3
Nasals, length
50-6
—
67-5
Interorbital breadth
40-0
—
44-5
Postorbital constriction
52-0
—
57-1
Braincase breadth
69-3
—
69-9
Toothrow (f — (»')
49-1
—
53-4
p* length
(.9-2)
—
21-8
(h1 breadth
5-8
—
6-8
nil length
6-0
—
6-8
Head & body
770
Tail
—
553
—
Hindfoot
—
237
—
Ear
—
72
—
RATIOS (per cent)
Tail/head & body
—
7a
—
Zygom. br./condylob. 1.
78
—
78
Braincase/condylob. 1.
A6
—
41
Braincase/zygom. br.
59
—
53
Palatilar l./condylob. 1.
39
—
41
hiterorb./postorb.
77
—
78
p*lc—m^
39
—
41
512 THE CARNIVORFS Or WEST AFRICA
female from Uganda are given for comparison. No field body measurements exist on
any African skin for comparison with tlie young Lake Chad specimen.
GLOSSARY OF TERMS
alveolus
anterior
antitragus
auditory bulla
auditory meatus
basal length
basilar length
bifid
biotope
braincasc
braincase breadth
bristle-hair
buccal
bulla
bursa, aural
canine
camassial
caudal
cheekteeth
cheekteeth, greatest
breadth
cingulum
circumanal
cline
condyle -
condylobasal length
cusp
deciduous
Tooth socket.
To the front; foremost; furthest from the tail.
A lobe, sometimes emarginate, near the base of the outer margin of
the ear pinna.
See bulla.
The external orifice of the ear; the earhole.
The distance from the most anterior margin of the foramen magnum
to the anterior limit of the premaxilla (i.e. exterior to the incisors).
The distance from the most anterior part of the foramen magnum to
the anterior margin of the palate (i.e. interior to the incisors).
Divided into two parts by a notch.
Habitat.
Strictly, that part of the skull actually housing the brain; the cranium;
in contradistinction to the rostrum it is that portion of the skull
lying posterior to the anterior line of the orbits.
The greatest transverse measurement across the braincase taken
usually just above the squamosal processes.
Usually the main constituent of the outer fur; see page 12.
On the cheek side of the teeth.
One of the paired subglobular bones seen on the underside of the
skull housing middle and inner ear structures.
A small pocket situated on the external rim of the ear pinna.
The tooth immediately posterior to the premaxilla ; in the Carnivores
almost always the tallest in the jaw, pointed and recurved; the
"dog-tooth".
One of the blade-like sectorial teeth in the Carnivores; see page 17.
Pertaining to the tail.
The premolars and molars together; see page 16.
Measured across the outside of the teeth of the upper jaw, usually,
but not necessarily, the posterior angle of the camassials, p*-p*-
A prominent girdle around the base of the crown of a tooth just
above the alveolus.
Around the anal orifice; see page 11.
A gradual and sequential change of character without significant
break such as would justify division into separate species.
A rounded process on a bone serving as an articulation with another
bone.
The distance from the most posterior face of the occipital condyles
to the most anterior face of the premaxillae.
Braincase; laxly, the whole upper portion of the skull without the
lower jaw.
A point or elevation, often subconical, on the crown of a tooth.
Used of teeth; falling out naturally at some period of life, not in
response to old age or disease.
513
5U
THE CARNIVOUhS 1)1^ WESr AFIUCA
Jt-nt.il forii
diastema
digit
distal
dorsal
cmarginatc
cpihyal
foramen
(pliir. foramina)
foramen magnum
frontal
genal
tjlenoid fossa
glossal
habitat
hallux
hyoid chain, hyoidean
apparatus
infra-orbital
inner
interdigital
interorbital breadth
interramal
jugal
jugal process
lambdoidal crest
lateral
longitudinal
A method ot expressing in succinct form the number of each category
of tooth characteristic of a given genus or species. It is in pseudo-
fractional form, citing the teeth in correct order from incisors to
molars for one side only of the mouth, the upper and lower jaws
respectively above and below a horizontal line, the total shown
being, however, that for the entire mouth and thus twice the sum
of the given figures. The number of incisors (3) and canines (i)
being constant throughout the Carnivores the formula is often
reduced to that of the premolars and molars alone; see page 16.
A gap between two teeth.
Finger or toe.
Further from the medial axis or point of attachment or origin.
On or pertaining to the back.
Having a notch or indentation in the mat gin.
The upper part of the hyoid arch.
A (usually) small aperture in a bone or between bones, circular or
elliptical in shape, tor the passage of a nerve, muscle or blood
vessel.
The large opening at the posterior end of the skull through which
the spnial cord passes.
One of the paired bones lying between the nasals and the parietals
and forming with them the roof of the skull.
Pertaining to the cheek.
The long subcylindrical cavity on the underside of the skull which
receives the mandibular condyle to form the hinge coiuiecting
the lower to the upper jaw.
Pertaining to the tongue; situated on the tongue side ot the teeth.
Ihe kind of place, vegetationally speaking, which an animal
normally inhabits.
The 1st digit of the hind limb.
A U-shaped series ot bones between the root ot the tongue and the
l.irynx; see page 377.
One of the category of teeth set most anteriorly in the skull and
nearest to its medial axis, in the upper jaw arising from the
premaxillae.
Situated below the orbit or eye-socket.
Nearer the medial ,ixis.
Situated between the toes.
The least distance between the upper rims of the orbits measured
across the top of the skull.
Situated between the two branches ot the lower jaw.
The middle bone of the zygomatic arch joining that arising from the
maxilla to that from the squamosal.
A pointed projection of bone on the upper side of the jugal, forming
the lower part of the orbital ring.
An alternative name for the supra-occipital crest.
Situated to the side of the main axis.
Lengthwise; running in a head to tail direction.
GLOSSARY
515
mandible
mastoid process
maxilla
meatus
mesopterygoid fossa
medial
milk dentition
molar
morphology
muzzle
mystacial
nares
nasal length
nomenclature
nuchal
occipital condyle
occipital crest
occlusal surface
oestrus
orbit
outer
palatal length
palatilar length
paratype
paroccipital process
pectoral
pelage
The lower jaw.
A process of bone immediately posterior to the ear.
That bone of the upper jaw which bears the canines .and cheekteeth
and forms the m.ajor part of the pal.atc.
A tubular p.issage or, more restrictcdly, its opening.
On the ventral face of the cranium posterior to the palate, the
channel, usually broad and deep, between the fuic pterygoid
bones, i.e. the posterior part of the nasal passage.
Situated in the middle or along the middle axis.
The teeth, usually simple in form and erupting soon after birth, which
precede and are rcpkiced by the perm.incnt teeth char.ictcristic ot
the mature animal.
One of the most posterior category of teeth, not preceded by any
corresponding milk-teeth.
The branch of zoology that deals with the form and structure of
animals.
That part of the face that lies anterior to the eyes.
Resembling a moustache.
The nasal passages; or their openings, as in anterior nares, posterior
narcs.
The distance from the most posterior point of junction of the two
nasal bones to the most anterior tip of one or the other of them
(not the medial line, as in some works).
The scientific naming of animals.
Pertaining to the back of the neck.
One of the pair of smooth processes of the exoccipital bone at the
posterior end of the skull lateral to the foramen magnum serving
to hinge the head to the neck.
A process, often flange-like, of the supra-occipital bone which forms
the upper posterior portion of the cranium.
The surficc of a tooth which closes against the corresponding tooth
in the opposing jaw.
The state of being on heat, or in season, of a female.
The eye socket; or, more especially, the bony ring or partial ring
surrounding it.
More distant from the central axis.
The distance from the most anterior point of the posterior margin
of the bony palate to the foremost edge of the premaxilla.
The distance from the most anterior point of the posterior margin
of the bony palate to the posterior alveolar rim of the incisors.
A specimen forming part of the original material used and mentioned
by an author as the basis of description of a new species or sub-
species and from which he has selected and designated a holotype.
An outgrowth of the occipital bone extending in a downward
direction, sometimes finger-like but sometimes spreading over the
posterior surface of the auditory bulla.
Pertaining to the chest.
A general term for the fur of an animal.
5i6
THE CARNIVORES OF WEST AFRICA
pcrincil
petiole
phylogcny
pinn.i
postcaiiinc gap
postdcnt.il p.ilatc
posterior
postorbital constriction
postorbital process
pregeuital
premaxilla
premolar
prescrotal
process
proximal
pterygoids
ramus
rostrum
sagittal crest
sectorial
septum
sinuous
sub-
sub-bristlc-hair
submental
subocular
superciliary
supra-anal
supra-occipital crest
suspensorium
taxonomy
Pertaining to the region between the anus and the scrotum or vulva.
Herein used to denote the slender stalk of a sub-bristlc-hair; see
page 1 6.
The evolutionary history of an animal.
The external ear flap or conch.
A space unobstructed by teeth posterior to the canines, evident
when the jaws are closed; see page 18.
A medial extension of the main palate backwards and lyiixg posterior
to the molars.
To the back of; hindmost; furthest from the head.
The shortest distance across the top of the skull posterior to the
postorbital processes.
An outgrowth from the frontal bone, in the carnivores usually fairly
long, forming the upper rim of the orbit.
Anterior to the genital organs.
One of the pair of bones forming the extreme anterior part of the
palate and rostrum below and at the sides of the nares, and from
which the upper incisors arise.
One ot the teeth lying between the canine and the molars and in the
adidt state always preceded by a milk tooth.
Situated in front of the scrotum.
A natural outgrowth from a bone or other structure.
Closer to the medial axis or point of attachment or origin.
The paired, vertical, thin, wing-like bones on the ventral face of the
cranium to the rear of the palate, forming the lateral walls of the
posterior part of the nasal passage.
One of the two branches of the mandible.
The, usually tapering, portion of the skull lying anterior to the orbits.
A longitudinal ridge of bone situated on the medial axis of the
cranium, sometimes lacking, sometimes well developed into an
erect plate.
Adapted to cutting.
A partition separating two cavities.
Having a number of curves : wavy.
In combination with shapes or terms (as subequal, subcylindrical),
approximately, roughly, almost.
Shortened to sub-bristle, one of the components ot the top fur of
mamnials; sec page 12.
Beneath the chin.
Beneath the eye.
Situated over the eye.
Situated above the anus.
A more correct term for the occipital crest.
One of the bones of the hyoid chain.
The systematic arrangement of the animal (or plant) world in a
natural order of evolutionary relationship. It is necessarily closely
associated with nomenclature, which is thus, for convenience but
GLOSSARY
517
terete
toothrow
total length
tragus
transverse
trifid
trilobed
tympanic bulla
type
type locality
ventral
vibrissa
zygoma
(plur. zygomata)
zygomatic arch
zygomatic breadth
somewhat laxly, included under the side-headings in this present
work.
Having a cylindrical or slightly tapering form.
The complete series of teeth on one side of the jaw from incisors to
molars; sometimes for the purposes of convenient measurement
limited to the distance from the anterior rim of the canine alveolus
to the posterior face of the last molar.
The greatest length of a skull from its most posterior point to the
front edge of the most forward bone or tooth.
A cartilaginous process sometimes found near the base of the inner
margin of the car pinna.
In a direction across the body from side to side.
Divided into three parts by two notches.
Having three lobes; trifid.
See bulla.
The specimen used in the original description as the basis of naming
a new species or subspecies. More properly referred to as the
holotype.
The exact place from which an original type specimen came.
Pertaining to or on the abdominal side ; on the under as opposed to
the upper or top side of a structure or animal.
A stout, stiff and generally very long, tactile bristle growing singly
or in small clusters, mostly in a few constant and well-recognised
sites on the body; see page 11.
The arched bone supporting the cheek on each side of the skull,
comprising processes from the maxilla and the squamosal con-
nected by the jugal bone.
The zygoma.
The greatest width across the bones forming the zygomatic arches
measured at right-angles to the main longitudinal axis of the skull.
a
NOTE ON THE AREA TAKEN AS WEST AFRICA
This and cognate matters were dealt with at greater length ni Rosevear (1965), where
reasons for the choice of boundaries were given, liriefly, the West African region is
taken as bonnded on the west by the coast line but including the island of I\rnando I'oo;
and on the north by the iSth parallel ot latitude. The southern and eastern boundaries
are taken as the River Sanaga (Canieroiui) from the sea to its source and thence the
watersheds dividing the Congo and Nile systems from those rivers that run either into
Lake Chad or into the Atlantic Ocean north of the 4th parallel. The area thus enclosed
IS indicated on the accompainnng map, page 521.
The southern boundary, which seems at least in some measure to be a natural one,
cuts across existing political units, the names of which can, thus, not be usefully
employed; the term "upper Cameroun" is used, therefore, to indicate the part of that
country which lies mside West Africa as defined above. "Extralimital" means outside
the region dealt with in this work.
The name Congo, in reference to extralimital ciistnbutions and other matters relevant
to this work, is used rather laxly to indicate the basin ot the river, especially to its north,
rather than the river itself or any political unit.
518
NOTE ON VEGETATION
The West African vegetation was dealt with fairly fully, together with photographs
of the different types and maps of the zones, in Roscvcar (1953); and in a shortened
version without illustrations but with a map in Rosevear (1965). The vegetation of the
continent of Africa south of the Tropic of Cancer, using a somewhat different termin-
ology but accompanied by a large coloured map, has been succinctly treated, in both
French and English, in Keay ct al. (1959).
The system of classification used in this present volume has been accepted and
commonly employed in biological works of varying kinds for half a century or more.
It is admittedly of a broad nature, taking little or no note of sub-associations of vegeta-
tion within the main categories; but it has nevertheless served, and continues to serve,
a useful purpose and will seemingly do so until far more detailed ecological studies of
both plant and animal life have been achieved.
The terminology used 111 this present work, as in the others of this series of mono-
graphs on West African mammals, differs from that commonly employed only in the
substitution of "woodland" for "savannah". The reason for this change is that it seems
to the author that it is important to bring constantly before the naturalist that trees do,
indeed, play an extremely important role in these zones, providing for many mammals
sources of food, shade, shelter and refuge in alarm. It seems likely, at least to the present
writer, that a good proportion of the open-country mammals, whether giraffes,
polecats, squirrels or tree mice, are more actively aware of the arboreal than of the
graminaceous constituents of these zones. The French term "savanne boisee" contams
an essential conception of this class of vegetation that should always be present in the
mind of the field zoologist. Moreover, in dealing with West African animals it is
important to emphasize the vast difference of appearance and ecology between these
zones and the extensive areas of open plains which cover much of East Africa, where
the tree species, though possibly basically the same, are so incomparably more scattered
as to offer little hindrance to vision over long distances.
It is scarcely possible to define the various zones succinctly and at the same time
clearly; and it is even less feasible to indicate their limits. However, the latter can be
gathered in a broad way from the accompanying small-scale map, page 521. In using
this two things must be borne in mind: the boiuidary lines are throughout much of
their length nothing more precise than the mere joining of distantly separated though
fairly accurately known pouits of change ; and, secondly, the division between zones
is, in fact, rarely clear-cut, there being, often, a belt in which the definitive species are
to a greater or less extent mixed. The differing belts of vegetation are mostly the out-
come of various climatic factors of which the chief are rainfall, atmospheric humidity
especially at the peak of the dry season, and the number of months that this season itself
lasts. Li succession from the coast inland to the Sahara, that is m conditions of ever-
increasing aridity, the chief zones are as follows :
519
5iO THE CARNIVORES OF WEST AFRICA
High forest. Typically consisting of trees in different strata from very tall to low, the
crowns of which form a complete canopy blocking out the sky (hence the alterna-
tive term "closed forest"). Grasses absent. Vast areas of the zone have, however,
been destroyed for farming and other purposes, completely altering its character
and opening up the ground to light; grasses gain admission on a temporary basis
until they are shaded out by the gradual regrowth of tangled shrubs and eventually
the forest; but on areas such as broad roadsides that are constantly kept open
grasses fmd a permanent home. There are therefore a number of very distinct
habitats within the high forest zone. "Rain forest" is another commonly used
alternative term. It must be added that "forest" as often used in East Africa in the
context of animal habitats is, except in Uganda, a very different matter and would
generally be regarded in West Africa as little more tlian rather dense woodland.
Invasive woodland. This is immediately contiguous to the high forest and is, in fact,
on climatically potentially forest land; but constant severe clearing of the original
forest for farming has in the past admitted abundant grass through which the fires
from the adjacent Guinea woodland have swept annually, eventually destroying
all regeneration of forest-zone, fire-tender species. Only the fire-resistant trees of
the Guinea zone could in these circumstances obtain a foothold; and the Guinea
woodland, therefore, has almost completely invaded the area except for a few
relic patches of forest wliich tor religious or other reasons were unfarmed and have
been preserved in their original state, fire being unable to sweep through them
owing to their intrinsically damp nature and the characteristic absence of grass.
Tlie zone is thus to the eye indistinguishable from the contiguous Guinea woodland
except by the occasional presence of forest patches unassociated with water courses.
The name "derived savamiah" is commonly applied to this region, a meaningless
term since the present vegetation is derived from nothing but is of a type invading
an area to which it climatically has no claim and is thus more logically invasive
savannah or, in this work, woodland.
Guinea woodland. This is characterized by a dense, continuous ground cover of tall,
coarse grasses, 2 to 3 metres high, amongst which stand tliick-barked, fire-resistant
trees mostly of small to moderate size and of twisted shape, their crowns rarely
touching to form a canopy. A few tree species of larger size also exist and sometimes
stand in close clumps. In most years the grass is deliberately burnt in the dry season
(November to March) and springs again in lush green form from its tufts. This, in
conjunction with the freedom of movement then possible, attracts animals
normally absent dicing the rains. High forest penetrates, sometimes deeply, into
the zone along water courses, so that it is possible for species appropriate to that
vegetation to occur m what from the map appears to be Guinea woodland. Because
of Its more fertile nature much of this "fringing forest" has now been destroyed
for farming; but elsewhere in the zone the effects of cultivation are neither so
marked nor so permanent as in the forest.
Doka woodland. This is of very similar appearance to the Guinea woodland except
for Its slightly less abundant grasses and its donnnance by the tree knowni in Hausa
as doliii (Isohcrliniiij.
MAP
521
522 THE CARNIVORES OF WEST AFRICA
Sudan woodland. Althougli the overall appearance of this important zone has a
unifying "atmosphere" which distinguishes it pretty clearly from the Doka on the
one side and the Sahel on the other the vegetation is. in fict, very variable and is
made up of a mosaic of communities each determined by geology or soil moisture.
Succinct description is, therefore, even more difficult than in the case of the other
zones. Broadly the Sudan may be distinguished from the Doka by the virtual
absence of that tree and by the markedly greater number of thorny species. These
are mostly different kinds oi Acdda; but the zone nevertheless differs from the
Sahel by the relative uncommonness m it o( Aaicia raddiana. The grass cover is of
lower height and sparser than m the preceding zones, so that vision is less obstruc-
ted, passage easier, and the annual tires less intense and damaging.
Sahel woodland. The tree cover here is much more limited both as regards variety and
numbers. The zone is, indeed, dominated by a single species of Aciuia, the flat-
topped .-1. raddiiiiici, which covers considerable areas, the individual trees, however,
standing at some distance from each other. There are stretches from which .-InifiiU
are lacking, fiirly thickly covered with the grotesquely shaped shrub Coimniphora.
Throughout two-thirds of the year, from about mid-October to mid-June, there
is little or no rainfrll.
Subdcsert. Here the rainfiU is too slight and the atmospheric hunndity too low to
support any tree growth. There are a few low scattered shrubs and, during the
very brief rains, a sparse growth of tufted grass and annual herbs, often growing
from bulbous roots which constitute a welcome source of food to many mammals.
There are sand-dunes, some stable, which provide subterranean shelter.
Desert. This does not exist within the limits taken for West Africa in this present work.
It is characterized by the complete absence of vegetation except in a relatively few
favoured locations such as oases or above undergroiuid water-courses.
Besides these main zones, which each cover very large areas and extend across the
continent from west to cast, there arc a number of minor types of vegetation which
owe their existence to local fictors of ramfill, soil, topography or altitude. The
commonest of these are:
Mangrove. This occtus in coastal and sub-coastal areas subject to flooding by brackish
water diurnally. To all intents and purposes these swamps are made up of two
arboreal species, the red mangrove [Rhizophora) and die white mangrove
{Avicennia). The former exliibits two clearly recognisable types of growth : the
tall primaiy growth on very soft, newly deposited silt — a biotope offering no
refuge to any but purely arboreal or purely aquatic mammals; and the low, tangled
secondary growth on firm, peaty, root-riddled soil, flooded only at the highest
tides, and of annually lessening submergence. These latter areas are thus increasingly
open to and invaded by terrcstial mammals.
Fresh-water swamp. This is high forest, within the main zone and often just inland
of the tidal mangrove swamps; but it comprises species that can withstand annual
seasonal flooding by fresh water from the overflow of rivers during the rains. There
are also more restricted areas of permanently inundated fresh-water swamp fed by
small inland streams and occupied almost solely by the Raphia palm.
VEGETATION 5*3
Coastal scrub. Although contiguous to the seaboard and thus apparently well within
the region of high precipitation and humidity which fosters the tropical rain forest,
the vegetation of these relatively small areas is almost Subdesert-like in its sparse-
ness, consisting largely of low, tufted grass and scattered shrubs, the latter often
seemingly connected with old termitaria. The underlying causes are various — past
destruction, lowered rainfall due to the direction of the coastline, and soil factors.
It IS not to be confused with strand vegetation, which consists of psammophilous
grasses and creeping herbs growing along sandy shores.
Mountain forest. This, which lies between about looo and 2000 metres and is pretty
constantly enveloped in cloud, partakes of all the characters of normal lowland
high forest except that with increasing altitude the trees become progressively
smaller and are, throughout, draped with mosses and lichens.
Montane grassland. True Montane grassland lies at 2000 metres or higher and consists
of more or less continuous short grasses and herbs, with a complete absence of
woody growths. The mistake is not infrequently made of confusing highly
degraded "savannah" lying at moderate altitudes with this.
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532 THE CARNIVORES OF WEST AFRICA
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INDEX
NOTES:
1. English or other vernacular names are printed in capitals.
2. No distinction is drawn between currently accepted names and synonyms; generic names are
spelt with a Capital.
3. Subspecies are referred to the recognised genus not to the species.
4. Page references are in Arabic figures (9); major references are bold (9).
5. References to Figure numbers are in italics (p).
6. References to Colour Plates are in Roman style numerals (IX).
AARDVARKS, $
AARDWOIF, 341
Acacia, 71, 262, 293, 346, $22
Acinonychinae, 380
Acinonyx, 376, 380, 383, 412, 438, 44s, 467,
492, 63, 66
adustus, Canis, 49, 5, 6
aegypti, Ichneumon, 268
aethiops, Cercopithecus, 449
afra, Genetta, 191, 192, 196, 205, 28
africana, Felis, 494
Mephitis, 95
airensis, Felis, 396, 401, 2, jj
aithos, Hcrpestcs, 268, 276, 278, 279, VII
albicauda, Herpestes, 331
Ichneumia, 163, 301, jj, jg, VIII
albicaudus, Herpestes, 300, 301
albinism, 14
alexandri, Crossarchus, 281, 282, 288, 339
allamanda, Galictis, 321
almodovari, Herpestes, 331
Xenogale, 333. 335
Amblonyx, 148
Anahystcr, 148, 149
ansorgei, Crossarchus, 264, 281, 282
ANT, DRTVER, 326
anthus. Cams, 35, 38, 40
antiquorum. Hyaena, 345
antitragus, 7
Aonyx, 141, 143, 148, 149, 22, 2}, 24, II
arabicus, Fennecus, 56
arborea, Nandinia, 237
Arctogalidiini, 229
Arctoidea, 29
Ariela, 248, 282
Artiodactyla, 5
asiaticus, Leo, 437
Atherums, 448
Athylax, 291, 293
Atilax, 163, 268, 291, 292, 316, 331, 332, }8
atilax, Herpestes, 292
aubryana, Genetta, 198
aurata, Felis, 382, 424, 425, 435, 454, iS, jg, XI
aureus. Cams, 35, 36, 3, 4
Lynx, 425
aurita, Viverra, 56
Avicermia, 522
538
INDEX
539
badger(s), 29, 92, no
HONEY, 6, III, 16
barbara, Genetta, 192
Hyaena, 352
Bdeogale, 245, 321, 323
BEARS, 6, 9, 29, 34
bettoni, Genetta, 198
bini, Genetta, 200, 213, IV
binotata, Nandinia, 163, 230, 31, 32, V
Viverra, 230
BINTURONG, 229
BOBCAT, NORTH AMERICAN, 425
Bosraan, William, 175
brachyura, Felis, 411, 413, 421
Bradypus, 94, 95
breeding, 24
Bridelia, 174
bristle-hairs, 12
brockmani, Mellivora, 126
brucei, Fennecus, 56
Trypanosoma, 390
brunnea. Hyaena, 345
buchanani, Mellivora, 112, 124, 127, ig
bulla, 15, 29
burrowing, 23
bursa, 7, 342, 374
BUSHPIG, 450
caesia, Vulpes, 49, 67, 70, 8
caffer, Herpestes, 266, 274
calabaricus, Anahyster, 148, 149
Aonyx, 153
Calictis, 266
caligata, Felis, 396
Calogale, 266
cana, Galerella, 310, 319, 320, DC
Canidae, 29, 30
Caninae, 34
Canis, 35, 56, 449, 3, 4, 5, 6
Canoidea, 29
canus, Mungos, 316
capensis, Aonyx, 130, 133, 149, 22, II
Felis, 412
Hyaena, 354
Icton)Tc, 94
Lutra, 148, 149
Mellivora, III, 16, 17, 18, tg
Mephitis, 95
Viverra, in
caracal(s), 401, X
Caracal, 382, 383, 401, 412, }4, 55
caracal. Caracal, 402
Felis, 382, 401, 435, S4, 55, X
camassials, 17
Camivora, }
carotid canal, 29
cat(s), 373, 382
black-footed, 373, 384
possa, 373
GENERAL MARGUERITTe's, 396
GOLDEN, 424
AFRICAN, 425, XI
HOUSE, 383
JUNGLE, 384
PAMPAS, 437
RECENT, 373
SAND, 395
AIR, 401
SERVAl, 411, 412
SILVER, 426
SWAMP, 373
TODDY, 229, 230
TRUE, 383
WILD, AFRICAN, 88, 384, X
EUROPEAN, 387, 390
HAUSA, 392
MID-BEIT, 393
CATFISH, 133
catus, Felis, 382, 383, 390
caurinus, Mungos, 252, 262
celidogaster, Felis, 382, 425, 427, 428, 430, 432
centralis, Canis, 49
Cercopithecus, 449
cercropioides, Musanga, 236
cerda, Canis, 56
Megalotis, 50
cerdo, Canis, 56
chalybeata, Felis, 425
chaus, Felis, 373, 384
cheekteeth, 16
cheetah(s), 6, 375, 492, 493
Chiroptera, 5
Chrysailurus, 382
chrysothrix, Felis, 425
chui, Panthera, 457
540
INDEX
Cistanche, 6i
ciVFr(s), 6, i6i, 164
AFRICAN, 166, 168, 2i
AQUATIC, 183
ORIENTAL, 16S
PALM, AFRICAN, 6, 229, 23O
ASLAN, 229
TWIN-SPOTTED, 23O
TWO-SPOTTED, 23O, IV
TREE, 231
WATER, 167
"civet" (perfume), ii
civetta, Civettictis, 163, 168, 25, 26, 27
Viverra, 166, i68
Civettictis, 163, 166, 2$, 26, 27
claws, 9
dew, 9
retractile, 9
climbing, 23
COATIS, 6, 29
Cogiiiauxia, 236
colocola, Felis, 437
colour, sensitivity to, 9
Commiphora, 522
communis, Vulpes, 64
concisa, Mellivora, in, 123, 126, ip
congica, Aonyx, 154, 155, 22, 23, 24
Civetta, 177
CONIES, 3
corsac, Canis, 64
cottoni, Mellivora, iii, 124, 126, tg
COYOTE, 36
crassicauda, Bdeogale, 321, 322
Cricetomys, 82, 418, 448
cristata, Felis, 384
Genctta, 179, 18S, 198, 28, III
croacuta. Hyaena, 354
Crocotta, 353, 354
Crocuta, 343, 345, 353, ^9, ^o
crocuta, Canis, 353
Crocuta, 353, 46, ^p, 50
Crossarchus, 248, 249, 252, 262, 279, 337, 339,
J7, VI
Cuon, 75
cuvieri. Hyaena, 354
Cynailurus, 494
Cynalopex, 64, 70
Cynhyaena, 75
Cynictis, 316
Cynofelis, 492
Cynoidea, 29
DASSIES, 5
Dasyurops, 209
delalandi, Aonyx, 149
denhami, Vulpes, 56
dentition, 15
symbols, 17
Dieba, 35
diet, 20
digits, 9
Dingo, 36
doedcrleini, Canis, 38, 42
dog(s), 30
BUSH, 167
DOMESTIC, 36
HUNTING, 75, 76, g
TYPICAL, 34
WLLD, 6
"dog-teeth", 16
DOLPHINS, 5
dongolana, Genetta, 179, 188, 193
Viverra, 193
dorsalis, Canis, 38, 40
Vulpes, 65
Dorylus, 326
dubbah, Hyaena, 345, 352
dubia, Genetta, 203, 206, 210, 213
DUGONGS, 5
ear(s), 7, 14
ebermaicri, Lycaon, 76
economics, 25
edwardsi, Vulpes, 70, 72
elegans, 292
ELEPHANTS, $
encrita. Hyaena, 354
Enhydra, 128
Eptesicus, 185
Eremaclurus, 383, 401
erminea, Mustcla, 93
Erythrocebus, 449
Euxerus, 82
eyes, 7
accommodation of pupils, 8
sensitivity to colour, 9
INDEX
541
famelicus, Canis, 66
faniiliaris, Canis, 35
fasciata. Hyaena, 345
fasciatus, Herpestes, 251
Mungos, 248, 281
fearonii, Felis, 493
fearonis, Felis, 494
feet, 6, 9
Felidae, 160, 373
felina, Genetta, 179, 188, 190, 192, 201
Felinae, 373, 380
Felis, 387, 382, 383, 437, 438, 439, 445, 467,
493. 2, 5>-59, X-Xl
Feloidea, 29, 160
felting, 13
FENNEC(S), 56, 388, I
Fennecus, 56, 1, 7,1
fennecus. Cams, 56
ferae, 5
FERMT, 92
Ferungulata, 5
Ficus, 288
fieldiana, Genetta, 187, 205, 210
Fissipeda, 5, 6
flavus, Guepardus, 492
Fossa, 194
fox(es), 6, 30
bat-eamd, 3 i, 34
DESEHT, 61
FENNEC, 57
PALE, 70
PALLID, 70
RED, 64
AMERICAN, 64
COMMON, 64
ruppell's, 66, 1
SAND, 70, I
TRUE, 63
foxi, Felis, 384, 393, 31, S2
frenata, Ictonyx, 105
fulva, Vulpes, 64
fulvus, Guepardus, 492
Galeopardus, 383
galeopardus, Felis, 412
galera, Mustela, 292
Galerella, 163, 245, 267, 307, 40, 41, IX
Galeriscus, 245, 295, 300, 321, 42, 43, Vm
Galictis, 321
Galidia, 292
Galidictis, 292
gambianus, Aonyx, 153
Herpestes, 262
Leo, 460
Lutra, 149
Mungos, 262, 281-3, 285, }4, 35, VI
gap, post-canine, 18
genet(s), 6, 161, 164, 177
BENIN, 200, IV
crested, 198, iii
european, 191
hausa, 214, iii
Johnston's, 217
kuhn's, 217
north african, i92
PARDINE, 209
pel's, 210, rv
RUSTY-SPOTTED, 1 86
SENEGAL, 193, III
SMALL-SPOTTED, 212, IV
Genetta, 163, 167, 177, 222, 224, 28, 29, III, IV
genetta, Genetta, 179, 188, 190, 191, 195, 196,
197, 202, 28
Viverra, 191
genettoides, Genetta, 179, 203, 204, 206, 210,
213, 28, rv
Viverra, 210
gerrardi, Nandinia, 237
glands, anal, 10
circumanal, 11
mammary, 11, 357, 375
perineal, 11, 171
prescrotal, 11
scent, 10, 29, 165, 171, 179, 225, 233,
240, 253, 269, 285, 295, 298, 303,
312, 342, 356, 375
supra-anal, 11
Glires, 5
gothneh, Mungos, 251, 261
gracilis, Galerella, 309, 320
Herpestes, 266, 307, 316
Mungos, 248
grayii, Lutra, 141
GRISON, 321
Grison, 322
Guepar, 492
542
INDEX
Gueparda, 492
Guepardus, 492
guineeiisis, Atilax, 293, 299
guttata, Felis, 492, 493, 494
guttatus, Acinon)Tc, 492, 510
hair, surface patterns, 1 3
various types of, 1 1
hamiltonii, Paradoxurus, 230, 231
Haplocliromis, 146
hartcrti, Vulpes, 70, 72
haussa, Fclis, 384, 392, 394, 399
head, 7
hearing, senie of, 8, 22
hecki, Acinonyx, 494, 510
Hemigalidia, 292
hcrmaphroditus, Paradoxurus, 230
Herpcrtes, 266
Herpcstes, 163, 245, 24S, 266, 292, 300, 308,
314. iif', 3^9, 331, 454. 3(>, VII
Hcrpcstinae, 29, 239, ??
HONEY BADGER, 6, III
HONEY GUIDE, GREATER, 120
hunting methods, 21
hya£na(s), 6, 341
BROWN, 344
SPOTTED, 353, 46
STRIPED, 344, 345, 46
Hyaena, 344, 46, 47, 48
hyaena, Canis, 344, 345
Hyaena, 345, 46, 47, 48
Hyaenidae, 29, 160, 341
Hyaeninae, 341
Hyaenoides, 75
hybridization, 188
Hydrictis, 140, 141
Hydrogale, 140
hyoidean apparatus, 377, 378
Hyracoidea, 5
HYRAXES, 5
Hystrix, 448
IBIS, SACRED, 272
Ichneumia, 163, 268, 300, 331, jh J9. VIII
ICHNEUMON, 268
FOREST, 276, VII
NIGERIAN, 278
WESTERN, 275, VII
Ichneumon, 266, 268
ichneumon, Hcrpestcs, 163, 264, 268, 308, 330,
454. }(<, VII
Vivcrra, 266, 268
Ictidonyx, 94
Ictomys, 94
Ictonyx, 29, 94, 104, 12, I J, 14
inauritus, Ursitaxus, in
Indicator, 120
indicator. Indicator, 120
INSECTIVORES, 5
insularis, Genctta, 206, 210
intensa, Genetta, 198, 200
Nandinia, 237
munguis, Lutra, 149
iris, 8
Isoberhma, 520
JACKAl(s), 6, 29
BIACK-BACKED, 36
GOLDEN, 36, J
SIDE-STRIPED, 36, 49, 3
jacksoni, Galcriscus, 321, 322, 329
JAGUAR, 442
johnstoni, Genetta, 188, 203, 217
jubata, Fchs, 492, 493
jubatus, Acinonyx, 492, 493, 6$, 66
jujuba, Zizyphus, 47
JUJUBE TREE, 1 84
kamptzi, Fchs, 460, 465, 490
keys, Aonyx, subgenera, 149
Canidae, subfamihes, 34
Caninae, genera, 35
Canis, species, 36
Feloidca families, 160
Fissipeda, families, 28
Fissipcda, superfamihes, 27
Genetta, species, 189
Herpcstinae, genera, 245
Hyaeninae, genera, 344
Lutrin.ie, genera, 140
Mustclidae, subfamihes, 93
Mustcliiiae, genera, 93
Viverridae, subfamihes, 163
Viverrinae, genera, 166
Vulpes, species, 65
543
KINKAJOU, 6
kuhni, Liberiictis, 337, 43
KUSIMANSE, 282, VI
lalandii, Aonyx, 149, 154
lanea, Felis, 494
Lasiopus, 300
legs, 9
lehnianni, Genetta, 217
leightoni, Poiana, 226, 227
lenoiri, Aonyx, 154
Lutra, 149
Leo, 437, 459
leo, Felis, 413, 437
Panthera, 459, 63, 64
Leonina, 437
ieopahd(s), 6, 437, 439
CLOUDED, 442
HUNTING, 493
SNOW, 442
W. AFRICAN FOREST, 458
W. AFRICAN OPEN-COUNTRY, 458
Leopardus, 424
leopardus, Felis, 439
Panthera, 453, 458
Leptailurus, 382, 383, 402, 411
leptura, Viverra, 193
Lepus, 82
leuconota, Mellivora, iii, 124, 126, ig
leucurus, Herpestes, 301
liberiensis, Poiana, 227
Liberiirtis, 336, 45
libyca, Felis, 384, 397, 413, }!, 52
Mustcla, 104, 105
Poeciliais, 104, 105, iz, 15
LINSANG(s), 164, 220, 229
AFRICAN, 220
ASIATIC, 220, 224
leighton's, 227
Richardson's, 225, V
Linsang, 220
lion(s), 6, 437, 459
ASIATIC, 489
Lipetus, III
lipostiaa, Felis, 422
locomotion, 162
locmpo, Herpestes, 301, 306
Lophuromys, 185
lowei, Felis, 384, 392
LUNGnSH, 133
lupaster, Canis, 39
Lupus, 35
lupus, Canis, 35
Lutra, 140, 149, 150, 20, 21, 22, II
lutra, Lutra, 141
Mustela, 140
Lutreola, 93
Lutrinae, 128, 22
lybica, Felis, 384, 396
lybiensis, Felis, 384
Lycaon, 75, 368, p, 10, 11
lynesi, Felis, 384, 393
LYNX, DESERT, 402
Lynx, 425
lynx, Felis, 384
inacrodon, Atilax, 292
maculata, Genetta, 198, 202, 204, 205, 209
Hyaena, 354
Viverra, 209
maculatus, Dasyurops, 209
Serval, 382
maculicoUis, Lutra, 130, 133, 140, 141, 150,
20, 21, 22, II
MAGARIYA, 47
major. Ichneumon, 268
MAMMALS, EUTHERLAN, 5
FLESH-EATING, 5
PLACENTAL, 5
MANATEES, 5
mandjarum, Crossarchus, 252
Mungos, 262
Mandrillus, 449
manguensis, Lycaon, 76
Mangusta, 266, 292
margarita, Felis, 395, 2, 53
margaritae, Felis, 396
marginata, Felis, 396
margueritei, Felis, 396
marguerittei, Felis, 396
MARTEN, 92
massaica, Fehs, 490
matschiei, Lutra, 141, 142, 147
mauritiana, Ziziphus, 184
megabalica, Acinonyx, 510
Felis, 494
544
Megalotis, 56
meinertzhageni, Fclis, 401
melanism, 14, 171
melanotis, Caracal, 382, 402
melanura, Galcrella, 310, 315, 318, 40, IX
melanurus, C^iiictis, 316
Mclinae, no
Melitoryx, in
Mellivora, 29, III, 16, 17, iS, \i)
Mellivorinae, no
mellivorus, Ratelus, 1 1 1
Ursus, III
Mephitis, 95
mesos, Hcrpestcs, 268, 278, 279
microdon, Paraotiyx, 154, 155
Xcnogale, 267, 329, 331
minimus, Vulpes, 56
MINK, 92, 93
mischlichi, Lycaon, 76
Monachus, 6
monachus, Monachus, 6
mongoose(s), 6, 161, 239
BANDED, 248, 251
KORDOFAN, 26l
NORTH-WEST, 262
SCHWARz's, 262
taibot's, 261, VI
BLACK-POOTED, 323, VIII
DWARF, 315
EGYPTIAN, 268
FOUS-TOED, 321
GAMBIAN, 262, 283, VI
INDIAN, COMMON, 248
SMALl, 272
kuhn's, 337
liberian, 336
long-nosed, greater, 329
LESSER, 279, 283
MARSH, 272, 291, 292, 293, VII
RED-TARED, AIR, 3l8
WESTERN, 318, IX
SLENDER, 307, 316
FOREST, 318, IX
GUINEA, 319, IX
TYPICAL, 266
WATER, 291
WHITE-TALLED, 3OO, 30I, VIII
MONITOR, NILE, 272
moult, 13
multivittata, Poccilictis, 105, 109
Rhabdogale, 105
mungo, Hcrpcstes, 248
Mungos, 163, 251, 263, 281, VI
Vivcrra, 248, 251, 280
MUNGOS, AFRICAN, 248
Mungos, 163, 248, 281, 282, 283, 285, 300, 316,
322, 331, J-/, 35, VI
Mungotidac, 163
MUSANG, 229
Musanga, 236
Mustela, 93, 104, 105, 292
mustcla, Galerella, 311, 321
Mustclidae, 29, 92
mustclina, Rhabdogale, 95
Mustelinac, 93
Mutica, 5
Myonax, 245, 267, 307, 309
Nandinia, 162, 230, Jl, }2, V
Nandinimi, 229
naso, Atilax, 331
Herpcstes, 266, 267, 329, 331
Mungos, 331
Xenogale, 283, 294, 330, 331, 3}, 44
ncglecta. Fells, 382, 425, 427
nictitating membrane, 9
nigcrianus, Mungos, 331
Xcnogale, 330, 333, 335, 44
nigra, Fclis, 384
nigricauda, Ichneumia, 301, 306
nigripes, Bdeogale, 321, 322
Felis, 373
Galeriscus, 322, 323, 42, 4}, VIII
nilotica, Lutra, 141, 142, 147
niloticus, Var.mus, 272
noltei, Crocotta, 354
Croaita, 371
Hyaena, 354
nostrils, 7
nubianus, Thos, 38
nubicus, Fclis, 410
obscurus, Crossarchus, 262, 279, 282, 337, 57, VI
occidcntalis, Herpcstes, 268, 275, 277, 278, 279,
VII
INDEX
545
OCELOT, 424
ochracea, Galerella, 245, 308, 309
Poiana, 225
ochraceus, Herpestes, 266, 307
ocreata, Felis, 390, 396
oertzeni, Canis, 70
Vulpes, 72
ogilbyi, Felis, 413
oralis, Poecilictis, 108
orientalis, Hyaena, 345
omata, Felis, 413
Osbomictis, 167
Otocyon, 31, 34
otteh(s), 6, 29, 92, 128
african ciawiess, i49
african long-clawed, i4i
cape clawless, i49, ii
small-toothed clawless, i54. i55
speckle-throated, i4i, ii
spotted-necked, i4i
TYPICAL, 140
TYPICAL AFRICAN CLAWLESS, I49
pads, 9
interdigital, lo
metacarpal, lo
metatarsal, lo
pedal, 10
palmar, lo
plantar, lo
pallida, Vulpes, 66, 67, 70, I
pallidus, Canis, 70
PALM, TODDY, 230
paludinosus, Atilax, 163, 291, 292, 331, }8
Herpestes, 292
Mungos, 331
paludosus, Athylax, 293
PANDAS, 6
PANTHER, 440, 442
Panthera, 377, 380, 383, 437, 439, 493, 60-64
panthera, Felis, 439
pantherina, Genetta, 209
Pantherinae, 380
Papio, 449
Paradoxurinae, 229
Paradoxurini, 229
Paragenetta, 188, 217
Paraonyx, 139, 142, 148, 150, 151, 154, 22, 2j, 24
pardalis, Felis, 424
Pardictis, 166, 219, 224
pardina, Genetta, 179, 188, 191, 201, 203, 209,
213
Pardus, 437
pardus, Felis, 437, 439
Panthera, 439, 458, 60-62
paroccipital process, 29
parvidens, Herpestes, 275
patas, Erythrocebus, 449
pelage, 11
pattern, 13
texture, 13
Perissodacryla, 5
persica, Salvadora, 46, 61, 301
petiole, 12
pharaon. Ichneumon, 268
phelypaea, Cistanche, 61
philippsi, Paraonyx, 148, 154, 155
phoenicura, Galerella, 310, 318, IX
phoenicurus, Mungos, 316
Phoenix, 231
picta, Hyaena, 75, 76
pictus, Lycaon, 76, g, 10, it
Pinnipedia, 5, 6
Plasmodium, 102
play, 24
pluto, Atilax, 299
Herpestes, 293
pococki, Felis, 413
podolaena, Cogniauxia, 236
Poecilictis, 29, 98, 104, 12, jj
Poecilogale, 94
poecilotis. Caracal, 402
Felis, 403, 406, S4, J5
poensis, Aonyx, 147, 153
Genetta, 188, 202, 203, 204, 2ii, 212,
IV
Lutra, 149
Poiana, 167, 220, 30, IV
POLANE, 220
polecat(s), 29, 92, 93
AFRICAN, 94
STRIPED, 6, 110
AFRICAN, 95, 12
SENEGAL, 104
Potamochoerus, 450
Primates, 5
54<5
Prionodon, i66, 220, 224
Prionodomini, 164, 220
Proboscidea, 5
Procyonidae, 6, 29
Profelis, 382, 383, 424, 454
Proteles, 341
Protelinae, 341
Protopterus, 133
Psammophis, 255
Pscudogenetta, 188, 214, 215
pulverulentus, Galerella, 268
pupil, eye, 8
RACOONS, 6, 29
raddiana, Acacia, 71, 262, 346, J22
Raphia, 522
RAT, Pharaoh's, 269
RATEl(s), 29, III, 16
AIR, 127
BLACK, 126
LAKE CHAD, 126
SPECKLED, 126
WHITE-BACKED, 12(5
ratel, Viverra, 1 1 1
Ratellus, iii
Ratelus, iii
ratlamuchi, Galerella, 268
Reade, Winwood, 277, 428, 454
reichenowi, Panthera, 440, 453, 455
rex, Acinonyx, 492
Rhabdogale, 94, 105
rhinarium, 7
Rhinolophus, 185
Rhizophora, 522
richardsoni, Poiana, 225, 227, jo, IV
richardsonii, Genetta, 220, 225
riparius, Canis, 38, 41, 43
robustus, Atilax, 292
RODENTS, 5
rothschildi, Poccilictis, 105, 108, 1}
rubida, Felis, 395
rubiginosa, Genetta, 179, 188, 208
rueppclli, Vulpcs, 66, S, I
rufa. Fells, 425
Hyaena, 354
ruScauda, Galerella, 308
rutila. Fells, 425, 427
rutilus, Felis, 425
saarensis, Vulpes, 56
sabbar, Canis, 66
saharae, Galerella, 310, 318
Herpcstcs, 316
Salanoia, 292
SALT BUSH, 46, 61, 301
Salvadora, 46, 61, 301
sanguinea, Galerella, 163, 308, 309, 310, 316,
40, 41, IX
sanguineus, Herpestes, 266, 316
savanicola, Felis, 384, 395
scent-marking, 10
Schaeflia, 35
schwarzei, Civetta, 170
Scotophilus, 185
senegalensis, Acinonyx, 510
Felis, 413, 460, 465, 490, 493
Genetta, 179, 188, 191, 193, 203,
2g, III
Irtonyx, 102, 104, ij, 14
Mustela, 95
Thos, 38, 40, 41
Zorilla, 95
SERVAi, 374, 412, XI
Serval, 382
serval, Felis, 382, 412, 434, 435, }6, 57, XI
Servalina, 383
servalina, Felis, 411, 413, 422
Genetta, 179, 191, 196, 198, 200, 202,
212, 213, 214
SERV ALINE, 411, 416, 42I
sharicus, Lycaon, 76, to, 11
sibilans, Psammophis, 255
sight, sense of, 22
signata, Mellivora, in, 113, 124, 126, 19
sikapusi, Lophuromys, 185
silvestris, Felis, 3S7, 389, 390, 401
Phoenix, 231
Simocyoninae, 75
Sirenia, 5
size, range, 6
skull, 14
SKUNKS, 29, 92, no
SPOTTED, AMERICAN, 95
smell, sense of, 8, 22
SNAKE, SAND, 235
snake-killing, by mongooses, 242, 255, 271, 306
sociability, 22
INDEX
547
soemmeringii, Cynailurus, 494,
soemmerringii, Acinonyx, 510
sole, 9
somalicus, Mungos, 262
soudanicus, Canis, 38, 39
Speothos, 75
Spilogale, 95
stoat(s), 29, 92. 93
striata, Hyaena, 345
Zorilla, 95
striatus, Bradypus, 94, 95
Ictonyx, 95. '2> '3, '4
stuhlmanni, Genetta, 180, 186
suahelica, Genetta, 186
sub-bristle-hairs, 12
sudanicus, Ictonyx, 103
suspensorium, 377
sympatry, in Genetta, 188, 203, 215
taenianotus, Herpestes, 248
tail, 10
talboti, Crossarchus, 251
Mungos, 261, VI
tapetum lucidum, 8, 129, 342, 374
teeth, 15
camassial, 17. 2
cheek, i6
dog, 16
sectorial, 17
temmincki, Felis, 424
thierryi, Crocotta, 354
Crocuta, 371
Genetta, 179. 188, 203, 214. HI
Hyaena, 354
thinobia, Felis, 401
thinobius, Eremaelurus, 383
thooides, Canis, 38, 42
Thos, 35, 38
throwing, by mongooses, 297, 305
Thryonomys, 46, 82, 418, 448
tigrina, Genetta, 179. 188, 190, 201, 205, 207,
210
Tilapia, 146, 449
TODDY-CAT, 229, ^30
togoensis, Crocotta, 354
Crocuta, 371
Fehs, 413
Hyaena, 354
Topsell, 348
torquatus, Felis, 390
tortilis. Acacia, 71
tragus, 7
tricolor, Canis, 76
Lycaon, 75
Trypanosoma, 390
Tubulidentata, 5
typicus, Lycaon, 76
Ratellus, :ii
UMBRELLA TREE, 236
underfur, 12
Unguiculata, 5
UNGULATES, EVEN-TOED, 5
ODD-TOED, 5
urinatrix, Mangusta, 292
Urolynchus, 382
Ursidae, 6, 29, 34
Ursitaxus, III
Urva, 266
vaillanti, Poecilictis, 108
VANSiRB, 292
vansire, Atilax, 292, 293
Varanus, 272
variegatus, Canis, 38
venatica, Felis, 492, 493
Hyaena, 76
venaticus, Acinonyx, 492
venator, Acinonyx, 492, 493
vibrissae, 11
villiersi, Pseudogenetta, 214, 215
vision, 7
colour, 9
Viverra, 56, 166, 167, 292
Viverricula, 167
Viverridae, 160, 161, 341
Viverrinae, 164
Viverrini, 164, 220
voang-shire, Mustela, 292
vulgaris, Genetta, 192
Lutra, 153
Thos, 35
Vulpes, 36, 56, «3, S, I
vulpes, Canis, 64
Vulpicanis, 35
.U8
INDEX
wagneri, Acmon)'x, 494, 510
WEASEi(s), 29, 92, 93
STRIPED, 6, 104, no, ti
SOUTHERLY, I09
webs, 9
WHALES, 5
whiskers, 1 1
WOLF, 36
Xenogalc, 245,267,283,294, 314, 329, 331, }}, 44
zebra, Mungos, 281
zerda, Canis, 56
Fennccus, 56, 1, 7, I
zibetha, Viverra, 166
Zizyphus, 47, 174, 184
ZORILLA, 95
Zorilla, 94, 95
zorilla, Mustek, 95
Viverra, 95
ZORRLE, 95
ISBN 0 565 00723 8