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LIFE-HISTORIES

OF

AFRICAN GAME ANIMALS

VOLUME I

BOOKS BY THEODORE ROOSEVELT
PUBLISHED BY CHARLES SCRIBNER’S SONS

LIFE-HISTORIES OF AFRICAN GAME ANI-
MALS. With Edmund Heller. Illustrated. 2
vols. Largei8yo . . . . . © + « « $10.00 net

AFRICAN GAME TRAILS. An account of the African
Wanderings of an American Hunter-Naturalist.
Illustrated. Large svo . ep wee fe Se 00KneE

OUTDOOR PASTIMES OF AN AMERICAN HUNTER.
New Edition. Illustrated. 8vo . . . $3.00 net

HISTORY AS LITERATURE and Other Essays.
TSO sh) Sah'o Suucy ia) ai Mearns We a anor

OLIVER CROMWELL. Illustrated. 8vo . $2.00 net
THE ROUGH RIDERS. Illustrated. 8vo . $1.50 net

THE ROOSEVELT BOOK. Selections from the Writ-
ings of Theodore Roosevelt. 16mo . 50 cents net

THE ELKHORN EDITION. Complete Works of
Theodore Roosevelt. 26 volumes. Illustrated.
8vo. Sold by subscription.

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| LIFE-HISTORIES

OF

AFRICAN GAME ANIMALS

BY

THEODORE ROOSEVELT

AND

EDMUND HELLER

WITH ILLUSTRATIONS FROM PHOTOGRAPHS, AND FROM DRAWINGS
BY PHILIP R. GOODWIN; AND WITH FORTY FAUNAL MAPS

VOLUME I

a JUN LS be
y Zaee LO. 7

ek ati, ey A \yse
onal

NEW YORK
CHARLES SCRIBNER’S SONS
IQI4

CoPYRIGHT, 1914, BY
CHARLES SCRIBNER’S SONS

Published April, 1914

A)
CHARLES Do. WALCOTT

SECRETARY OF THE SMITHSONIAN INSTITUTION

Sp Ny

eo

PREFACE

Tue life-histories of African game mammals offer an
almost virgin field for investigation and study. The pres-
ent treatise is a faithful account of what we have ourselves
observed; it sets forth much of what is now known; but its
real value must lie in its being treated primarily as a sug-
gestion of what is still open for discovery in this vast field
of animal psychology and adjustment to environment. The
discovery of new species and races based upon the study of
preserved specimens of game animals, has already pro-
gressed very far; but the more attractive field which in-
cludes the habits of the game remains yet to a great extent
unexplored. This field is peculiarly open for investigation
to big-game hunters, and to all other men who go far afield
and obtain first-hand knowledge of the conditions under
which the game animals live. The closet naturalist, with
his technical knowledge of the structure of animals, can be
trusted to perform the work of classification to a mathe-
matical degree of precision; but we cannot obtain from him
a trustworthy account of the behavior of animals in their
natural environment, or learn from him the value to the
animals of the various structures or characteristics which
he has shown them to possess. Much knowledge regarding
the habits of game is acquired by the successful sportsman.
Yet it is often infinitesimal in quantity compared to what

vil

Vill PREFACE

may be acquired if the outdoors observer will direct his in-
vestigations along the broad lines covering the life-history of
the species with which he comes in contact. To carry out
such investigations successfully it would be necessary to
spend many hours and days, perhaps even weeks and
months, observing certain individuals or family groups of
game. This is quite beyond the limits of time alloted the
average sportsman. Nevertheless much can be learned by
the collected evidence from many fragmentary observa-
tions, providing only these are accurate. A great mass of
accurate fragmentary observations will often spell far more
progress in investigations of this kind than the observations
of a few trained individuals over an extended period of time.

The specimens of game animals most familiar to the
writers are those secured by the Smithsonian African expe-
dition under the direction of Colonel Roosevelt, which are
now preserved in the National Museum at Washington.
This collection consists of some six hundred specimens of
big-game mammals from British East Africa and the upper
Nile regions, comprising more than seventy species or races,
nearly all of which are represented by series of various ages
and sexes. Besides this collection, Edmund Heller has
examined at the National Museum the Paul J. Rainey col-
lection from British East Africa consisting of some four
hundred specimens of big game, the Abbott collection from
Kilimanjaro, the Carl Akeley collection from British East
Africa in the Field Museum, the Tjader collection in the
American Museum of Natural History, the British Museum
collections, the Powell-Cotton collection from northern
Uganda and Mount Elgon, the Berlin Museum collection,
the Congo Museum collection at Brussels, the Paris Museum

PREFACE 1x

of Natural History collection, and a considerable number of
smaller collections of game animals.

The country traversed by the writers jointly during their
association on the Smithsonian African expedition in 1909-
1910 covered the region of the Athi Plains north to Macha-
kos and westward to Nairobi, thence from Kijabe southwest
to the Loita Plains or Sotik district and northeast to Lake
Naivasha. From this point a journey was taken over the
summit of the Aberdare Range through Nyeri to the south-
west slopes of Mount Kenia. Returning to Nyeri the jour-
ney was continued northeast, skirting the slopes of Kenia to
its northeast base at Meru. A further expedition was made
from Meru northward to the Northern Guaso Nyiro River,
continuing up country along its course and across the Lai-
kipia Plains to Nyeri and thence to Naivasha. A journey
into the Uasin Gishu country was made from the summit of
the Mau Escarpment at Londiani northwest to Sirgoit Lake
and the headwaters of the “Nzoia River. After returning
from the Uasin Gishu Plateau the journey was continued
westward by way of a steamer across the Victoria Nyanza to
Entebbe and Kampala, and thence by way of Hoima to
Butiaba on the northeast shore of the Albert Nyanza. Em-
barking at this point in boats the route led down the Nile to
Rhino Camp near the present site of Wadelai, but upon the
opposite or west bank of the river. From Rhino Camp the
voyage was continued after an interval to Nimule and thence
overland to Gondokoro. A trip was made westward of
Rejaf. Here the voyage was farther continued down the
Nile to Lake No at the entrance of the Bahr el Ghazal and
thence to Khartoum, where the natural history operations
were discontinued. This journey covered a period of eleven

x PREFACE

months’ field work, almost every day of which was devoted
to active operations. More recently, in 1911 and 1912,
Edmund Heller again traversed much of the same highland
country in British East Africa with Paul J. Rainey. In
addition to the regions previously visited he journeyed into
the desert district lying between the Northern Guaso Nyiro
River and Mount Marsabit and also visited the coast dis-
tricts of Mazeras, Maji ya Chumvi, and the Taita Hills and
the inland districts in the Kakumega forest south of Mount
Elgon. Previously to these two journeys he accompanied
Carl Akeley to British East Africa in 1905, visiting the
highland region about the Athi Plains, Lake Naivasha, and
Molo on the summit of the Mau Escarpment, in addition to
the lowland districts near Voi.

At present we know so little regarding the habits and
life-history of the various game animals that we find our-
selves at a loss to account for many of their most obvious
peculiarities. For instance, the distribution of almost any
game animal offers some features which with our present
knowledge are inexplicable. It is well known that every
animal, or plant for that matter, has strongly within it the
tendency to enlarge its distributional area until it covers
the habitable earth. It is equally well known, although
not so universally kept in mind, that the distribution of each
species marks the limits within which it is able successfully
to compete with its environment. It would appear, there-
fore, a comparatively easy matter to determine the factors
which are accountable for the distribution of any species;
and yet no task in natural history is more difficult.

This is one of the fields open to sportsman and to
traveller alike. Dozens of factors enter into each case; and

PREFACE xi

no two cases are quite alike, each species being largely a
law unto itself. Generalizations concerning habits and dis-
tribution are accordingly very difficult to prove true until
much investigation has been undertaken concerning all the
species or groups concerned. ‘The distribution of one species
may depend upon the distribution of its food plants or ani-
mals, of another upon its natural enemies, of another upon
climatic conditions; while yet others may be limited in dis-
tribution only by natural boundaries such as large bodies
of water or high mountains. Usually the factors consist
of a combination of some of these or all of them acting in
varying degrees.

Again, take the question of the extent to which an ani-
mal is nocturnal. In our observations on African game we
were not able to make out whether or not under natural
conditions some of the antelopes were more nocturnal than
others. But in American game this is certainly the case.
The whitetail deer will often feed at all hours of the night,
as we can personally testify, and this when there is no moon.
But Mr. Shiras informs us that moose do not feed—at least
by the water’s edge—later than ten or eleven at night;
whereas the whitetail will continue feeding until dawn.
Caribou are diurnal. Wapiti feed freely in moonlight, but
move about very little on moonless nights. Blacktail move
about more freely on dark nights than wapiti, but hardly
as freely as whitetail. |

The chief purpose of the present publication is to give
our own observations, and in some cases to add what is
already known, regarding the life-histories of African game
animals, so that sportsmen who combine love of hunting
with a taste for faunal natural history, and outdoor ob-

xil PREFACE

servers of all kinds, may direct their efforts along new lines
of investigation and toward a more complete understanding
of the life-histories of the various species.

In dealing with the systematic side of the work the
authors have kept in mind the desirability of keeping the
recognition of races well within the powers of discrimina-
tion possessed by the average non-scientific outdoors man,
whether sportsman, explorer, or nature lover. Some de-
scriptive naturalists have recently proposed distinctive
names for new races of game mammals based upon very
slight characters in color or form, differences which can
scarcely be detected by an expert, much less by a layman.
In still another and even more numerous class of cases, the
supposedly -distinct species or races are based upon indi-
vidual differences in specimens, or upon immaturity. The
described races of elephant, giraffe, and buffalo are chiefly
of this character. The preserved specimens of these great
mammals are so few that it has been possible to describe
races only from the study of individual specimens taken
usually in widely remote localities and often of very different
ages and even of different sexes. The comparison of mate-
rial of such diverse character has in many cases led to the
description of individual differences as racial. At present
all the museums combined do not contain sufficient speci-
mens representing these great mammals to make possible
the exact determination of the subspecific characters of the
geographical races. The difficulty of studying the largest
species applies in a general way to the bulk of the big-game
animals. Besides the limited number of specimens avail-
able, there are physical difficulties which often lead the
species-maker astray. The mere size of the great specimens

PREFACE xlil

prevents the optical comparison of form or coloration when
the specimens are arranged side by side in a series, the eye
being incapable of taking within its area of vision at close
range more than one specimen, or part of one, at a time, so
that comparison by measurements must be resorted to.
The smaller mammals, on the other hand, can be compared
much more minutely because a large number of them can
be brought within the range of vision at the same time; and
also, because of their smaller size, larger numbers of them
are available in museums for study. The specimens of large
mammals being limited in number, and their size being so
great, minute discrimination cannot be employed in deter-
mining their characters. 3

A “species,” as we have used it throughout the text,
may be defined as the most limited collection of individuals
showing some one structural or color character or combina-
tion of characters distinct from the character or characters
possessed by any other group of individuals. A subspecies
or race we may define as a closely related group of indi-
viduals occupying a certain portion of the area occupied
by the species to which it refers, and exhibiting intermedi-
ate characters in part or in all of the individuals occupying
the intermediate regions between the centre of development
of the race and the centre of development of any other race
of the same species. In other words, the subspecies is a
geographical race linked with any other subspecies of the
species to which it refers by individuals exhibiting inter-
mediate characters. We regard degree of difference as the
essential feature in the determination of species and hold
that geographical isolation is not of itself of specific
value if the individuals concerned show but slight, even if

XIV PREFACE

constant racial characters. The giant eland inhabiting the
Lado Enclave and Bahr-el-Ghazal Province is a good ex-
ample of the case in point. The Nile race differs very
slightly from its geographical ally which is found, as far as
we know, only in the Senegal region on the West Coast of
Africa and is separated from the Nile form by some two
thousand miles. Although so widely isolated geographi-
cally from the Senegal eland, Taurotragus derbianus, the
degree of difference in coloration or other characters of the
Nile race is so slight that we prefer not to give it specific
rank, and refer to it merely as a race Taurotragus derbianus
gigas.
Genera we have treated in a similar way; that is, we
have made degree of difference the essential feature and
have grouped together only species equally related. Con-
venience has not been considered a determining factor, no
account being taken of the number of species within the
genus. A few cases of antelope genera demanding segrega-
tion in order to emphasize the relationships of their various
species have come under our notice. Hunter’s antelope of
the Tana River is a case of this sort. This species has been
taken out of the genus Damaliscus with which it neither
agrees in horn shape nor coloration and has been placed in a
new genus, Beatragus. The genus Kobus as generally used
is a further instance of a similar sort. Three groups of not
especially close relationship and all of wide distribution
over Africa have been associated together. This genus is
now split into: Kobus, the waterbucks of large size with
wide-spreading, slightly recurved horns and shaggy-haired
bodies; Adenota, the kobs of small size with s-shaped horns
and short-haired bodies; and, finally, Onotragus, the lech-

PREFACE XV

wis with lyrate, wide-spreading horns and hairless pasterns
on both fore and hind feet. This arrangement shows at
once the true relationship of such a completely isolated
member of the lechwis as the Nile lechwi, Onotragus
megaceras, which is usually referred to as Mrs. Gray’s water-
buck, an altogether meaningless and misleading name.

We are well aware that objection may be taken to our
use of the words “species” and “genus.” This is unavoid-
able, for as yet there is no general standard of agreement

’

as to their use. Of course “species” is an artificial term,
which, no matter how defined, can only approximately ex-
press the facts in nature. No definition will entirely and
precisely meet the case; and any limitation we use in defin-
ing species will still necessarily leave the term in one case
out of parallel with the term in another. If we had all the
connecting links before us it would be practically impossible
as regards any animal to do more than mark the “specific”’
differences in purely artificial and conventional fashion; for
even if species are due to sudden mutations and not to in-
finitely slow changes, these mutations must, in the immense
majority of cases, each be so comparatively small that the
use of the word “species” would appear an exaggerated
way of noting the difference. Moreover, as regards the
“species”’ of some animals, such as the arctogeal wolf—
including, say, the Texan, extreme boreal, and Indian forms
—and the common zebra or bonte-quagga, the gradations
are even at this stage of the world’s history practically com-
plete between extremes which, if it were not for this inter-
grading, would unquestionably be accepted as specifically
distinct. It is impossible to arrange either species or sub-
species satisfactorily in linear fashion, so as to show the rela-

XVI PREFACE

tions of each to the others, simply because nature does not
develop them in one line, but by an indefinitely varied sys-
tem of branching in all directions. Yet ina book it is neces-
sary to adopt some linear system—as we have adopted it in
dealing with the Bovide, for instance—even though we well
know that this system is faulty.

Moreover, in some forms the variation will be in one
set of characteristics, in some forms in a totally different set.
In certain African antelope—the eland for instance—color
is a more permanent characteristic than osteological detail.
On the other hand, the American black bear, without change
of color or external peculiarity, shows here and there such
changes in skull structure as to have given rise not only to
the erection of subspecies but even of species by certain
learned systematists. The lion and tiger differ more from
one another in color and external peculiarities than do any
other pair of the big cats; and yet their skeletons and skulls
are practically indistinguishable.

Again, the presence or lack of interfertility between
allied forms may have no seeming reference to the degree of
variation in bodily structure. The differences between
horses, asses, and zebras are so purely differences of color
and external anatomy that it is difficult to tell from the
skeletons of fossil remains the type to which they belonged.
On the other hand, the common ox, the humped ox, the
bison, the yak, and the gaur are sundered by such marked
skeletal as well as external peculiarities that many system-
atists put them all in different genera, and there is never
the slightest difficulty in telling the fossil forms apart.
Yet the mules of the ass and horse are infertile, whereas the
various species of Bovide mentioned above, seemingly far

PREFACE Xvii

more widely separated than the horse from the ass, are fer-
tile inter se.

It must be understood, therefore, that “species” is as
purely a term of convenience as “genus.” It designates all
the animals of a certain type which it is convenient to group
together under one name, both to indicate their more or
less close relationship to one another, and their greater or
smaller dissimilarity from other forms. As the term is one
of convenience and should indicate relationship, it is well
not to over-refine it in using it on ordinary occasions—that
is, not to become a species-splitter—and, as far as possible,
to use it so that there may be some rough approximation
between what is meant by species in one case and in an-
other. In technical scientific treatises the use merely of
generic and specific terms may not be sufficient to indicate
the finer differences; in such event it is unwise to make a
fetich of binomial Latin terminology—in itself also a pure
convenience—and it is well to use freely a trinomial, or
quadrinomial or pentanomial terminology. But in ordi-
nary writing, not only for scientific laymen, but for intelli-
gent laymen of all kinds, and also for scientists not so much
absorbed in details as to lose sight of the importance of the

’

whole, it is well to use the word “‘species” in as large, as
ample, a sense as possible. Any needed further definition
can be made by the use of such words as subspecies, vari-
eties, geographical races, and the like.

Mr. George Shiras, out of his own experience, recently
supplied a case in point. Mr. Shiras, although he would
disclaim the title, is a faunal naturalist whose photographs
of and notes on the big game and various other mammals

and various birds of the North American wildernesses are

XVill PREFAGE

of unique value. He is a singularly competent observer.
Recently he was speaking of his first trip to Alaska and how
he had been interested in reading about the “Alaskan jay”
—only to find it, as he expressed it, “the same old whiskey-
jack,” identical in every essential with the familiar whiskey-
jacks of the woods of Maine, Canada, Michigan, and the
Rocky Mountains. He said that he knew the blue jay, the
Florida jay, the big-crested jay of the Rockies, all obviously
distinct species; and he knew the whiskey-jack, or Canada
jay, also as a separate species; but that to write about a
local variety of the latter as the “Alaskan jay”’ as if it were
a separate species in the sense that is true of the Florida jay,
or blue jay, or Steller’s long-crested jay, merely served
to confuse him—that is, to confuse an intelligent, outdoors
layman-naturalist, a keen observer. Mr. Shiras is emphat-
ically right in his complaint. Give the Alaskan form of
the whiskey-jack a third Latin name, by all means, to dis-
tinguish him in writing treatises exclusively for specialists
—if it gratifies them; but in books for general reading by
intelligent men call it the whiskey-jack, mentioning only if
necessary that the allusion is to the Alaskan form.

Let us illustrate what we mean by using as examples the
big cats: the lion, tiger, cougar, jaguar, and leopard. These
represent five very distinct animals. The last two come
closer together than is the case with any of the others, but
even as regards these the distinction is obvious. The use
of the word species as regards each of these big cats conveys
a definite, graphic, and easily grasped idea, which corre-
sponds to the facts in the case. But each is split into vari-
ous forms. The cougars of Brazil, Patagonia, the Andes,
Colorado, Oregon, and Florida all differ from one another,

PREFACE xix

and all have been made into species, the Latin names of
which convey no intimation that they do not differ from one
another as much as all collectively do from the lion, tiger,
or leopard. We think such a system of nomenclature a
mistake, even from the standpoint of the pure specialists.
It is a case for trinomial Latin nomenclature; call the ani-
mals all Felis concolor (or Felis puma, or by whatever name
the last delver into the bibliography of the subject has dis-
covered was the first name mentioned in some obscure book
a century ago), and add hippolestes, or oregonensis, or what-
ever other title is necessary to show what particular regional
form is meant. Follow the same course with the tiger and
the lion. Then we shall treat, as we ought to, of the big
cats as lions, tigers, cougars, leopards, and can add in each
case where it is necessary the limiting adjectives—Man-
churian tiger, Masai lion, Patagonian puma—without en-
cumbering ourselves with such an elaborate and minutely
divided terminology as serves merely for the darkening of
wisdom.

It would be well for ordinary purposes to adopt some-
what similar treatment for the word genus. It is an excel-
lent word when used to indicate both the resemblances of
different animals and their common differences from other
somewhat similar groups. It is highly desirable to include
only closely related forms in each genus. But if too much
split it loses much of its value. To make one genus of lions,
another of tigers, another of leopards, is to rob the word
genus of much of its value. Exactly as with species, it is
wise to remember that genus is a pure term of convenience.
In scientific treatises written by specialists for a limited
number of other specialists, there is no possible reason for

xX PREFACE

not using Latin names which will indicate genus, subgenus,
species, subspecies, and variety, if great exactness in the
conveyance of the idea is needed; there is, as we have al-
ready said, nothing sacred in the use of Latin binomial ter-
minology. But in ordinary scientific writing for laymen—
when we say scientific writing for laymen, we mean the kind
of writing which found its highest expression in the works of
such masters of science as Darwin and Huxley—it is better
to use terminology which is fairly clear and fairly easy to
understand. ‘This does not mean that we can revert to the
Linnzan genera and species; but it does mean that we can
try to use genera and species in somewhat the same sense
as Linnzus used them, of course keeping in view the enor-
mous mass of information which has been accumulated
since the great Swedish naturalist first reduced biology
from chaos to system.

In order to define the species more clearly we have made
all the geographical forms of subspecific rank by taking
the first described one as the typical race and repeating
the specific name. The first known form of any specific
group is necessarily originally described as a species, but
later, upon the recognition of additional races, the original
must be reduced so as not to confuse our conception of the
species. The Grant gazelle, for instance, was originally
described as Gazella granti from Ugogo in German East
Africa, and for a considerable number of years this re-
mained the only form described and the original locality
the only place from which it was known. Later, upon the
exploration of East Africa by sportsmen, many specimens
were collected in various parts of the country and gradually
the various races we now recognize were described, as our

PREFACE Xxi

knowledge, based upon the accumulation of specimens,
advanced. The original locality thus becomes the home of
the typical or first-described race, Gaxella granti granti, and
the other described races, such as robertsi, brighti, notata,
roosevelti, etc., occupy adjacent districts, and these areas
as a whole represent the range of the species or Grant
gazelle in its entirety.

Considerable difficulty has been encountered in map-
ping the ranges of the various species and races owing to
the vastness of the area covered and the lack of records
for many of the species over much of the area. The dis-
tribution of the game animals in the territory through which
the writers have travelled, particularly that of the highland
region of British East Africa, is well known, but beyond
this territory the distribution in a detailed way is unknown,
and it has been necessary to map it somewhat approx-
imately. The actual spots from which specimens have
been collected or recorded are marked on the maps by
numerals referring to each species and race, and these rep-
resent faithfully the real data upon which the mapped
areas are based. The interest in the large game mammals
in equatorial Africa is so keen among sportsmen that at
the present time their distribution is better known than
any other group of animals to be found in the same ter-
ritory. Owing to the present fairly satisfactory state of
our knowledge of the distribution of the game animals it
has been thought worth while to attempt to show their
ranges by means of maps. It is hoped that these maps will
lead to a clearer understanding of the distribution, and form
a basis upon which to build in greater detail. In cases
where the data have been insufficient or where a single

Xx PREFACE

form is considered no map has been prepared showing the
ranges of such species and races.

The sportsmen who have shot and written about East
African game animals have come largely from temperate
Europe or North America, and have to a considerable
degree looked upon the African game animals in the light
of their knowledge of northern species. In this way they
have often come to regard differences in coloration between
individuals of the same species as seasonal affairs, one coat
being attributed to the summer season and another to the
winter. As a matter of fact, however, the mammals of
equatorial Africa have no definite season for shedding their
coats nor are they subject to any seasonal climatic change
which would necessitate such a change. The climate over
the greater part of the highlands is uniform or so irregular
that no definite seasons can be said to prevail. On the
coast and in the lower desert districts flanking the coast
a more definite climatic division into dry and rainy season
exists, but the rainy seasons are usually two annually, and
so short that a mammal cannot adjust its pelage to the
change. In some species there is a great individual varia-
tion in coloration, as in the case of the male white-eared
kob and Nile lechwi, in which a large per cent of the indi-
viduals become black in adult life, but such changes are
never seasonal. Similarly erroneous ideas regarding a def-
inite breeding season in East Africa for large mammals
are quite prevalent among sportsmen, but there is no
climatic necessity for such a habit. Young of almost all
ages can be seen at the same season throughout the high-
lands, but there does seem to be in some species a certain
time when the young are born in greater numbers. In

PREFACE Xxili

the coast district and the low country, where the rainy
seasons are well marked, there may be a more definite
breeding-season, but upon this point we have no exact
knowledge, owing to the difficulties of observing game
animals in such bushy districts and the lack of data refer-
ring to such districts.

In the preparation of the present work Colonel Roose-
velt has written the first draughts of the life-history account
under each species, and of the introductory chapters on
Game Reserves and on Concealing Coloration. Mr. Heller
has prepared the first draughts of the other introductory
chapters, and of the technical descriptions (including the
nomenclature used, the coloration, and the faunal maps
showing the range of each species). But each has reviewed,
added to, and assented to the work of the other; and their
responsibility for the entire book is joint.

The Secretary of the Smithsonian Institution, Doctor
Charles D. Walcott, has kindly permitted the mammal
material in the National Museum to be drawn on freely for
study and description, and the data derived from this source
have been of invaluable assistance.

The photographs which have furnished the illustrations
have been received from various sources. Many of those
taken from life in East Africa and Uganda have been sup-
plied by Carl E. Akeley. Doctor William T. Hornaday has
donated for use a large number of photographs of the Afri-
can animals in the New York Zoological Park under his
charge. By far the larger number of photographs of heads
of game animals have been received from James L. Clark
and represent heads mounted in his studio in New York.
The groups and mounted specimens in the United States

XXIV PREFACE

National Museum have also furnished numerous illustra-
tions, as well as the specimens of live animals from Brit-
ish East Africa donated by William N. McMillan to the
National Zoological Park. For the photographs of the
McMillan mammals we are indebted to Mr. A. B. Baker,
the assistant superintendent of the park. The Field Mu-
seum of Chicago has supplied the photographs of the un-
rivalled groups of Somali mammals mounted by Carl E.
Akeley in that institution. To all of these the writers wish
to acknowledge their indebtedness.

THEODORE ROOSEVELT,

Epmunp HELLER.
New York, October 1, 1913.

CONTENTS

CHAPTER I

PAGE
ize (COUNTRY. AND» rs: HisTORY i) 6) 2 SOO a ee I

CHARTER

DERIVATION OF THE Fauna, GEOGRAPHICALLY AND PALEONTO-
POCICATE VY) MIRA acts Maen Commu sehr ih a) tet tei ac! |e. hy aa Re

CHAPTER Ii

Tue Fiora or East anD Mipp_e ArFrRIca AND Its RELATION TO

IAERESLUATINUAUMIL A Ties STS A, “ype ard dal cal Ca, teat lay heen tans SiS
CHAPTER IV
CoNCEALING AND REVEALING COLORATION AND THEIR RELATION
HOM NATURA. OREECIION) | yo Meas. /s,/\0y de. eat pees tlre eee aaa
CHAPTER V
SMM ERESER WIESE aioe 24 Soke ln BT Wels a ale te aes PES
CHAPTER VI
ER LACE: eee hse Nola anni Maia Een M2 eC RP a Oe See 6 76

CHAPTER VIE

kp SL EOPARD. AND: THE CHEERTAM 2) aio Me EWE ne a nd 229

CHAPTER: Vill

Tue Spotrep Hyena, StriPED HyENA, AND Huntinc Doc . . -251
xXxXV

XXV1 CONTENTS
CHAPTER IX

Pics 2
: CHAPTER X

Tue HrprorpoTaMus

CHAPTER (Xi

Tue RETICULATED AND CoMMON GIRAFFES

CHAPIER 2

Roan, SABLE, AND Oryx

CHAPTER, Sait

WILDEBEEST, DaMALIscus, AND HAaRTEBEEST

CHAPTER) XIV

BUFFALOES .

PAGE
270

289

301

348

404

ILLUSTRATIONS

On the Northern Guaso Nyiro River
Photogravure from a drawing by Philip R. Cee

Characteristic vegetation of East equatorial Africa
Lion group in the United States National Museum .
East African lions at National Zoological Park

Method of a lion’s attack on a zebra .
From a drawing by Philip R. Goodwin.

Cowboys roping a lioness

Races of the leopard

Rainey cheetahs .

Spotted and striped hyenas

East African hunting dogs

The forest hog and wart-hog of East Africa

Living specimens of hippopotamuses .

The races of the giraffe-—Masai, Uganda, and reticulated
The Masai and Uganda races of the giraffe

Defensive action of teticulated’giraffe =. 2. 6s 6) %
From a drawing by Philip R. Goodwin.

Reticulated giraffe
The races of roan antelope,—East Africa and Nile
Roosevelt sable

Showing individual variation in head markings in the Beisa oryx
XXVii

Frontispiece

FACING PAGE

44
166

XXVIll ILLUSTRATIONS

Mounted specimens of the topi damaliscus .
White-bearded wildebeest

Kongoni from the Athi Plains of British East Africa
East African and Nile races of the Lelwel hartebeest
East African buffalo

Living specimen of the East African buffalo

FACING PAGE

354
362
376
398
406

412

MAPS

1. Faunal areas of East equatorial Africa. (Color) \

2. Life zones of East equatorial Africa. (Color)
Distribution of the taces/of the lion), |. ws asia 6 ee ae
Distubution-of the races:of the eiraffe: ) 4. - 5 he ee
Distribution of the races of sable and roan antelopes .... .

3
4
5
6o* Discubution of the racesiof the Oryx ‘s55 =.) <3. 6) wh ee
7» Distribution of the races of the damaliscus . << - 9. *) =. «= °-
8

. Distribution of the races of the white-bearded wildebeest and hunter
ATCO MEM, denen kn! twom Conhotee ee Me ye armenia atic, Peat ie

9. Distribution of the races of the Coke hartebeest . . . . ..~ .
10. Distribution of the races of the Lelwel hartebeest. . . . .. .

11. Distribution of the races of the African buffalo . ......

. between pages 24-25

PAGE
227

319
S57
345
357

373
395

401

419

*,* The numerals 1, 2, 3, etc., on the maps mark the exact spots where specimens have been collected or
observed by reputable sportsmen, and the lines limiting the distribution are drawn around these numerals so

as to map the approximate area occupied by the race to which the numeral refers.
© Represents type locality or exact spot from which the type or original specimen came.

ate

nf

Rik GM i

LIFE-HISTORIES

OF

AFRICAN GAME ANIMALS

iti ta

| ail rom Ae

LIFE-HISTORIES OF AFRICAN
GAME ANIMALS

CHAPTER i
THE COUNTRY AND ITS HISTORY

Tue establishment of British protection over East
Africa and Uganda is of very recent birth, dating back
scarcely over a score of years. The history of the inland
exploration of the country is very little older. Only within
the last half century have we learned the more important
details of the topography of the interior, the position of the
great equatorial lakes and their connection with the Nile
watershed. It is indeed difficult to realize that a coast which
has been known to Europe since the fifteenth century could
hide from civilization for three centuries and a half all
knowledge of its hinterland. Ptolemy’s map, in the mid-
dle of the second century, actually gives a better idea of
this hinterland than that given by the European maps of
the middle of the nineteenth century.

The Portuguese navigator Vasco da Gama was the
first European to visit the coast. In 1493, during his cir-
cumnavigation of Africa, he touched at Mombasa and
Malindi, and later carried to Europe the first knowledge

concerning the East Coast. Mombasa and the neighbor-
1

2 AFRICAN GAME ANIMALS

ing coast for the following two hundred years was subject
to the control of various peoples. Sovereignty fluctuated
at various times between Portuguese, Turks, Arabs, and
negro tribes. Throughout much of this long period, during
which the country remained unknown, the seaports were at
war with civilized powers or besieged by the tribes living
near the coast and were effectually prevented from sending
any trading or exploring caravans into the wilderness. One
of the most powerful deterrents to inland travel were the
fierce Masai, who harassed and controlled the interior high-
lands. The occasional looting of the coast districts by
wandering bands of Masai kept the tribes who were in
touch with civilization in mortal fear of these savage
warriors.

The first Englishman to reach the coast was Commo-
dore Blankett, who touched at the mouth of the Juba
River in 1799, where a landing party from his fleet were
treacherously murdered by the natives. After this incident
he sailed southward to the island of Zanzibar, where his
fleet was well received. A half century later, in 1844, a
German missionary, Ludwig Krapf, recently expelled from
Abyssinia, entered Mombasa and established the first Chris-
tian mission. A year later he was joined by another Ger-
man missionary, John Rebmann, who founded a second
mission a few miles inland, among the Warabai tribe. Reb-
mann, with a few native followers, and armed only with the
faith he represented, penetrated inland in 1848 and dis-
covered the giant snow-capped mountain of Kilimanjaro,
journeying to within fifteen miles of the great volcano.
The dome or crater attains a height of nineteen thousand
seven hundred feet and can be seen from a few score miles

THE COUNTRY AND ITS HISTORY 3

inland, although situated some one hundred and seventy
miles from the coast.

The following year Ludwig Krapf made a longer jour-
ney inland and discovered Mount Kenia, the second highest
mountain on the continent of Africa. Kenia is situated
two hundred and fifty miles from the coast and attains an
altitude of seventeen thousand feet. Krapf did not reach
the mountain, but saw the peak from a distance of forty
miles from the Kitui district southeast of the peak.

The first Arab caravan to brave the perils of the far
interior and cross the much-dreaded Masai and Nandi
country reached the Kavirondo district in 1857. Within a
few days’ march of the shores of the Victoria Nyanza they
were stopped by the authority of the King of Uganda from
entering the country under his control, but they learned
from the natives of the existence a short distance westward
of a great body of water. During this year Speke, after
leaving Burton at Lake Tanganyika, which they had just
discovered, journeyed northward alone and reached the
southern shore of the great Nyanza.

Upon his return to England he organized an expedition
to further explore the vicinity of this newly discovered
lake, which he had shrewdly guessed formed the head-
waters of the Nile. With Grant he set out in 1862 across
German East Africa and eventually reached the lake.
They skirted the western coast to the vicinity of Kampala,
where they obtained guides from the King of Uganda, who
led them to the Ripon Falls where the Nile issues as a cata-
ract from the great lake. Speke and Grant, however, did
not trace the Nile from its birth at the Ripon Falls west-
ward to the second lake, the Albert Nyanza. Owing to

+ AFRICAN GAME ANIMALS

the difficulties or danger from native attack in the region
traversed by the Victoria Nile, the explorers chose a route
westward through the central portion of Unyoro and thence
northwest across the Victoria Nile to the Bahr el Jebel or
upper Nile. They thus missed by a few miles the discovery
of the Albert Nyanza of which the natives had given them
accounts. On their journey down the Nile they met at
Gondokoro Sir Samuel Baker who was ascending the river
on a mission of discovery concerning its source. He re-
ceived from Speke and Grant information concerning their
discoveries and of the native reports of another large lake
on the borders of western Unyoro. Baker soon afterward,
in 1864, ascended the Nile and discovered this lake which
he named the Albert Nyanza. The amount of natural-
history material brought to Europe by Speke and Grant is
really marvellous considering the limited means of trans-
portation at their disposal and the great length of their
journey. They brought collections of mammals, birds, rep-
tiles, fishes, insects, and plants, accompanied by extensive
field notes.

Subsequent explorers of the same district for the follow-
ing thirty years failed to equal these initial efforts to give
to the world a knowledge of the fauna and flora of equatorial
Africa. Stanley with the immense resources at his com-
mand is a notable example of this sort. He followed in the
paths cut by these two intrepid explorers, but added very
little to the discoveries made by his predecessors, and he
further accumulated only an infinitesimal number of speci-
mens or amount of information concerning the animal and
plant life of the regions he visited.

Speke and Grant collected fourteen specimens of hoofed

THE COUNTRY /AND ITS HISTORY 5

animals. Two of these were striking new species and were
named after the explorers: the sitatunga, Limnotragus
speke1, and the great-horned gazelle, Gazella granti. Sev-
eral other new species of big game were in the collection,
notably the white-bearded wildebeest and Baker’s roan
antelope, but they were named later from specimens secured
by more recent explorers.

While Speke and Grant were exploring Uganda, Heuglin,
a German naturalist, was penetrating the unknown region
of the Bahr el Ghazal, a western affluent of the White Nile.
Theodore von Heuglin was more naturalist than explorer,
his work being undertaken without any political or geo-
graphical incentive. He made extensive collections of the
mammals, birds, and plants of the upper Nile districts dur-
ing 1863-4. To this exploration is due the discovery of
the giant eland, the largest of all known antelopes. The
giant eland is of rare and local occurrence, and since its
discovery in 1864 remained quite unknown until within
the last five years, when it was rediscovered in the region
where Heuglin first obtained it. He also discovered and
named the lelwel hartebeest, the Nile bushbuck, and the
remarkable white-withered lechwi, since called Mrs. Gray’s
waterbuck. This latter species he failed to fix in literature
by a description, but simply gave to it a name, Adenota
megaceros. A fellow countryman, Fitzinger, in recording
the collection on its arrival at Vienna, mentioned the name
proposed by Heuglin and added a short description which
gives the name standing and fixes it for its discoverer.
Later Gray, of the British Museum, described it as Kobus
maria, from a specimen obtained subsequently by an Eng-
lish consul.

6 AFRICAN GAME ANIMALS

Following Speke and Grant we have a host of explorers
entering equatorial Africa successively down to the present
time. Doctor Georg Schweinfurth, a German botanist and
anthropologist, followed in the footsteps of Heuglin in the
Bahr-el-Ghazal region during 1863 to 1871. He described
the region and its inhabitants and added a list of the mam-
mals observed; but his efforts were chiefly devoted to the
botany of the region. Another German, Baron von der
Decken, ascended Kilimanjaro in 1862 to an altitude of
ten thousand five hundred feet.

In 1875 Stanley first appeared in Uganda as an explorer.
By following Speke and Grant’s old route from the East
Coast through German East Africa he reached the southern
shores of the Victoria Nyanza. His explorations were then
devoted to a circumnavigation of the lake and the mapping
of its shore-line, which gave to the world the first accurate
account of the lake and its approximate extent. After
finishing this task he continued on through Uganda and
explored the Albert Nyanza and the region southward to
Lake Tanganyika. Uganda was again visited by him in
1888 by way of the Congo on the relief to Emin Pasha. No
natural-history data, however, were gathered by Stanley,
whose work was chiefly geographical and political.

One of the most famous of, at all events the most unique
figure among, African explorers, is Emin Pasha or Edward
Schnitzer, best known by his Arabic name and title. Emin
was an explorer-naturalist of German-Hebrew extraction
who entered the upper Nile district in 1876. He displayed
remarkable administrative talents, and until 1890 reigned
as governor of the Bahr-el-Ghazal Nile and the adjoining
districts of the upper Congo, a post to which he was ap-

THE COUNTRY AND ITS HISTORY 7

pointed by Gordon. A series of his letters relating to the
political management, the people, and the fauna and flora
of the region have been published, and these testify amply
to his capacity as an administrator and a naturalist. At
various times he made collections of the smaller mammals
and birds of his provinces which he sent to the British
Museum, and these formed one of the earliest sources of
our knowledge of the smaller fauna of the Nile. The big-
game animals seem not to have interested him. One
proof of this lack of interest was his failure to report the
presence of so remarkable a beast as the white rhinoc-
eros, which was really abundant near the stations of
Wadelai and Kiro, which he occupied for some time while
governor.

The first naturalist to enter the territory which is now
British East Africa was the German botanist Hildebrandt,
who explored in 1875 the Taita district and the country
northeast to the Kitui district. Besides making collec-
tions of plants, he collected the smaller mammals, reptiles,
insects, and mollusks. Of the big game he collected the
horns of the impalla, the common waterbuck, and the black
rhinoceros. Soon afterward, in 1878, came Doctor G. A.
Fischer, another German naturalist. He first visited the
Tana River, which he ascended a considerable distance,
discovering during the exploration the small coast race,
closely allied to the Grant gazelle, known as Peters gazelle.
In the forests bordering the river he collected two new
species of apes, a colobus and a mangabey, neither of
which have since been secured by naturalists visiting the
country. Four years later, in 1882, he reached the Rift
Valley and discovered Lake Naivasha. During his eight

Sone AFRICAN GAME ANIMALS

years of travel in East Africa, Fischer collected many species
of small mammals and birds.

Joseph Thomson spent the years from 1883 to 1884 ina
journey of discovery through East Africa, proceeding from
Mombasa westward to Kilimanjaro and northwest to Lake
Naivasha. Journeying northward beyond Naivasha, he dis-
covered the much larger lake, Baringo. From this point
he travelled westward and was the first European to reach
the slopes of Mount Elgon. Leaving Elgon, he continued
on down the course of the ’Nzoi River to its mouth on the
northeast shore of the Victoria Nyanza. During his journey
he discovered the gazelle which was named for him by
Doctor Giinther from a few pairs of horns which he pre-
sented to the British Museum. His account of the animals
met with is very meagre, but the plants which he collected
during his travels received considerable attention in the
narrative he published.

We owe to Sir John Willoughby the first extensive ac-
count of the game animals of East Africa. Willoughby
together with Harvey and Hunter were the first men to
visit the country on an expedition solely devoted to the
shooting of big game. The region from Mombasa west-
ward to the slopes of Mount Kilimanjaro was visited by
Willoughby and Harvey, who reported for the first time the
presence of the gerenuk and both the lesser and greater
koodoo in the coast district of East Africa. Hunter made a
journey up the Tana River, where he discovered the peculiar
hartebeest, Beatragus hunteri, which has been named for
him, and further reported the first occurrence of oryx,
topi, and oribi in East Africa.

After the initial visit of these three sportsmen many

THE COUNTRY AND ITS HISTORY 9

others entered the country on similar missions. In 1888,
an American naturalist, Doctor W. L. Abbott, visited Kili-
manjaro, spending more than a year on its southwestern
slopes making collections of the mammals and birds of the
region. To his efforts are due the discovery of a rare forest
duiker, Cephalophus spadix, a large species related to the
West African yellow-backed duiker. This species has not
since been obtained by any sportsman visiting the district.
He also secured the first specimens of the fringe-eared oryx
and of the red forest duiker. Doctor Abbott made large
collections of both mammals and birds which still remain
the most extensive preserved from the region.

A year previous to Doctor Abbott’s expedition Count
Teleki, accompanied by Lieutenant von Hohnel, made his
remarkable journey through British East Africa, travelling
northward to the Abyssinian border, where he discovered a
large inland sea which he named Lake Rudolf, and a smaller
one northeast of it which he christened Lake Stefanie.
During this expedition he made the first ascent of Mount
Kenia, reaching the snow-line and naming some of the topo-
graphical features found near the summit. Later he as-
cended Mount Kilimanjaro to the ice cap. The trophies of
big game which he preserved, however, have never been:
classified. We have only the account of his travels, contain-
ing a brief list of his big-game trophies and notes on his
collections of plants and insects, to give us a knowledge of
his discoveries.

No one has contributed more to the literature concerning
east equatorial Africa than Sir Harry Johnston. This is
particularly true of the natural history. He first entered
the region in 1884 and made an exploration of the higher

10 AFRICAN GAME ANIMALS

altitudes of Kilimanjaro. The results of this expedition
were published a few years later in book form. From 1899
to 1901, as special commissioner, he explored Uganda and
ascended Ruwenzori. Later he published two volumes cov-
ering his own explorations in Uganda, and summarizing the
work of others in the territory. This work is the most
elaborate general account of the natural history extant.
Johnston secured in the Uasin Gishu Plateau the first
specimens of the five-horned giraffe, Giraffa camelopardalis
rothschildi, but this race was not distinguished until several
years later when the specimens collected by Powell-Cotton
led to the description of the race as new. ‘The explorations
of Johnston brought to light many new species of the smaller
mammals, birds, and plants.

The birth of British East Africa may be said to date
from 1888, at the time of the formation of the Imperial
British East Africa Company by royal charter from the
British Government. For the following seven years the
country was exploited commercially by the company. In
1895 the British Government took over the territory as a
protectorate and the work of constructing the Uganda Rail-
way was begun. During the tenure of the country by the
company the territory was often designated Ibea, a term
formed of the initials of the company. The word has since
come into use in zoology as a specific term for many of the
species discovered in the region.

One of the men whose name is closely associated with
the early history and discovery of many of the game ani-
mals is the present governor of Uganda, Mr. F. J. Jackson.
He first entered the country in 1887 on a shooting trip in
the region about the slopes of Kilimanjaro. Soon after the

THE COUNTRY AND ITS HISTORY ot

founding of the Imperial British East Africa Company
he proceeded, in 1889, on an expedition of exploration for
the company to Uganda. The old caravan route was taken
from Mombasa to the Kikuyu country westward across the
Rift Valley to the southern slopes of Mount Elgon and
thence to Uganda. During his long residence in East
Africa he has devoted much of his time to shooting expe-
ditions, which have given him a wider experience and knowl-
edge of the game of East Africa than that possessed by any
other sportsman. In the Badminton Library, Big Game
Shooting, he has given us the most comprehensive and
accurate account which we have of the field natural history
of the large game animals. To his zeal as a collector
is due the discovery of many of the described species
peculiar to East Africa. The large hartebeest, Bubalis
gacksont, which was named for him by Oldfield Thomas, was
discovered during his first Uganda expedition. Later he
discovered an allied species in central Uganda, Bubalis in-
signis. The first specimen of the bongo, a pair of horns,
which reached Europe were obtained by Jackson from
Wandorobo bushmen, but they were unfortunately erro-
neously assigned by Sclater to the nyala of South Africa.
Later Isaac, for whom the species was named, obtained more
complete specimens from the Wandorobo, which enabled
Thomas to discover the really close affinity of this rare for-
est species to the West African bongo. The white-bearded
wildebeest and the Kilimanjaro red duiker were further
discoveries based on Jackson’s collections. To his efforts
are due also many discoveries of new species among the
smaller mammals and birds of East Africa.

The Tana River and the desert region of East Africa were

12 AFRICAN GAME ANIMALS

visited in 1892 by an American sportsman, William Astor
Chanler, in company with Von Hohnel, who had accom-
panied Count Teleki some five years previously. Chanler
devoted his efforts to an exploration of the Northern Guaso
Nyiro River, which he followed from the slopes of Mount
Kenia to its termination in a swamp in the midst of the
desert. He discovered the existence of the Lorian swamp
and considered it to be the termination of the river, and in-
dependent from the Tana River system as well as the sea.
During the explorations in the vicinity of Kenia he dis-
covered the small species of rock reedbuck, Oreodorcas
chanleri, which has been named for him.

In 1894 another American, Doctor A. Donaldson Smith,
reached Lake Rudolf by a new route from the northeast,
and then continued southward across the Guaso Nyiro to
the Tana, and down the Tana to the sea. In addition to
big-game hunting he collected some twelve hundred speci-
mens of vertebrates, eighteen hundred insects, and three
hundred plants. Six years later, in 1900, this explorer made
another expedition from Somaliland west to the north end
of Lake Rudolf, and thence west to the Nile, at Fort
Berkeley, being the first white man to explore the region
between Rudolf and the Nile.

In 1893 we find two Neumanns appearing in East Africa,
one, Arthur H. Neumann, an English elephant hunter, the
other, Oscar Neumann, a German naturalist. A. H. Neu-
mann journeyed inland from Mombasa following the old
caravan route to Kitui southeast of Mount Kenia. From
this point he continued north, skirting the eastern flank of
the great mountain, and then travelled northward to the
Lorogi and Mathew Ranges, the flanks of which he followed

EMEP COUNTRY AND IYS HISTORY 13

to the shores of Lake Rudolf. Neumann was a mighty
hunter and well acquainted with the distinguishing char-
acteristics of the various species of game animals in the
country which he traversed. During his elephant shooting
he found time to collect specimens of many of the species
of big game, and to these efforts are due the discovery of
the reticulated giraffe, Neumann hartebeest and the
notata Grant gazelle.

Oscar Neumann entered British East Africa from the
German protectorate by way of the Rift Valley. From the
Rift Valley he journeyed across the Loita Plains to the
shores of the Victoria Nyanza, which he followed northward
to Mount Elgon and then west into Uganda. The route
followed by him upon his departure from the country was
very much the same as that taken by Joseph Thomson
some years previously and now commonly known as the
old caravan route to Uganda. From his collections have
been described the Uganda bushbuck, Tragelaphus dama,
the steinbok, Raphiceros neumanni, and the Masai lion,
Felis leo massaicus. Many small mammals and birds were
also discovered and described by Neumann.

During this same year, Doctor J. W. Gregory made a
geological and exploring tour of Mount Kenia and the Rift
Valley. While engaged in this work he made collections of
the smaller mammals, birds, reptiles, invertebrate animals,
and plants. A few years later, in 1897, Lord Delamere
entered the country from the north by way of the Galla-
land desert. Much of his time during this journey was
given to elephant hunting about Mount Marsabit and the
region north of the Guaso Nyiro River. Among the speci-
mens collected during this trip and presented to the British

14 AFRICAN GAME ANIMALS

Museum by Lord Delamere were two new species, the Somali
wart-hog, Phacocherus delamere1, and the desert bushbuck,
Tragelaphus delamerei. A notable discovery, in 1900, was
made by Major Gibbons, who discovered the white rhi-
noceros in the Lado Enclave while engaged on his explo-
ration of Africa from south to north.

Two years later Major Powell-Cotton entered upon his
exploration of the region northwest of Mount Elgon to the
banks of the Nile. He traversed the highlands bordering
the Turkana country and the region about Mount Agora,
much of which had never before been visited by a Euro-
pean. A detailed account of this region and the hunting
experiences which befell this sportsman are given in his
book, “In Unknown Africa.” The chief object of this
exploration was the gathering of specimens of the game ani-
mals inhabiting the district. Two subspecies of giraffe
have been named from this material, Giraffa camelopardalis
rothschildi and G. c. cottoni and the highland oribi, Ourebia
montana cottont, besides several other less distinct races.

The most recent big-game animal to be discovered in
East Africa is an obscure forest species, the giant forest pig.
Lieutenant Minertzhagen was instrumental in bringing this
remarkably distinct pig to the notice of naturalists. In
1905, while engaged in exploring the forest regions of Mount
Kenia, he secured from some of the Wandorobo pieces of
the skin of a large pig. Later he secured other similar
pieces of skin, together with two skulls of a similar pig in the
Kakumega forest lying at the base of the Nandi Escarp-
ment. These skulls enabled Oldfield Thomas, of the British
Museum, to determine the relationship of the animal, which
proved to be a new genus and quite intermediate between

THE COUNTRY AND ITS HISTORY 15

the very aberrant wart-hog and the common domestic pig
or wild boar. The pig has been named Hylocherus mi-
nertzhagent, and since its discovery, in 1904, several complete
specimens have been secured from the Wandorobo. Mi-
nertzhagen further secured, near Kenia, a new species of
oribi, Ourebia keny@, the range of which is confined to a
small tract of country skirting the southeast base of the
mountain.

The British Museum sent in I905 a natural-history
expedition to the Ruwenzori Range. The territory cov-
ered by the expedition was confined to the slopes of the
mountain, where no large game was encountered, but where
many new species of small mammals, birds, insects, and
plants were secured.

The Smithsonian African Expedition, under the direc-
tion of Colonel Theodore Roosevelt, made collections of
large-game animals in British East Africa, Uganda, and the
upper Nile districts during 1909 and 1910. The chief object
of the expedition was the collecting of specimens of all of
the game animals occurring in the territory traversed.
Among the specimens secured by the expedition were a
new subspecies of sable antelope, Egocerus niger roosevelt,
and three new races of hartebeest; a Nile race of Jelwel,
rooseveltit, a Rift Valley race of coke1, nakure, and also a
highland form, kongoni. In addition, several new species
of smaller antelopes were distinguished in the collec-
tion as well as a large number of small mammals and
birds.

During the past eight years large numbers of sportsmen
have shot over the game fields of British East Africa.
The highland region is, without doubt, the best-known big-

16 AFRICAN GAME ANIMALS

game field in the world. The game animals of this portion
of Africa are now almost as familiar to us as our own deer,
bison, or bear. Notwithstanding this familiarity with the
healthful highlands, there yet remain large tracts of country
in the desert lowlands which are either wholly unknown or
known only to a handful of hardy adventurers.

CHAPTERS I

DERIVATION OF THE FAUNA, GEOGRAPHICALLY
AND PALEONTOLOGICALLY

A—GEOLocIc FORMATION

THE greater part of the area comprised in British East
Africa and Uganda is of Archean age, being gneissic in
structure and derivation. All of the desert lowlands and
the higher grass-lands which flank them are comprised in
the gneissic area, together with the hills and mountains
which rise above them, including the lofty fault range of
Ruwenzori. Capping the central portion of the Archean
region we find a more recent volcanic or eruptive rock
series composed of various lavas. The whole highland
region of both East Africa and Uganda, including the lofty
volcanoes of Kenia, Elgon, and Kilimanjaro, are of this
formation. The Rift Valley has been the most recent
source of the lava cap which covers the highlands. The
lavas of this age have been erupted from fissures in the
vicinity of the Rift Valley, which comparatively recently
formed a great arch of eruptive rock, from the apex of
which the lava flowed east and west over the gneissic
region. Later the central portion of the arch dropped a
few thousand feet, forming the great rift, now known as
the Rift Valley. Muff in his account of the geology of
British East Africa * has described this formation as the

* Colonial Reports, Misc., No. 45, Jan., 1908, p. 27.
17

18 AFRICAN GAME ANIMALS

“falling of the keystone” of the great lava arch. Pre-
vious to this other masses of lava poured from the volcanic
centres of Kenia, Elgon, and Kilimanjaro, and these were
later augmented greatly by the Rift Valley flow. Elgon
and Kilimanjaro, both of which are crowned by well-pre-
served craters, are the most recent volcanic sources of this
lava flow, while Kenia, being much older, has been denuded
of its crater by weathering and is now capped by a pre-
cipitous peak representing the old volcanic neck. The
Rift Valley exhibits a score of smaller and still more recent
volcanoes; indeed, it is of such recent formation that its
many independent drainage systems have not been estab-
lished long enough to allow erosion to unite them into a
common system. The “falling of the keystone” which
established the Rift Valley has been of quite recent date.
The Ruwenzori Range, although in close proximity to dis-
tricts covered by recent lava flows, is a series of gneissic
or crystalline peaks formed by faulting and is without any
evidence of volcanic structure.

The only rocks not of eruptive or igneous origin are a
narrow belt of corals and sandstones bordering the coast
and extending inland from the seashore to a distance of
approximately fifty miles. The newest or most recent of
such formations is the coral limestone forming the coast
plain, which is of Pleistocene age. Dipping under the coral
formation we find a series of shales of Jurassic age, which is
exposed for several miles in width near the station of Chan-
gamwe. Beyond Changamwe successive exposures of yel-
lowish sandstone are encountered between Mazeras, Mari-
akani, and Maji ya Chumvi stations, and beyond these a
series of grits as far inland as the Taru Hills. The sand-

DERIVATION OF THE - FAUNA 19

stones and grits are the oldest sedimentary deposits and
have been assigned to Triassic age. The dip of all of these
beds is toward the coast as is that of the Archean gneiss
which underlies the Taru grits and comes to the surface
near Mackinnon Road station. The narrow coast belt of
sedimentary rocks is a very incomplete geological series
without any formations of Tertiary age or any older than
a doubtful Triassic sandstone. No vertebrate remains
have been found in any of these formations.

From the gneissic rocks have been derived the bright-
red gritty soil of universal occurrence in the desert nyika.
In the grassy uplands a black cotton soil is met with on
grassy steppes, notably the Kapiti and Athi Plains. Its
dark color is apparently due to vegetable humus and alka-
line salts from the red soil. The volcanic lavas are made up
chiefly of trachytes on the higher altitudes, with phonolites
underlying them, which are found exposed at the lower
levels near the Archean rocks. The lavas give rise to a
bright-red clayey soil containing a large per cent of ferric
oxide and resembling in color the red gneissic soil of the
desert lowlands. So wide-spread are these various soil con-
ditions over areas extending through several faunal zones
that no botanical or zoological significance in distribution
has been found attributal to their influence.

B—DERIVATION

In all the literature pertaining to zoology there is noth-
ing, perhaps, more unstable than the theories of derivation
based upon paleontological evidence. The discoveries of
new forms of animals or groups in new horizons and locali-
ties which are constantly taking place give a kaleidoscopic

20 AFRICAN GAME ANIMALS

effect to this field of theory. Notwithstanding the present
apparent futility of such speculation, participation in it is
irresistible. We may justify such indulgence of the fancy
as a groping toward the light or a legitimate attempt to
illuminate the obscurity surrounding the remote past. In
the northern hemisphere or holarctic region the paleo-
mammal records are at least partially complete in some
few groups, but in Africa the paleontological records are so
absurdly scanty that we are left with hardly a straw to
support our speculation. Africa is so dark paleontologic-
ally that we may almost consider ourselves engaged in
mere guesswork.

Localities in which mammal remains have been found
in Ethiopia, or Africa south of the Sahara Desert, are limited
to a few small deposits. In the Cape region are two de-
posits of Pleistocene age containing species allied to or iden-
tical with those existing to-day in the same district. The
most important of these localities is Barkley West, situated
on the Vaal River, near the boundary of the Orange River
Colony. Fraas described, in 1907, from this locality, a
species of Damaliscus, a hippopotamus, and a mastodon.
From the Zululand coast, in Natal, Scott* has described a
buffalo, a black rhinoceros, a hippopotamus, and an ele-
phant; all forms closely related to living African species.
Broom has recently described from the Orange River a
Pleistocene species of hartebeest allied to Coke’s, but ex-
ceeding it in size, and a large horse from the shore deposits
near Cape Town. More recently part of a mandible of a
Dinotherium, a primitive elephant bearing mandibular
tusks, together with remains of various other animals, have

* “Geol. Surv. Natal and Zululand,” 1907, p. 253.

DERIVATION OF THE FAUNA 21

been discovered on the eastern shore of the Victoria Nyanza,
near Karungu, British East Africa, in beds of Miocene age.
Other fragments supposed to belong to a Dinothertum have
been found in the lower Omo River near its entry into Lake
Rudolf.

North of the true African region we find extensive de-
posits of mammal remains near the Mediterranean. The
oldest of these are the Fayum beds of Eocene, Oligocene,
and Middle Miocene ages, situated near Cairo. In the
Eocene beds have been found remains of the toothed whales,
sirenians, and generalized mammals allied to tapirs and
elephants. Overlying these in the Oligocene beds remains
of giant hyraxes and primitive proboscidians have been
obtained. A large mammal, Paleomastodon, allied to the
elephant, has been found in the Miocene beds, together with
a primitive rhinoceros.

Mammals clostly related to the existing African species
are first met with in the Pliocene of Algeria. The water-
buck, the gazelle, and an antelope allied to the eland occur
in beds of this age, together with remains of a horse, H7p-
parion, a hippopotamus, and the sabre-toothed tiger. Dur-
ing the Pleistocene age which followed there existed in
North Africa many species of antelopes. Beds of this age
in Algeria contain the remains of several species of harte-
beests, two duikers, a reedbuck, several gazelles, two sable or
roan antelopes, a koodoo, one eland, and two species of buf-
falo. This is almost as great a number of species as we find
living to-day anywhere in Africa south of the Sahara Desert.

Our data for the determination of the geological history
of African mammals are thus very scanty. Almost the only
Ethiopian records are Pleistocene, which is too recent to con-

22 AFRICAN GAME ANIMALS

tain marked ancestral forms. North Africa, on the other
hand, has been since Pliocene time in close connection with
Europe and Asia and dominated by the same fauna. The
Pliocene fauna of North Africa was practically the same as
that occurring contemporaneously in southern Europe and
Asia. The same fauna exists in the game fields of East and
South Africa to-day. Before the Pleistocene dawned in Eu-
rope the great bulk of the antelopes and carnivores of Afri-
can affinities had disappeared and were replaced by a fauna
similar to that which exists there to-day. The deer, sheep,
ibex, bear, and boars which characterized this fauna en-
tered North Africa by way of the land connections at Suez
and Gibraltar, but were not able to cross the Sahara Desert
into tropical Africa. Africa in Miocene* time was cut off
as a continent from Europe, but unfortunately we have only
a few beds in the Fayum referable to this age. They con-
tain only primitive proboscidians and» rhinoceroses. In
addition to this evidence we have the Miocene Dinotherium
from the Victoria Nyanza. What evidence we have would
lead us to assume that the antelopes had not yet reached
Africa in Miocene time. Eurasia, however, in the Miocene
reached its climax of antelope forms, supporting eleven
genera, four of which still survive in Africa. During the
Oligocene and Miocene of Eurasia there appeared many
new genera either identical with or closely allied to those
found to-day in Africa. Such genera are represented by
the aardvark, the scaly ant-eater, the black rhinoceros, the
chevrotain, the springbuck, the gazelle, the sable or roan
antelope, the oryx, the koodoo, the bushbuck, the eland, the

* Mathew, “Outlines of the Continents in Tertiary Time,” Bull. Am. Mus.
N. H., XXII, P- 353, connects Africa during the Miocene with Eurasia; but the
distribution of antelopes during that time does not support such a connection.

DERIVATION OF THE FAUNA 23

giraffe, the hyena, the civet, and the mongoose. Nearly
all of these forms continued to exist in southern Europe and
Asia until the close of the Pliocene. The commonly ac-
cepted theory, however, derives the present African fauna
from southern Europe and Asia. The facts which we have
at present certainly support this theory, but the case is
not well balanced, Eurasia with its extensive Miocene beds
being weighed against the Miocene void in Africa. What
mammals inhabited Ethiopian Africa during middle or
early Tertiary times we do not know, as no fossil-bearing
beds of these ages have been discovered. It is certain,
however, that Africa is one of the oldest continents and may
well have been the birthplace of some groups of mammals.
Osborn,* from the evidence found in the Fayum beds, con-
siders it probable that Africa was the original home of the
elephants, sirenians, toothed whales, and hyraxes. Not-
withstanding the presence in the Mediterranean region of
Africa of early Tertiary generalized types of Proboscidea,
Hyracoidea, and ungulates, it is doubtful whether such forms
gave rise in any important degree to the present African
fauna. These Miocene forms exhibit such primitive char-
acters that their relationships with the modern fauna can-
not be well established. Owing to the incompleteness of
the fossil records, the direct ancestral forms of the living
antelopes are still unknown. The Miocene antelopes of
Europe and Asia were as specialized as the existing species,
while, in the older beds of Oligocene and Eocene age, allied
types are quite wanting.

The large island of Madagascar, which to-day has a very
peculiar fauna of lemurs, insectivores, and rodents, does not

* Osborn, “Age of Mammals,” 1910, p. 73.

24 AFRICAN GAME ANIMALS

throw much light on the derivation of the continental Afri-
can fauna. The present fauna of the island gives conclu-
sive evidence of its separation from the African continent
before the influx of the present big-game fauna, probably
some time before the Miocene. Just what its connection
was previous to this age is doubtful, but its fauna may, as
suggested by Sclater,* represent the ancient African fauna.
Since its separation several strays have reached it; in the
Pleistocene two species of pigmy hippopotamus and more
recently the bush pig. A fair per cent of its Chiroptera
have, without doubt, recently reached the island, so that
to-day its fauna is of a mixed character. This is equally
true of its earlier fauna. The lemuroids, which now char-
acterize it so remarkably, reached the island long after the
insectivores had attained their present specialization.

Immediately after the close of the Cretaceous period in
Africa there existed a connection with Eurasia and Mada-
gascar which allowed an intermigration of the land fauna.
During most of the Eocene, Oligocene, and Miocene, Africa
was isolated from Eurasia but connected with Madagascar.
This long interval gave ample time for the development of
specialization in the mammals of Ethiopia. But we are
practically without any records of the mammals inhabit-
ing Africa at this period. The fragmentary evidence of the
Fayum beds 1s quite insufficient for the building of a picture
of the life of the vast continent of Africa which was then in
outline quite as at present. Much of Africa has been dry
land since the Permian age, and the great bulk of the in-
terior is to-day covered by Archean rocks, which would indi-
cate a much greater antiquity.

* Sclater, “‘Geog. Mams.,” p. 108.

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DERIVATION OF THE FAUNA 25

C—PRESENT DISTRIBUTION

The big-game fauna of British East Africa and Uganda
is made up of three elements, or rather has been derived
from three different faunal areas. The chief source of this
derivation has been the South African region, to which
three-fourths of the species are closely allied. A consider-
able number are closely allied to Somaliland or northeast
African species, while a very few, chiefly forest animals, have
been derived from the Congo region. The large carnivo-
rous mammals and the elephants are so generally distrib-
uted that there is little faunal significance to their dis-
tribution. The faunal divisions recognized are on this
account based upon the hoofed mammals solely.

The region may be divided into two large faunal areas
which coincide with the two important drainage areas.
The greater of these in area and number of species is the
coast and Rift Valley drainage; the smaller, the Nile
watershed. The Rift Valley, though consisting of numer-
ous small independent lake-drainage systems is closely
allied faunally to the coast area, from which it differs by
almost no peculiarity. The real faunal barrier is west of
the Rift Valley, at the meeting of the Nile watershed and
these independent systems. Throughout the region this
barrier is marked by high mountains and plateaux. The
coast and Rift Valley drainage area is characterized in its
upper or highland portion by rolling, grassy plains flanked
on their higher parts by the forests which clothe the moun-
_ tain ranges of the district. On the lower borders the grassy
plains merge gradually into the dry scrub of the thorn-
bush deserts of the lowlands. In the immediate vicinity

26 AFRICAN GAME ANIMALS

of the coast the desert scrub gives way to a humid tropical
vegetation of palms and forest trees. The great herds of
game are the characteristic feature of the grassy upland
steppes. They occur less numerously in the desert scrub
and are quite rare in the forest. Many more species are
peculiar to the coast-drainage area than to the Nile water-
shed. To this region are confined the two gazelles, gerenuk,
impalla, sable, oryx, lesser koodoo, rock reedbuck, dikdik,
wildebeest, the reticulated and Masailand giraffes, the
common waterbuck, Coke hartebeest, Hunter antelope,
steinbok, and Haggard oribi. Within this area the most
important physical barrier is the Tana River. This stream
is broad and deep enough to act as a barrier across the
desert portion of the coast slope from Mount Kenia east-
ward to the sea. The aversion which most antelopes have
for crossing rivers is due no doubt chiefly to the fear of
attack by the crocodiles which haunt the streams. The
ungulates which range as far south as the Tana River but
do not cross to the south bank are Grévy zebra, reticu-
lated giraffe, Hunter antelope, beisa oryx, Rainey gazelle,
Ginther dikdik, and Somali wart-hog. We find a less
number of species reaching their northern limit on the
south bank. The best marked of these are Peters gazelle,
Masailand giraffe, Coke hartebeest, fringe-eared oryx, stein-
bok, and Masailand wart-hog.

The southern limits of the Tana or British East African
fauna are marked on the east by Kilimanjaro and the
ranges of Usambara and Pare which link it with the coast.
Westward the fauna is continued on down into German
East Africa by way of the Rift Valley as far as Ugogo, where
the lesser koodoo and fringe-eared oryx reach their southern

DERIVATION OF THE FAUNA 27

limit. The southern limits of the range of a northern carni-
vore, the striped hyena, coincides with that of the two
antelope, but the gerenuk is stopped by the highlands of
Kilimanjaro. Extending northward and meeting this fauna
in German East Africa we find two characteristic Zambesi
antelope, Lichtenstein hartebeest in the Rift Valley and
the sable on the coast slope, the latter penetrating as far
north as Mombasa.

The Nile watershed comprises all the country west of
the Mau escarpment or its continuation northward into
Abyssinia. The Mau proper is a high plateau extending
from the German border northward to the vicinity of
Mount Elgon, retaining an elevation of from eight thou-
sand to ten thousand feet unbroken. North of Elgon in
the latitude of Lake Rudolf it falls away and loses its char-
acter both as an escarpment and a faunal barrier. It rises
again in the Aybssinian highlands and effectually prevents
the desert lowlands of the Red Sea coast from coming in
contact with the Nile lowlands. The Nile faunal area is
better characterized by what it lacks than by any peculiar
animals it possesses. The kobs, genus Adenota, and the
Bubalis lelwel type of hartebeest are common to the whole
area, besides which there are a considerable number of dis-
tinct species peculiar to certain parts of the drainage sys-
tem. In contrast to the coast slope it lacks the dikdiks,
wildebeests, oryx, steinbok, rock reedbuck, gerenuk, ‘and
impalla. Gazelles occur only in its lower desert levels on
the northern frontier.

Two fairly well-marked regions may be defined within
this area due to differences in altitude and moisture. The
higher or Uganda region comprises the Victoria Nyanza

28 AFRICAN GAME ANIMALS

lake region and the territory extending northward and east
of the Nile. To this area are confined the common eland,
the quagga zebra, the Cape buffalo, and the klipspringer.
Uganda proper is to a large extent dominated by an
extensive growth of tall cane grass which forms suitable
cover only for buffalo and elephant. Northward, however,
and on the eastern or Mau drainage of the lake the open
plains and bush country support much game.

North of the Uganda region we meet with a drier and
lower region, the Bahr-el-Jebel Nile. In character this
region resembles the Soudan of which it is a part. It con-
sists of level grassy steppes interspersed with thorn trees
and bush and subject to a distinct wet and dry season.
The Nile in this region proves a very important faunal
barrier to much of the big game. To the region west of
the Nile which may be known as the Bahr el Ghazal dis-
trict we find limited the white rhinoceros, the giant eland,
the Congo giraffe, and the typical lelwel hartebeest. The
region east of the Nile or the Uganda Soudan has an even
larger number of species confined to it or rather prevented
by the Nile from entering the Bahr el Ghazal. Such species
have reached it recently from the southeast. The most
important of these recent arrivals are the black rhinoceros,
quagga zebra, common eland, Uganda giraffe, Roosevelt
hartebeest, and red-fronted gazelle. The Abyssinian buf-
falo, Nile oribi, roan antelope, Thomas kob, and Nile bush-
buck are common to both sides of the Nile.

Besides the regional distribution of animals into areas
limited by various physical barriers, we encounter within
the zoogeographical areas a zonal distribution which 1s
based on climatic and floral factors and is quite independ-

DERIVATION OF THE FAUNA 29

ent of other barriers. The zones occur in East Africa as
broad bands of territory extending parallel to the coast,
owing to the uniformity with which the land rises as we
proceed inland to the high central plateaux. Within each
zone the altitudinal and climatic conditions and plant species
are fairly uniform. The zones, owing to the irregular dis-
tribution of the flora, can be further subdivided into plant
associations such as grassy plains, thornbush thickets, or
dense forests. By using the zonal names in connection
with the plant associations, we are enabled to define very
precisely the habitats of the various animals inhabiting
the region. The zones are primarily upon a floral basis,
being founded upon the ranges of various trees and shrubs
to which the distribution of a majority of the animals con-
form. The zones with the plant species peculiar to them
are described at length in the chapter concerning the flora.

The whole region under consideration falls naturally
into five life-zones. Beginning at the coast, we find a nar-
row belt of lowland varying in width from five to twenty
miles and extending from the Lamu Islands southward.
This tropical belt is marked florally by the occurrence of
the cocoa-palm, and climatically by the greater amount of
moisture it receives in comparison to the desert which suc-
ceeds it. Peculiar to the tropical coast zone are the sable,
Egocerus niger roosevelti, the suni, Nesotragus moschatus de-
serticola, the blue duiker, Cephalophus monticola heck1, and the
maritime oribi, Ourebia montana haggardi. Thesable extends
no farther north than Mombasa, but the other species of ante-
lope extend to the northern limits of the zone near Lamu. The
sable and the oribi are more characteristic of the parklike
country bordering the groves of cocoa-palms than of the

30 AFRICAN GAME ANIMALS —

zone proper, but they do not extend beyond this intermedi-
ate strip into the dry desert. Other game animals occurring
in the zone, but not characteristic of it, are the yellow reed-
buck, Cape buffalo, the bushbuck, the elephant, the lion,
and the leopard.

Immediately succeeding the coast belt the desert nyika
stretches inland, covering almost one-half of the entire
region. In British East Africa it is broken by the elevated
highlands, but northward, in the latitude of Lake Rudolf, it
extends unbroken to the Nile. The nyika is low, the great
bulk of it less than two thousand feet elevation, with a hot,
dry climate and subject to a long dry season and one or
two short rainy periods annually. The whole region is
covered by scattered thorn-bush or by impenetrable jun-
gles of thorn trees, euphorbias, aloes, and sansevierias.
Grassy plains are rare, as well as absolute deserts devoid of
vegetation. The nyika zone resembles the Somaliland
country closely, and many of the animals which are char-
acteristic of it have been derived from this source. Its
two most peculiar and wide-spread types are the gerenuk
and the oryx. Others are Grévy zebra, Hunter antelope,
both the greater and the lesser koodoo, the reticulated
giraffe, Peters gazelle and Giinther dikdik. A host of
other game animals occur in the nyika, but are not confined
to it. Such are Coke and Neumann hartebeest, the topi,
the impalla, the common waterbuck, the bushbuck, Kirk
dikdik, the common eland, the Cape buffalo, the Kilimanjaro
giraffe, quagga zebra, the wart-hog, the bush pig, the ele-
phant, the lion, the leopard, the cheetah, and the hunting
dog.

The region most frequented by sportsmen is the high-

DERIVATION OF THE FAUNA 31

land veldt zone, which is characterized by its rolling prairies
of grass and its verdant appearance. Green rolling prairies
are, however, only one of its features. Much of it is cov-
ered by scattered growths of acacia trees or weird candela-
bra euphorbias. The altidudinal range extends from three
thousand to eight thousand feet, with correspondingly great
differences in temperature. The region, however, is uni-
formly well watered, but the rainy seasons are irregular
except in the western or Uganda section. There is scarcely
an antelope which is peculiar to the whole zone, although
many are peculiar to certain parts of it. Antelope which
are found only in the veldt zone are the wildebeest, Thom-
son gazelle, Jackson, Uganda, Kongoni, and Nakuru
hartebecsts, klipspringer, steinbok, and the five-horned
giraffe. Many others share this zone with the nyika.
Such are the black rhinoceros, quagga zebra, common
eland, topi, bushbuck, waterbuck, kob, roan, dikdik, oribi,
reedbuck, suni, and various species of duiker, wart-hog,
and bush pig.

Extending above the veldt zone we find the highland
forest, ranging from six thousand to ten thousand feet in
altitude. It is a uniformly dense assemblage of trees of
mixed tropical and mountain affinities and strikingly dif-
ferent in appearance from our northern forests. The well-
known families to which our oaks and beeches belong are
not represented, their places being taken by trees belong-
ing to families of the higher flowering plants peculiar to the
tropics. The pines and firs are likewise lacking, the only
“representatives of the conifers being a single species of
juniper and two species of African yew. The rose family,
to which most of our fruit-trees belong, is likewise conspicu-

32 AFRICAN GAME ANIMALS

ous by its absence, being represented by only a few species
of brambles and alpine shrubs. The forest is a mixture of
many species of trees, vines, and undershrubs. The chief
forest-trees are species of figs, olives, camphor, cloves,
sapote, and various genera related to the euphorbias,
acacias, and coffee. Various species of large vines link the
forest together, while prickly undershrubs allied to Acan-
thus add to the impenetrable nature of the tree growth.
In the higher altitudes the yew and juniper predominate
and give the elevated region a more northern aspect. Bor-
dering the forest at its upper altitudes is a dense zone of
bamboos, which at a distance has the appearance of fields
of tall grass. The bamboo 4rundinaria alpina occurs from
nine thousand to ten thousand feet as a pure growth to the
exclusion of all other vegetation, not excepting under-
shrubs. It forms a waste quite uninhabited by animal life,
although elephant, and very rarely roan antelope, at times
penetrate it.

The bamboo is lacking on Kilimanjaro and the high
mountains in its vicinity. In the Nile district of Uganda
and the Lado Enclave a lowland bamboo occurs, Oxytenan-
thera abyssinica, which is a member of another genus and
quite distinct from the peculiar high-mountain species
found on Kenia, the Aberdares, the Mau and Kikuyu
escarpments, Mount Elgon, and the Ruwenzori Range.

Peculiar to the forest we have the bongo, the giant for-
est pig, and Abbott duiker. The dense nature of the
forest has made it possible for these animals to escape dis-
covery until within the last few years. They are still
nearly unknown to white hunters, being familiar only from
the specimens shot or pitted by the Wandorobo bushmen.

DERIVATION OF THE FAUNA 33

But one or two instances of either the bongo or forest pig
having fallen to the rifle of the white man have thus far
been recorded. The Abbott duiker is still only known by
the original specimen secured by Doctor Abbott a score of
years ago in the forests of Kilimanjaro. These three ex-
clusively forest animals are of West African affinities and
have reached East Africa by way of the straggling forests
through Uganda and the upper Congo. Other game ani-
mals which are confined to the forest are the red duiker,
the bush pig, the suni, the bushbuck, the elephant, and the
leopard, the last being the only large carnivore which
enters the forest. Many of these game animals are found
widely distributed in the lowland forests of Uganda and
the tropical coast belt.

The moorland zone succeeds the forest on the high
mountains of Kenia, Kilimanjaro, Ruwenzori, Elgon, and
the Aberdare Range. It extends from the upper limits of
the bamboo at ten thousand feet to the lower margin of
the snow-fields at fourteen thousand five hundred feet.
The country between these altitudes is an open moorland
of boggy soil overgrown by dense masses of shrubby al-
chemillas and tussocks of reeds and grass. Tree-heaths
occur throughout the lower levels of the zone and gro-
tesque tree-like groundsels and lobelias in the upper alti-
tudes. Great numbers of rodents occur in this high, cold
region, but the antelopes are represented by only a single
resident species, the Abyssinian duiker, Sylvicapra, grimmi
altivallis. ‘The Cape buffalo occasionally reaches the moor-
land as a straggler. There is a record of one on Kenia
which perished recently in the snow at fifteen thousand
feet, where its remains still lie. Elephants are also oc-

34 AFRICAN GAME ANIMALS

casional visitors, their tracks being quite numerous in the
boggy soil of the moorland. Two carnivores occur, the ser-
val cat, which is resident in the moorland, and the leopard,
which is simply a visitor in the region.

CHAPTER Ch

THE FLORA OF EAST AND MIDDLE AFRICA AND ITS
RELATION TO THE FAUNA

Two floral types stand out conspicuously in East Africa
and give the landscape its peculiar African character.
These two, the acacias and the giant euphorbias, are the
dominant tree forms, and are wide-spread throughout the
steppe and bush country. Although both genera are cos-
mopolitan, there is no other country in which the euphorbias
assume the giant, candelabra-like tree growths or where
their association with flat-topped acacia trees occurs.
These peculiar tree growths extend throughout the whole
region occupied by the game fields, and dominate the real
Africa of the sportsman. Both of them have, however,
received erroneous names at the hands of sportsmen. The
acacias are usually referred to as mimosas, a genus which
occurs in the region, but only as a small prickly shrub in
swampy districts. The tree euphorbias are often referred
to as cactuses because of their close resemblance in form
to the giant species of the American deserts, with which
they have no real relationship, being closely akin to some
of our common roadside weeds, the spurges. In the lower,
drier parts of this region two other conspicuous tree growths
appear which are much more peculiarly African, but are
less wide-spread. These latter, the grotesque gouty bao-

bab and the branching doum-palm, are not found beyond
35

36 AFRICAN GAME ANIMALS

the Ethiopian region. The country dominated by these
four arboreal types extends from the coast inland to an ele-
vation of seven thousand feet, covering practically all the
region except a few forested highland areas and a narrow
coast strip. The forests of the highlands and mountains
are composed largely of trees which belong to peculiar
African genera, but exhibiting no such peculiarity in ap-
pearance as the baobabs and the branching palms. The
great bulk of the plants which occur in East Africa are
members of genera found in India or are wide-spread in
tropical Asia and America. The vegetation is nowhere
tropical or luxuriant in growth except in the highland for-
ests. Even at the coast the rainy season is short and the
country has a semiarid appearance. The greater part of
the lowland country below four thousand feet is desert in
character with xerophytic plant forms chiefly.

The thorny acacias in either tree or bush form are dom-
inant everywhere below an altitude of six thousand feet or
outside of the forested areas. They are primarily plains or
riverside trees and are never associated compactly into
dense forests. The Leguminose are represented by many
other genera in the region under consideration, and exceed
the other families in number of species. Next in number
of species is perhaps the Rubiaceae, of which the coffee-bush
is typical; it occurs as one of the native shrubs in Uganda.
The great bulk of the smaller trees and shrubs of the high-
land forest belong to this family. The Rosacee are only
represented in the high mountain forests and alpine regions,
where many of the moorland shrubs and a few trees belong-
ing to the order are found. The great bulk of the trees
and shrubs belong to the tropical orders of Sapotacea,

FLORA OF EAST AND MIDDLE AFRICA 37

Apocynacea, Burseracee, Tiliaceg, and several others, all
of which are unfamiliar to botanists from north-temperate
zones. The conifers are found only in the highlands, where
they are represented by a typical juniper, a tree of wide
range, and by two species of yews of the tropical genus
Podocarpus. Palms are quite an inconspicuous feature in
East Africa; but four or five species occur and they are
only abundant locally.

Large areas in the highlands of British East Africa and
in Uganda and the Nile Valley are covered by a heavy
growth of grass, forming treeless plains of considerable
extent. One or two species of grass usually predominate
in the plains, but many different species are represented
in the various districts where grass veldt occurs.

The whole region falls naturally into five floral or faunal
zones based primarily on climatic differences due to mois-
ture and altitude and characterized by peculiar tree or plant
species. The zones succeed one another as the country
rises from the coast to the highlands in the interior and
extend parallel with the coast as broad bands of territory.
_ The higher zones, which are found only on mountain slopes
or summits, occur as islands in the lower zones.

The animal life is dependent chiefly upon certain plant
species, and on this account conforms quite closely in
distribution to the floral zones in which such species occur.
Throughout the zones the distribution of the plants com-
prised in each is by no means uniform, for each zone is made
up of several distinct plant associations. Because of the
dependence of the animals upon certain plant associations
they seldom occur generally distributed in the zone. Plant
associations are really the most important floral factors in

38 AFRICAN GAME ANIMALS

the distribution of animals and are the most precise means
of defining their ranges.

Extending along the coast from Lamu southward is a
narrow strip of country which is well characterized by the
groves of cocoa-palms which grow luxuriantly on the low
coral coast plain and the hill country inland to an altitude
of five hundred feet. This stretch of low country may be
known as the tropical coast zone. It receives consider-
ably more rainfall than the dry desert country flanking it,
and supports a luxuriant vegetation during the short rainy
season, but is semiarid in character, with little semblance
of tropical luxuriance. The cocoa-palm marks the zone
precisely, extending from the seashore inland to the sum-
mits of the low coast hills.

On the Uganda Railway the zone extends from Mom-
basa to Mazeras, having a width here of fifteen miles.
Farther north it narrows down to a few miles in width,
owing to the coast hills approaching the coast and confining
the coastal plain. At the Tana River it broadens again and
extends up the alluvial valley a score of miles. Still far-
ther north beyond Lamu it gives way entirely to the desert,
which here reaches the seashore. Several plant associa-
tions are found within the zone. A mangrove association
composed largely of three species of mangroves, Rhzzo-
phora mucronata, Bruguiera gymnorrhiza, and Avicennia
officinalis, occupies much of the territory about the inlets
and the tidal portions of streams. Forests occur in places
on the slopes and crests of the coast hills. They are quite
dense assemblages of a variety of trees, chief among which
is the giant Trachylobium hornemannianum, a wide-spread-
ing, white-barked tree. Other common forest-trees are

FLORA OF EAST AND MIDDLE AFRICA — 39

Afzelia cuanzensis, and species of Combretum and Albizzia,
which are mixed with an undergrowth of scrubs allied to
Acanthus. Much of the tropical coast zone is covered by
scattered bush and grassy areas, interspersed with solitary
trees of Acacia stenocarpa, Bauhinia reticulata, Erythrina
tomentosa, baobabs, and occasional doum-palms and cande-
labra euphorbias.

Succeeding the tropical coast belt we find the desert
nyika zone stretching far inland as a dry, thorn-bush-
covered steppe. The nyika covers the greater part of the
region of east equatorial Africa and in it are found most of
the floral and faunal peculiarities of East Africa. All of
the country lying between the altitudes of five hundred
and three thousand feet are comprised in it. The Uganda
Railway traverses the zone from the station of Mazeras to
Makindu, a distance of one hundred and sixty miles actually,
but by the railroad some two hundred miles. Northward
the nyika broadens out tremendously, attaining in the
region north of Mount Kenia a width of over four hun-
dred miles, and extending unbroken from the seashore
inland past Lake Rudolf to the Nile. The region is quite
level, with a gentle slope to the coast, and occasional
isolated hills or small mountains of gneiss rising a few
thousand feet above the plain. A heavy, bright-red clay
soil covers the region universally, except where the gneiss
rocks project above it.

The nyika is subject to a long, dry season during which
most of the trees and bushes are leafless and the soil is
bare. The vegetation is on this account fitted in every
possible way to resist evaporation and is typically xerophytic.
During the short rainy season, which is irregularly dis-

40 AFRICAN GAME ANIMALS

tributed from April to June, the country blossoms out lux-
uriantly, all of the naked bushes and trees leafing out and
the ground becoming covered with a wealth of flowering
herbs and grasses. The transformation is so complete that
any one passing through the region at this time cannot con-
ceive of the desolation which is characteristic of the dry
season. The two most characteristic trees are the massive
baobab, Adansonia digitata, and the branching doum-palm,
Hyphene coriacee. Associated with these and scarcely less
characteristic are the two giant euphorbias, Euphorbia can-
delabrum and Euphorbia nytke, and several species of flat-
topped acacias. One of the characteristic shrubs of the
region, and one which impresses itself indelibly upon the
memory of the traveller, is the wait-a-bit bush, 4cacia mel-
lifera, which is armed with short recurved thorns. The tree
acacias are chiefly Acacia tortilis, which form groves of large
trees along dry watercourses, and Acacia verugosa, the
green-barked acacia, which is found near running water or
where the conditions are moist. A striking but irregularly
distributed plant in this zone is a cycad, Encephalartos hilde-
brandti. The great bulk of the bush vegetation is made up
of various species of Grewia, Commiphora, Euphorbia, Aloe,
Sansevieria, and Acacia.

The plant associations we meet with in this zone are
dense thorny bush, grassy plains with scattered tree
growths, and riparian flora. The nyika is characterized
primarily by the uniform distribution of its thorn-bush.
Much of the bush grows in dense thickets, which can only
be penetrated by cutting a way through with a knife or
axe. Such thickets are made up largely of small acacia
trees, the dagger-like leaves of several species of Sanseveria

FLORA OF EAST AND MIDDLE AFRICA 41

and Aloé, arborescent and twining euphorbias, and thorny
Commiphora bushes. Immediately bordering the coast
tropical belt a grass-covered park-like prairie is found, where
the bushes grow far apart and where there is an increase of
moisture sufficient to allow a fair growth of grass during
much of the year. The sable, Haggard oribi, and the
Swahili reedbuck are limited in their distribution to this
transitional area. Grassy or park-like plains of small extent
occur at intervals throughout all of the nyika, but they
form a very small proportion of the area. North of Mount
Kenia in the desert country which extends northward to
Abyssinia and Somaliland, we find the riparian vegetation
quite different from that of the coast nyika, where the dry
watercourses are bordered chiefly by green-barked and flat-
topped acacias, with only rarely a doum-palm. In this
more northern desert we find a heavy growth of doum-
palms bordering the dry stream beds and marking their
courses amid the low acacia and bush scrub of the desert.
Associated with the palms are thickets of the gray Salva-
dora persica bushes and groves along the smaller stream
beds of Acacia tortilis trees. The wait-a-bit, Acacia melli-
fera, with its spreading branches and hooked thorns reach-
ing out in every direction to grasp the passer-by, covers
most of the desert together with Grewia populifolia and
thorny Commiphora bushes. The baobab is absent from
this district.

On the sides of the hills and mountains this steppe veg-
etation is augmented by groves of tall, palm-like Euphorbia
nyike or the columnar Euphorbia candelabrum, the flat-
topped Acacia stenocarpa, the papery-barked Boswellia
trees, and a few aloe bushes.

42 AFRICAN GAME ANIMALS

We have now reached the high veldt or prairie zone,
where the great herds of antelope and other hoofed game
are found most abundantly. Open grassy plains are char-
acteristic of this elevated region, but nowhere are the grass
areas of great extent. The highland veldt is generally of
park-like appearance, interspersed by small grass plains
which seldom exceed a score of miles in breadth. It is a
distinctly rolling country and in places rough and much
broken by cliffs and rocky ravines. The predominating
soil is the red clay of the nyika, with the exception of the
plains area, which is covered by a black cotton soil support-
ing a heavy growth of grass. The older hills and moun-
tains are of gneiss, but much of the country is more re-
cent and covered by lava rocks. These predominate in
the Rift Valley, especially where much of the country is
traversed by trachytic cliffs and broken by recent lava
flows.

The high veldt zone covers the greater part of the
country between the altitudes of three thousand two hun-
dred and nine thousand feet. In its upper levels it extends
into and well above much of the forest as peninsular or in-
sular areas. Extensive areas of grassy veldt occur on the
summit of the Mau plateau at eight thousand and nine
thousand feet confined by the wall-like edges of the dense
highland forests. On the northern slopes of Mount Kenia
the veldt region breaks the forests’ girdle of the mountain
and extends as a wide peninsula upward to the alpine moor-
land where it attains an altitude of ten thousand feet. The
highland veldt zone is much less in area than the nyika.
It covers the head-waters of the coast streams, the greater
part of the Nyanza drainage, and the Uganda region west-

FLORA OF EAST AND MIDDLE AFRICA 43

ward almost to the shores of the Albert Nyanza and the
Bahr-el-Jebel Nile.

Much of the tree vegetation which is characteristic of
this zone occurs also on the border of the forested regions
rising above it. The quaint straggling Croton elliottianus,
the African yew, Podocarpus gracilior, the olive, and Albizzia
fastigata are trees of this sort. A very characteristic tree
or small shrub is the gray sagebush-like camphor-bush,
Tarconanthus camphoratus, which covers large areas in the
Rift Valley, Loita Plains, and North Kenia districts. Asso-
ciated with this are Carissa bushes, trees of Acocanthera
abyssinica from the wood of which the deadly arrow poison
is derived, a small, slender. tree, Dodonea viscosa, and sev-
eral species of the oak-like Combretum trees. From the
nyika we have as stragglers both of the giant euphorbias
and several tree acacias which are wide-spread in the zone
and give it a characteristic African appearance. Kigelia
ethiopica with its pendulent sausage-like fruits is a com-
mon tree in the Nile watershed.

The grassy plains which are the characteristic plant
feature of the zone are covered by a heavy black cotton soil
in marked contrast to the red soil of the hill and forested
areas bordering them. -During the rainy season a luxuriant
growth of grasses a few feet in height covers the plains, but
gradually it is grazed down by the great herds of game
until only mere stubble is left in the dry season. The most
extensive grass plains included in this area are the Athi,
Kapiti, Loita, Laikipia, Naivasha, Uasin Gishu, and the
whole eastern Nyanza lake steppe.

A characteristic bush of this region is the gall-covered
Acacia fistula which forms a scattered growth over much

44 AFRICAN GAME ANIMALS

of the plains region. The lower altitudes of the zone
are typically acacia parks, grass interspersed with bush
country and scattered trees. The elevated grassy plains
above seven thousand feet, such as occur on the Mau sum-
mit and on Kenia, are covered by a dense perennial growth
of grass which crowds out all other vegetation.

The riparian vegetation is dominated by the yellow-
barked tree, Acacia verugosa, which forms a narrow fringe of
forest along all the streams. Associated with this are often
trees of the rough-barked Acacia stenocarpa. In the lower
levels of the streams on the nyika border the large red-
fruited fig, Ficus stuhlmanni, is a conspicuous tree. In
this region the immense fronds of the Raphia palm are
often seen among the riverside trees. Throughout the
Uganda region and also much of East Africa the tall feath-
ery papyrus is the dominant swamp and watercourse plant
and is characteristic of this zone. On the Nile, however,
it covers large areas far down into the nyika to an altitude
of fifteen hundred feet. At the higher levels of the veldt
region the African clove, Eugenia cordata, grows abundantly
on the banks of streams, and with it are occasionally asso-
ciated a few willow-trees. A conspicuous riparian bush or
small tree is the Cassia didymobotrya, with its perennial
wealth of large yellow blossoms.

In East Africa proper, that is, in the Rift Valley and
coast drainages, no forests occur in the high veldt zone,
but in Uganda much of this zone is occupied by a tropical
forest in character similar to that which spreads over the
Congo basin. This forest is found in patches from its east-
ern extreme, the Kakumega forest at the base of the Nandi
Escarpment westward to the Semliki Valley and the Congo

9 : PEt bane

Euphorbia Forest, Euphorbia nyike Sausage Tree, Kigelia ethiopica
Amala River headwaters Nzoia River, B. E. A.

Flat-topped Acacia, Acacia verugera Candelabra Euphorbia
Acacia Forest near Lake Naivasha Loita Plains, B. E. A.

¥ Shak : oe

Haunts of Bongo, Forest Hog, and Red Duiker |
Highland Forest on Mau Escarpment near Ravine Station

Giant Groundsels, Senecio keniensis Dragon Tree Doum Palm

Alpine vegetation on Mt. Kenia at 13,000 feet Dracena papahu (Ivory Nut Palms) Hyphene
Nyeri, West Kenia coriacea

CHARACTERISTIC VEGETATION OF EAST EQUATORIAL AFRICA

FLORA OF EAST AND MIDDLE AFRICA . 45

drainage. It is continued westward along the shore of the
Victoria Nyanza around the west shore of the lake to Buddu
and thence west to Ruwenzori and the Albert Nyanza.

The conifers and olives of the East African highlands
are lacking in these lower altitudes, where the trees are
chiefly Sapotacee, Meliaceeg, and other tropical orders.
The forests are often composed of gigantic trees which grow
in dense clusters, but in great variety of species. On the
edge of the forest the brilliant red flowers of the Spathodea
tree are often found associated with a fringing growth of
red-berried sumac-like Haronga trees. In open spaces in
the forest, fields of the wild banana, Musa ensete, occur.

The highland forest covers much of the high plateau
region between the altitudes of six thousand and ten thou-
sand feet. In British East Africa it extends roughly from
Mount Kenia westward and southward to the Aberdare
Range and the Kikuyu Escarpment above Nairobi, where it
reaches its southern point. In the vicinity of Lakes Nai-
vasha and Elmentaita it straggles across the Rift Valley
to the great Mau forest. On the Mau Plateau the forest
extends nearly unbroken from the German border north-
ward to the latitude of Mount Elgon. The Elgon forest
in its lower levels is connected with the Mau by way of
the ’Nzoia River drainage. From the Mau crest the forest
continues down the west face of the Nandi Escarpment,
where it is continuous with the Kakumega forest, one of
the tongues of the Uganda forest, which in its turn is con-
tinuous with the Congo forests in the Semliki Valley. The
Kakumega and Uganda forests are, however, lowland and
quite different in character from the highland forests, but
the continuity of the whole area of tree growth has allowed

46 AFRICAN GAME ANIMALS

the extensive dispersal of forest animals. By using the
shelter of riparian tree growth it is possible for an animal
to travel within the forest from eastern Kenia westward
to the Congo watershed. By this route several West Africa
game animals, notably the bongo, giant pig, red duiker,
and Abbott duiker, have reached the coast drainage of
East Africa. Formerly this forest was much more extensive
between the Aberdare Range and Mount Kenia, where large
areas of the intermediate region have been deforested by
the Kikuyu tribe in order to give them new soil for cultiva-
tion. Similarly destructive agricultural methods have sep-
arated portions of the Elgon and Nandi forests, which were
formerly continuous. Isolated patches of this forest area oc-
cur in remote regions like the Taita Hills in the coast nyika.
To the north of Kenia isolated patches of forest again occur
on Lololokwi, Uarages, and many other peaks of the Mathew
Range as far northward as Lake Rudolf. In the extreme
southeast lie Kilimanjaro and Usambara on the German
border, covered by an extensive forest area which is isolated
by many miles of desert from the highland forest of Kenia.
Westward in Uganda on the elevated slopes of the Ruwen-
zori Range the same type of forest exists above six thousand
feet altitude. Ruwenzori is surrounded by, or rather its
snow-crowned peaks are set in, a wide belt of highland for-
est. All of the highland forest is in the volcanic area and is
underlain by lavas, chiefly trachytes and phonolites. Within
the forest the soil is a black leaf-mould, but when this is
removed a red clay soil is exposed, which is somewhat dif-
ferent in composition from that so universal in the nyika.
Owing to the great quantity of ferric oxide in the soil,
weathering converts it into a hard iron pan formation

FLORA OF EAST AND MIDDLE AFRICA 47

which cannot again support tree growth, and on this ac-
count deforestation in the highlands of East Africa is
attended by irreparable devastation. The only highland
forests outside of this volcanic area are the isolated patches
on the Taita Hills, the Mathew, Ruwenzori, and Usambara
Ranges, which are in a gneissic region.

The forest is made up of a great variety of trees, only a
few of which occur universally throughout itsarea. The yew,
Podocarpus milanjiana, and the black olive, Olea laurifolia,
are the only trees of this class which can stand the extremes
of moisture existing between the moist east and the dry west
slopes of the forested mountains. A wide-spread tree of
the dry western slope is the Abyssinian juniper, Juniperus
procera. A more important distributional factor than the
division of the forest into a dry western and a moist eastern
section is formed by the altitudinal assemblages of trees we
find characteristic of the lower edge of the forest, the inner or
true forest and the upper or timber-line edge of the forest.
Characteristic of this lower edge of the forest, which fringes
the open, grassy plains or continues as long tongues border-
ing streams down into the plains country, are the straggly,
spidery Croton elliottianus, the Brachylena (sandalwood),
and the acacia-like Al/bizzia fastigata. A conspicuous tree
of the forest edge is Dombeya nairobiensis, which is usually
covered by a wealth of white apple-like blossoms. On the
moist slopes we have added to these the large-leaved shrubby
tree, Macaranga kilimanjarica, the columnar Cusonia spicata,
tree-ferns, Lonchitis pubescens, and the peculiar tree-lilies,
Dracena. The poison-bush, Acocanthera abyssinica, the
gray olive, Olea chrysophylla, the yew, Podocarpus gracilior,
and the widely distributed juniper, Juniperus procera, are all

48 AFRICAN GAME ANIMALS

confined to the drier western forest edge. The grassy plain=z
do not, as a rule, merge gradually into the forested area
through an intermediate park-like country. The forest
trees seem capable only of growing in masses where they are
protected from the maximum quantity of sunlight. This
massed condition of the forest is particularly in evidence
on the Mau summit, where the grassy plains are bordered
by a solid wall of trees as dense and as tall as the central
parts of the forest.

The true forest is composed largely of two wide-spread
trees, Podocarpus milanjiana and Olea laurifolia, and on
western slopes often exclusively of Juniperus procera. As-
sociated with these trees is found the giant of the forest,
the camphor-tree, Ocotea usambariensis. Another tree of
immense size and wide-spread distribution is the killer fig,
Ficus mallocarpa, which in its youth twines about smaller
trees, eventually enveloping them by the rapid growth of
its knarled trunk and literally smothering them to death.
The pillar-tree, Wehea africana, with its smooth white bark
and mast-like trunk, is a conspicuous forest tree. A less
common tree is the shaggy-barked Walbergia ugandensis.
The large, glossy-leaved, magnolia-like tree, Conophryngia,
is another wide-spread species in the forest. One of the
large trees is the buttressed-trunked Chrysophyllum, which
towers above the other trees and bears only a small crown
of leaves at its apex. In certain parts of the forest the
small, straggling Trichoclaudus malosanus trees grow in
great abundance, especially in juniper forest. The com-
monest forest undergrowth is the prickly-leaved Acanthus
arboreus. ‘The bramble or blackberry, Rubus, is a common
shrub in the forest, where it grows in thickets in the lower

FLORA OF EAST AND MIDDLE AFRICA = 49

aititudes. Several shrubs and small trees belonging to the
Rubiacee are also common undergrowths. Great vines,
several inches in diameter and many feet in length, form a
network in some of the highland forest of the Mau and in
parts of Kenia. The rubber-producing Landolphia vines
often form a considerable part of this network.

The upper forest at timber-line is marked by a dense
growth of bamboo of a peculiar high-mountain species,
Arundinaria alpina. The bamboo forest occupies a zone
about a thousand feet in vertical height, ranging from tim-
ber-line at ten thousand feet down to nine thousand feet.
In this belt it grows to the exclusion of all other trees, the
stems usually standing a foot or two apart and so close that
it is quite difficult to squeeze through the forest without
cutting a path. The only trees which are stragglers in
this zone are Podocarpus milanjiana and Olea laurifolia.
On the upper timber-line edge a large tree of the order
Rosacee appears, Hagenia anthelmintica, which is character-
istic. Practically no animal life is confined to the bamboo
zone and very few animals live within its borders, owing
to the soil supporting no undergrowth. The bamboo forms
a desert waste between the life of the forest proper and the
alpine regions. A narrow fringe straggles down into the
forest along streams to an altitude of seven thousand five
hundred feet.

One of the marked peculiarities of the African timber-
line is the absence of dwarfing, which is so characteristic a
feature of the tree-growth at high altitudes in our northern
climes. The forest-trees grow fewer in numbers at the upper
levels in equatorial Africa, but they are no smaller in size.
Much of the dwarfing in the north is due no doubt to the

50 AFRICAN GAME ANIMALS

pressure of heavy masses of snow which must be borne for
several months. Such a condition never occurs in tropical
Africa, where the snow-fields are constant in area and far
removed from the forests. Only the herbaceous moorland
vegetation reaches the snow-fields, but does not mix with
them.

The moorland which rises above the bamboo zone is a
well-marked treeless belt extending from ten thousand
feet to the snow at fourteen thousand five hundred feet.
It is divisible into two lesser floral zones: a lower tree-heath
belt ranging up to twelve thousand feet and a higher zone |
reaching to snow-line, characterized by the presence of the
tree Senecios. The vegetation in the moorland zone is
closely allied to that of the alpine regions of Eurasia and
North America. Many of the genera and a few of the
species which occur are common also to the north temperate
zone. The great bulk of such plants belong to the orders
Rosace@ and Ericacee. The dominant shrub or small tree
of the lower zone is Erica arborea, a species found also in
the alpine regions of southern Europe. Several species of
the wide-spread holarctic genus Alchemilla form the carpet
of the moorland. A. argyrophylla, a stout, shrubby species,
is the chief of these and grows in a dense mat one or two
feet in depth over the entire area of the tree-heath zone.
Another European genus, Hypericum, commonly known as
Saint-John’s-wort, occurs abundantly in the form of small
trees at the lower or timber-line edge of the heath zone.
Hypericum lanceolatum is an abundant tree, and its large
golden flowers, which are borne in great profusion, give the
lower alpine region a striking appearance. A small gray-
leaved tree, allied to our sage-brush, Stobe kilimandcharica,

FLORA OF EAST AND MIDDLE AFRICA 51

appears with the Hypericum and the tree heaths in groves
on the lower edge of the zone.

The upper half of the moorland zone is given a weird
appearance by the prevalence of forests of tree groundsels,
Senecio keniensis. This plant exhibits a striking variety
of form ranging from a globular mass of large radical leaves,
in appearance like a cabbage, in youth, through a thick-
stemmed, barrel-shaped plant a few feet in height, and
finally to a tall woody-stemmed, branching tree capped by
immense club-shaped heads of dense leaves. Straggling
forests of groundsel occur from twelve thousand feet to near
the snow-line, but at the upper level they gradually dwindle
in numbers and size, and those which reach the snow are
only a few small, radical-leaved forms. Groundsels of sev-
eral species are found throughout the world as common
roadside weeds, but nowhere except in the highlands of
Africa do they exhibit such large tree forms.

Associated with the groundsels are two species of lobelia,
Lobelia gregoriana of the middle altitudes and a tall, woolly
pubescent form, Lobelia telekii, which is abundant in the
tree groundsel forest and continues up to snow-line. Both
of these are peculiar in form and quite far removed from
the lobelias of our gardens.

Throughout this upper alpine zone the Alchemilla is
lacking, its place being taken by the heavy tussocks of a
grass-like rush, Ucinia, which covers the hummocky, boggy
soil to the snow-line, where it alone carries on the struggle
with the cold. Growing everywhere over the whole moor-
land are found the daisy-like white and pink flowers of
Heliochrysum in clumps and isolated patches.

The alpine vegetation of equatorial Africa is distinct

52 AFRICAN GAME ANIMALS

from that of the lower zones, and is closely allied to that
of the north temperate zone, from which source it has evi-
dently been derived. The heaths, hypericums, alchemillas,
junipers, and brambles are the chief groups of northern
derivation. It is possible that some of these reached East
Africa from the Mediterranean region by way of the ranges
flanking the Red Sea and the Abyssinian highlands. The
region, however, is widely isolated from such a source, being
without connecting mountain chains and having the great
barrier of the Sahara Desert between. These northern
species have, no doubt, reached central Africa at various
times as strays, wind-borne, or by migratory birds or oth-
erwise. It is scarcely conceivable that they could reach
the region by progress through the lowlands at some time
when the climate may have approximated that of southern
Europe. Moreover, we have no evidence of the climate
having been any colder to any considerable degree during
any past period in tropical Africa. A small element of the
alpine flora has ascended the mountains from the plains
below. The groundsels, lobelias, bamboos, and grasses are
examples of this sort. A strong argument of the accidental
acquiring of the present flora is the fact that not a single
northern type of mammal exists on the high mountains.
This is not likely to have been the case had northern plants
been able to reach the region by way of the lowlands. The
species of mammals which occur at high altitudes have all
been derived from the lower levels. None of these show
more than a slight amount of specialization, notwithstand-
ing the great climatic and environmental changes to which
they are now subject. It is evident that the lack of any
marked physical characteristics in the mammals of the

FLORA OF EAST AND MIDDLE AFRICA 53

alpine regions show the species to have been of quite recent
immigration into the region.

The affinities of the forest flora are with West Africa
and to a lesser degree with Abyssinia and South Africa.
The flora of the nyika is distinctly East African with
scarcely any West or South African types, and forms a part
of a wide-spread association from the Red Sea to the Zam-
besi. The tropical coast belt is similar in affinities to the
nyika.

CHAPTER IV

CONCEALING AND REVEALING COLORATION AND THEIR
RELATION TO NATURAL SELECTION

INEVITABLY any great scientific discovery contains,
when first promulgated, some error; and equally inevitably
some of the followers of any great leader of thought tend
to push or twist his doctrines into fantastic extremes, and
then to turn his thought into a system of fossilized rigidity.
This happened to Plato and Aristotle; and it has happened
to Darwin. The real work that Darwin did was in con-
vincing practically all men of trained intelligence that the
animal world, including man, has developed by evolution,
not by an infinity of special creations. The law of evolu-
tion is as fundamental to our understanding of life as the
law of gravitation to our understanding of the inanimate
universe. It is less widely accepted merely because its
discovery was more recent; the good people who on different
grounds still reject it are merely occupying the same quite
natural attitude that other good people of their stamp once
occupied toward the then newly promulgated doctrine of
gravitation. Less than a century ago the Catholic Church
kept on the index all books that admitted that the earth
went round the sun, and orthodox Protestant clergymen
treated as heretical any doubt whether the world had been
made in six ordinary days and was more than seven thou-

sand years old.
BA

COLORATION 55

But full recognition of the greatness of Darwin’s achieve-
ment does not mean our acceptance of the immense impor-
tance which he ascribed to natural selection in bringing about
evolution. Above all, it does not mean acquiescence in the
views of his professed followers on this matter. In England
and America, and largely in Germany, Darwinism has been
made into a fetich by most scientific men, with results
nearly as deadening to intellectual liberty as anything done
in the name of theology by the medieval schoolmen. In
consequence outsiders have had to step in to correct the
error of the professed scientists. The absurdity of the com-
placently arrogant materialism associated with the name of
Haeckel, the showing that this new type of dogmatic intol-
erance is as unscientific as it is degrading, was left to Bou-
troux and Bergson, philosophers so thoroughly acquainted
with physical science as to be able to perceive its limita-
tions. The curiously unscientific mysticism of the Weis-
mann school was also left to be exposed by men who were
not professional scientists.

In this book we are dealing only with a tiny corner of
the field which must be covered. But even in this tiny
corner, what is above said applies. One of the series of
phenomena to which especial prominence has been given
in bolstering the theory of natural selection is that con-
nected with the coloration (and certain details of the bodily
form) of animals. We propose to deal with this in so far
as it relates to the big game of Africa, using certain other
animal forms for purposes of comparison and explanation.

By Darwin natural selection was treated as the chief,
and by his more fanatical followers as the only, factor in
evolution. Most American and English scientific men have

56 AFRICAN GAME ANIMALS

no real acquaintance with the writings of men like Cope,
who with masterly ability set forth the claims of those who
believe in the inheritance of acquired characteristics as
likewise a considerable factor; it is only in France, where
the outlook is broader, that Cope’s eminence is recognized.
In America and England natural selection as a complete or
nearly complete explanation of evolution is still on the
whole the fashionable scientific doctrine. Yet even in
America and England there is now a wide-spread and con-
stantly growing belief that, although natural selection un-
doubtedly plays a great part in the survival or extinction
of species, it only plays a minor part in the creation of
species, and that none of the explanations hitherto ad-
vanced of the creation of species is by itself sufficient to
explain, and indeed that all of them together are not suffi-
cient to explain, the major part of the phenomena.

By far the most satisfactory recent book on the subject
is “The Making of Species,” by Dewar and Finn. The au-
thors keep ever in mind Huxley’s statement that “Science
commits suicide when it adopts a creed.” In other words,
they try not to pin their faith to dogmas and, above all, not
to accept one explanation as true merely because they have
no other to offer. Doubtless the greatest service to science
is rendered by the man who can give the right explanation
of phenomena. But the most serious injury is done by
the man who persuades people to adopt the wrong explana-
tion. In our present state of knowledge it is a prime
necessity that we shall be continually saying: “I do not
know.” As yet we know almost nothing as to how species
originate. The theories of closet specialists help us very
little. Experiments carried on under wholly artificial con-

COLORATION ~ 57

ditions, as on animals in captivity, help us more, but must
be most carefully scrutinized or they will lead us into
error—as has been well shown by McAtee in his paper on
“The experimental method of testing the efficiency of
warning and cryptic coloration in protecting animals from
their enemies,’ in the Proceedings of the Philadelphia
Academy of Natural Sciences. Further help, indispensably
necessary, must be rendered by first-hand observations of
species in their natural surroundings and in their natural
conditions. We propose to render a very little help in this
way, by recording observations, far from being as extensive
as they should be, but faithful and painstaking as far as
they go. We wish to give our actual field experiences and
observations on the coloration of the big-game animals of
Africa, comparing it with the coloration of various other
animals, especially the big-game animals of North America,
with which we are also familiar at first hand.

The neo-Darwinian ultra-upholders of the natural selec-
tion theory, ranging from Mr. Wallace through Professor
Poulton to Mr. Abbott Thayer, all explain almost all the
phenomena of animal coloration and of the development of
appendages on animals by the theory of natural selection.
They hold that almost all the details of an animal’s colora-
tion, etc., have a direct utilitarian purpose, which they
name, and a particular kind of usefulness, which they point
out; and they treat these as survival factors. But as re-
gards many cases they differ from one another, and indeed
contradict one another and take directly opposite views
concerning what this useful function is. Mr. Wallace, for
instance, believes that many striking markings of a highly
advertising quality serve as recognition marks between

58 AFRICAN GAME ANIMALS

individuals of the same species. Mr. Thayer considers
these same markings as obliterative, as concealing. Now,
of course, both these theories cannot be true; as a matter
of fact on this particular point, while we can say that the
theory of Mr. Wallace is not proved and is probably incor-
rect, but is possibly true, we are obliged to say of Mr.
Thayer that his theory is obviously without any founda-
tion in fact and ought not to need serious discussion. We
mention it at all merely because scientific men of reputa-
tion, like Professor Poulton, and scientific journals of stand-
ing, like the 4uk, have given it solemn, and at times fa-
vorable, consideration. Mr. Wallace’s view that the white
tail of a running rabbit serves as a warning or directive
mark must be seriously treated; but Mr. Thayer’s theory
that this white tail hides the running rabbit from a fox,
snake, or weasel ought not to need any treatment whatever,
serious or the reverse.

Mr. Wallace’s theory is that concealment is the general
and ordinary purpose served by the coloration of animals
—of African antelope, for example—but that their face and
rump markings, and in the case of African antelopes their
horns, serve as recognition marks, to enable the individuals
of each species to recognize one another, and above all to
enable those of opposite sex to be sure that they mate
within the species. We think that the facts observed in
the field do not bear out these theories. Mr. Wallace gives
pictures of the horns of twelve species of African antelope
to show that they serve as recognition marks. In nine of
the twelve species—kob, waterbuck, lechwi, gazelle, etc.—
these horns are confined to the male, and assume their char-
acteristic shape only when he is adult. It seems a very

COLORATION 59

wild theory indeed to suppose that the old males need
recognition marks, while the female and young males do
not; and not one fact has been advanced in support of the
theory. All the facts are the other way. A waterbuck or
kob will recognize a hornless cow or a horned bull of the
same species with equal facility, and at a distance such
that the horns cannot be distinctly seen. A tenth species
mentioned by Mr. Wallace, the hartebeest, has horns in
both sexes; but they are relatively short, whereas this
particular antelope is of extraordinary shape and can be
recognized by its shape alone at such a distance that the
horns cannot be made out at all. It does not seem wise to
advance such a theory as this concerning antelope horns as
recognition marks when no observed fact supports it and
many observed facts are inconsistent with it.

As regards the conspicuous face and rump markings there
is room for argument. We do not, however, believe that
these markings serve any purpose of recognition for sexual
purposes. Cases of melanism and albinism are well known
in many animals, often appearing only in certain localities.
White deer and white waterbuck are not infrequent. In
every case the animal seems to associate with others of its
kind precisely as if it were normally colored; and if a female
it will frequently be found with a young one, usually of the
ordinary color. Black and brown bear cubs, black-maned
and light-maned lion cubs are found in the same litter.
Where such startling color patterns as are implied in being
white all over or black all over totally fail to confuse indi-
viduals of the opposite sex as to what species the animal
belongs to, it seems unlikely that they require guidance in
such matters by the display of a few spots of white or black

60 AFRICAN GAME ANIMALS

or a slightly different tinge to the coat. Terns which have
been dyed indigo or carmine and turned loose are soon
accepted by their fellows, and even by their mates, on the
old basis. Mr. Spencer Trotter found a male scarlet tan-
ager breeding although the hind back and the belly were
green; but the startling aberration in color had not con-
fused its mate. In one herd of topi we saw an individual
with a blazed or white face, like that of its relative, the
South African blesbok; but the other topi treated it pre-
cisely like one of themselves. Animals with dichromatic
color phases, like the red and gray phases of the American
screech-owl, and of the cougars we have collected in the
Colorado and Arizona mountains, mate together wholly
without regard to color. Still more striking are the facts
in the case of the white-withered lechwi and the white-
eared kob. These animals belong to very closely allied
genera and are of about the same size, and they dwell in
the same neighborhood along the White Nile—one in the
swamps, the other in the dry plains alongside the swamps.
Sometimes these herds meet on the edges of the swamps.
The females are so alike in shape and color that it needs
an inspection close up, including a look at their hoofs, to
tell them apart. If animals really needed striking recogni-
tion marks to enable the individuals of one sex to recognize
the individuals of the other sex of the same species, there
could not fail to be numerous hybrids between the kob and
lechwi. As a matter of fact, although both species are
abundant in the same region, such hybrids never occur, or
occur so rarely that they have never been observed. It is
evident that the sexes have no difficulty whatever in recog-
nizing one another and discriminating against the animals

COLORATION 61

of the opposite sex belonging to the other closely allied
species.

All these facts are fairly conclusive that there is nothing
in the theory so far as sexual recognition is concerned. That
there is also very little in the theory as regards ordinary
or non-sexual recognition, we have no doubt. A roan ante-
lope, for instance, has bold face markings—an eland, in some
of its forms, none; but there is evidently no more difficulty .
about recognition in one species than in the other. Dewar
and Finn believe that the rabbit’s white tail is of no use, and
advance substantial reasons why it cannot serve as a warning
to other rabbits. Nevertheless, to us it does seem as if a con-
spicuously exhibited white tail must serve some functional
purpose of advertisement where there is the kind of display
that we see in the various hares, the white-tailed deer, and
the antelope squirrel. Mr. Seton produces facts to show
that it may serve as a guide to the young. The prong-
buck of America does, when excited or curious, raise the
hair on the white rump with a chrysanthemum effect, so
that it flashes in the sunlight; and it is possible that this
may enable the pronghorns to recognize one another at a
distance of a mile or two—although, as their bulging, tele-
scopic eyes are better than those of any other American
game, they can probably recognize one another’s ordinary
markings by vision at such a long distance that the posses-
sion of another method of recognition seems a matter of
surplusage. Moreover, Mr. Nelson’s exceedingly inter-
esting study of one species of southern jack-rabbit seems to
prove beyond question that, for whatever purpose, the
animal does deliberately make use of directive coloration.
This jack-rabbit is concealingly colored when crouched

62 AFRICAN GAME ANIMALS

motionless and trusting to concealment. When it runs it
hitches its skin so as to bring the white of the belly up on
the side toward its disturber—a most remarkable example
of seemingly purposeful advertising coloration.

So much for recognition marks. But by the ultra nat-
ural-selection men the chief stress is always laid on conceal-
ing coloration. Instead of treating concealing coloration
as something which is sometimes present and sometimes
absent, which is sometimes useful and sometimes not—in
other words, instead of treating it from the scientific stand-
point of the observer who prefers rather unsatisfactory and
partial truths to brilliant and comprehensive falsehoods—
men like Professor Poulton and, above all, Mr. Thayer treat
the extreme neo-Darwinian theory of protective or con-
cealing coloration as a law of nature. It is nothing of the
sort. It is at best, as Dewar and Finn point out, a work-
ing hypothesis which is useful as throwing light on certain
phenomena, and which does apply in a large number of
cases. It breaks down if the attempt is made to apply it
universally or nearly universally, and is demonstrably false
as applied to all sexes and ages of many thousands of species
of the higher vertebrates, mammals, birds, and even reptiles
(to speak only of species which we have personally studied).
Professor Poulton has conducted a number of experiments
under highly artificial conditions, and has drawn certain
conclusions, partly from these experiments, but much more
largely from what can hardly be called anything except
closet theorizing. Mr. McAtee, in the paper quoted above,
has shown why these particular experiments and conclu-
sions have small value; this being done as an incident to
showing why most experiments under artificial conditions
have small value.

COLORATION 63

The best examples of experiments carried on under con-
ditions so artificial as to make the experiments not merely
useless but misleading, and of conclusions which are of
even less value than the experiments, are furnished by Mr.
Thayer. Mr. Thayer’s experiments have the value that
attaches to puzzle-pictures in the Sunday papers. They
show that by the exercise of ingenuity it is possible to find
for every mammal or bird, of every conceivable type of
coloration, some combination of light, background, and sur-
roundings in which it will be inconspicuous. Mr. Thayer
has recorded the results of his observations in a beautifully
printed and illustrated book, containing a great many
photographs of live wild birds and mammals taken by other
men, some photographs of dead or tame animals or of arti-
ficial imitations of wild animals taken by himself, and some
pictures painted by himself. The latter are designed to
show that peacocks are concealed from their enemies be-
cause they are colored like the sky and the foliage, wood-
ducks because they are colored like water-lily flowers and
leaves, blue jays because they are colored like shadows on
snow (most blue jays spend most or all of the year where
there is no snow, and against the snow their coloring is
revealing hundreds of times for once that it is concealing),
and spoonbills and flamingoes because they are colored like
clouds and sunsets—it being supposed that sharks and
crocodiles, and the minute gasteropods on which the fla-
mingoes feed, are misled into believing that the sun is always
setting whenever flamingoes are near. It is a little difficult
to develop the theories for which Professor Poulton stands
to a higher point than- Professor Poulton has developed
them; but Mr. Thayer has accomplished the feat. He

64 AFRICAN GAME ANIMALS

upholds the views that the coloration on practically all
animals (including even the most vivid and advertising
colors and patterns) is really concealing—not that it was
concealing in some remote geologic past, but that it is
actually concealing to-day; and he attributes this to “nat-
ural selection, pure, simple, and omnipotent.” He shows
both daring and versatility in his use of facts. Skunks,
crows, scarlet tanagers, white herons, and the like he cou-
rageously declares to be concealingly colored. Nor is he
daunted when his theory requires him to show that the
same sky is of two diametrically opposite hues, so that it
may, under the same conditions, harmonize both with the
deep blue of a peacock’s neck and the dazzling white of a
prongbuck’s rump. He wishes to show that each object
fades into the sky; so he paints in colors the picture with
the peacock in it, making its neck and the sky of the same
brilliant azure tint; and he leaves uncolored the picture with
the prongbuck in it, so that in this case the sky and the
rump are both white. What would happen if the two ani-
mals changed places and wandered each into the other’s
sky, he does not explain. Professor Poulton, presumably
speaking for and to the leading scientists of Great Britain,
highly praised the scientific worth of this book. It seems a
pity that he could not have included in his praise the equally
striking contention of the late Brother Jasper that “the
sun do move” round the earth.

Professor Poulton’s theory, developed both by him and
by Mr. Thayer, is that the eyes of a creature’s prey or
adversary take into account the most minute details of
coloration. This may be so in the case of such an animal
as a tree-toad. It is certainly not so in most cases, how-

COLORATION 65

ever. With multitudes of animals no notice is taken of any
prey unless it moves, and when it moves all small detail is
lost. A lizard will pay no heed to the most brilliantly col-
ored insect when motionless and will seize anything that
moves, even when itself at the moment in jeopardy. I once
watched a lizard poised motionless by a large motionless
insect on a small tree; I moved my hand to capture the
lizard, which caused both the latter and the insect to shift
positions; whereupon like lightning the lizard seized the
insect, and with its mouth distended by its prey it dodged
round the tree trunk to avoid its foe. Any angler could
enlighten doubters as to the fact that no resemblance in
minute detail is needed in order to get fish to take a fly.
From time to time fly-fishers experiment with artificial flies
showing this minute resemblance, only to return to the
regulation flies, which certainly do not minutely resemble
any known insect. A trout which is eager after a “red
ibis” or “‘brown hackle” must possess a splendid visual
disregard of minute detail.

What is true of wild fish in a state of nature is also true
of wild birds in a state of nature, as is proved by the experi-
ences of shore-shooters with decoys, both as regards water-
fowl and as regards bay-birds or shore-snipe. The ultra
protective-colorationists lay stress on the great importance
of the “ripple marks” on the flanks of ducks, and the exact
tint of the legs of waders, and other minute details, from
the standpoint of the creatures that look at them. Well,
the conduct of water-fowl and shore-birds and of birds of
prey toward all decoys shows that these minute details are of
no consequence whatever until the birds are so close that
they could by no possibility have either any concealing or

66 AFRICAN GAME ANIMALS

any revealing value, and that even then they may be com-
pletely ignored. By actual experience shore-shooters have
found that after a certain rough resemblance to the wild bird
has been attained further painting of the decoy to make the
resemblance minutely accurate is a waste of time and brings
no advantage. Instead of legs the shore-bird decoys stand
on sticks which are thrust into the sand. The duck decoys
which bring best results are commonly painted so roughly
that none of them would for a moment be accepted in even
the cheapest book or collection as fit copies of the originals.
Yet the species of water-fowl and shore-snipe which decoy
at all will come to these decoys as readily as to any others,
and often will light among them and swim or walk round
them for some minutes. For unknown reasons certain
ducks and waders will not come to decoys; but the species
that do come evidently fail to notice the minute details,
and some of them fail to notice any details, coming down
to decoys which are mere caricatures. Birds of prey some-
times strike at decoys. I have myself known an eagle to
stoop at decoy ducks, and when it failed with the first one
attacked, to repeat its swoop at another, evidently not
understanding what had happened. There could be no
better proof of the unimportance of detail in the eyes both
of at least some of the birds that prey and some of those
that are preyed on under the ordinary conditions of their
lives.

There are similar facts to be adduced regarding mam-
mals. In Africa, in former times, bushmen, customarily,
and at least one white man, Mr. Jackson, the present gov-
ernor of Uganda, on one occasion, rigged themselves as arti-
ficial ostriches and thereby were able to approach gazelles

~

COLORATION 67

and other animals closely, although, of course, their legs bore
no earthly resemblance to ostrich legs, and indeed the neck
and head were all that was really simulated. East African
natives sometimes equip a donkey with the scalp and horns
of a hartebeest and use it with success as a stalking-horse;
which goes to show that oryx, zebra, hartebeest, and the
like pay as little heed to detail as any ducks that ever were
tolled to the rudest wooden decoys. A hunter I knew in
Montana many years ago was once bringing home part of
the carcass of a deer on his back, when he was pounced on
and knocked down by a cougar. As soon as the cougar
realized that it had to do with a man, it drew back; evi-
dently it supposed it was attacking a deer; and in the stupid
boldness of hunger it paid so little heed to detail as to make
its assault in spite of the slow pace at which the man was
moving and the wholly unnatural position of the deer.

Of course, it is unwise to generalize in confident fashion
on a few data; but as far as they go these experiences with
fish, birds, and mammals tend to show that vertebrates pay
no such heed to minute details of color and shape as is
assumed by the ultra neo-Darwinians in supporting their
theories of the all-sufficiency of natural selection in origi-
nating species, and of the extraordinary and well-nigh uni-
versal potency of concealing coloration as a survival factor
as regards all beasts that prey or are preyed on. As for
insects, any one who has ever seen a hornet or wasp trying
to catch flies in a tent and continually pouncing on any
black spot must realize that their vision is so poor that they
cannot take in details of even an obvious kind.

Boutroux has dwelt on the need of using common sense
in testing the extravagances of scientists no less than the

68 AFRICAN GAME ANIMALS

extravagance of theologians. Really, there ought not to be
need of appealing to anything except common sense when
we are asked to consider some of the vagaries of the ultra
concealing-colorationists. One of their latest fads is that
white is a concealing color, even at night; that beasts with
white sterns, like so many ruminants and rodents, are con-
cealed from their foes at night by their white sterns because
white is invisible against the sky-line. War-ships in time
of war are most often seen against the sky-line, and are
always so seen by torpedo-boats, which it is the especial
object of the big ships to elude at night. Well, no naval
officer in his senses would dream of painting his ship white
so as to escape observation at night under any normal con-
ditions; as soon as war comes all ships are painted some
neutral tint, because as a matter of actual experience every
one recognizes the fact that, taking into effect moonless and
moonlit nights and all cloud effects and weather condi-
tions, white is on the average the most revealing or adver-
tising of all colors at night. In the old days, on the West-
ern ranches, we always preferred white horses for night
work whenever it was desired to be conspicuous. Scien-
tists ought not to show less common sense than naval men
and cow punchers.

Of course, every color and every conceivable combination
of colors may under some circumstances be concealing or
revealing, even although under ordinary conditions the re-
verse is true. It isa question-of averages. )) There arc
nights when no color can be seen, and then the color is a
negligible factor. There are other nights or portions of
nights when under some given conditions of background
or cloud effect or position of the moon, black or brown or

COLORATION 69

gray or white may each in turn be more conspicuous or
less conspicuous than any of the others. But half of
the hours of darkness are moonlit, and if the moon is
very bright, white, black, and the bold colors generally,
are almost as visible as by day. Taking into account
moonlit and moonless nights, starlit and starless nights,
clear and stormy nights, and all the different cloud effects,
and the probable backgrounds of sky, soil, and vegetation,
white is on the average by all odds the most conspicuous
color at night, and the neutral tints, the dull grays, drabs,
and yellow—browns are the least conspicuous when seen
from the ordinary view-point of the ordinary foes. The
white rump of a standing or running prongbuck or black-
tail deer or bighorn is under most conditions of a highly
revealing or advertising value at night as well as by day;
and the same is true of a running white-tail deer or rabbit.
We state this as the result of innumerable personal observa-
tions of these animals when in their own homes under nor-
mal conditions.

Two of Mr. Thayer’s ‘‘proofs by experiment” may be
mentioned to illustrate the worthlessness of experiments
when carried on under the influence of a perverse obsession.
Mr. Thayer’s thesis is that a// forms of coloration are con-
cealing; but next to countershading he considers “ruptive”’
and “secant” coloration schemes as most potently con-
cealing. He regards vertical striping as a type of concealing
coloration of peculiar potency, brought about purely by the
“omnipotent” power of natural selection, because of its
survival value. He adduces the striped African antelopes
(all of them tragelaphs, except one duiker) as cases in point.
He records one experiment which he asserts proves the truth

70 AFRICAN GAME ANIMALS

of his assertion. Ata meeting of the American Ornitholo-
gists’ Union in Washington, as he relates, he showed to forty
naturalists ‘‘a small, stuffed deer (a South American species)
that wore from its dorsal line down its sides two white
stripes in imitation of those of certain African antelopes.”
He states that at ten yards’ distance these (artificial) stripes
rendered the deer invisible to the “forty naturalists,” but
that when they were removed and the deer left colored as
it actually was colored by nature it became clearly visible.
Apparently neither Mr. Thayer nor any one of the “forty
naturalists”’ was struck by the sufficiently obvious fact that,
according to his own account, he proved that the actual color-
ation of the deer was not concealing, and that it had to be con-
cealed by artificially giving it a coloration it did not 1n nature
possess. He showed, therefore, that nature did not give
the deer a concealing coloration, and that Mr. Thayer him-
self had to step in to supply the omission. In other words,
he proved that natural selection had not been “simple and
omnipotent,” and that concealing coloration was not a sur-
vival factor; that, if it played any part in protecting the
deer, it was a minor part as compared to other factors, such
as habit and cover. Now, only one in ten of the species
of African antelope has stripes; very few have spots.
No antelope outside of Africa is striped. None of the deer
have stripes, and very few, when adult, have spots. The
small ruminants, such as the one to which Mr. Thayer
thoughtfully gave the concealing coloration it lacked, in-
clude the South American brockets, the Asiatic muntjacs,
and other little deer, and antelopes like the duikers, dikdiks
and reedbucks. All of these are unstriped.

Mr. Thayer’s experiment, therefore, proves, so far as it

COLORATION 71

proves anything, either that stripes are not concealing, or
that if they are concealing then natural selection has only
been able to develop them in one-twentieth of the animals
to which they would have been of benefit. Nor is this all.
Of the groups of African antelopes which carry stripes, two,
the bushbucks and the elands, seem to be slowly losing
them. The centre of development of these two groups
was probably the African equatorial forest, from which
both have spread eastward and southward. The farther
from the forest they go the less evident the stripes become,
until in the South African eland the stripes completely
vanish, and in the males of the East and South African
bushbucks they practically vanish. Yet where conditions
happen to be identical the practically monochrome bushbuck
and eland prosper as well as their striped and spotted kins-
men. In the Lado we found the non-striped duiker even
harder to see than the harnessed bushbuck of the same
habitat. In the dry bush of East Africa the unstriped
gerenuk is as difficult to make out as the striped lesser
koodoo of the same habitat. Considering all these facts,
the conclusion seems inevitable that, on the whole, the
striped coats confer no advantage and are if anything
slowly dying out, although not in consequence of natural
selection. The fact probably is that as regards deer and
antelope habit, or the combination of habit and cover, is
the survival factor of such overwhelming importance that
the precise color of the coat is of no consequence one way
or the other. This is certainly the case at the present time,
and it was probably the case as regards these particular ani-
mals since a comparatively remote geologic past.

The coloration of most groups and genera of African

72 AFRICAN GAME ANIMALS

big game is very ancient and of long standing and is often
more constant than the osteological and horn characters
upon which the genera are usually based. Such fixity must
have been produced by color inheritance through countless
generations. Present environment, therefore, acts very
slowly, if at all, on the color scheme; its only appreciable
effect being usually one of tone or shade on the colora-
tion generally, and this effect being probably due to the
inheritance of acquired characteristics rather than to natu-
ral selection. One of the strongest proofs of the long-
standing nature of color pattern is found in the young of
such striped game as zebra, koodoo, and bongo, which
show absolutely down to every minute detail the color pat-
tern of their parents. There is in these early stages no clew
to the remote ancestral coloration of the race, nor is there
moreover in the earliest foetal stages in which color is as-
sumed any change from the adult livery in essential pat-
tern. But the young of the unstriped eland and unspotted
and unstriped bushbuck do show traces of ancestral spots
and stripes.

The other “experiment” of Mr. Thayer’s to which I
wish to refer is that made with imitation oryx heads and
zebras. He wishes to show that its stripes protect the
zebra when it is among the reeds at a drinking-place; and
that the bold black and white markings on the head of
the oryx likewise “simulate reeds” at a drinking-place.
He contrasts each picture with a picture of the same ani-
mal, or animal’s head, colored gray or brown and counter-
shaded in ordinary fashion. He points out with triumph
that (in his pictures of artificial animals) the whole zebra
in one case and the oryx head in the other are more difh-

COLORATION 73

cult to see than the monocolor countershaded animal and
animal’s head; he announces that this proves that the stripes
on the zebra, and the bold black and white markings on the
oryx head, are striking instances of concealing coloration—
developed by natural selection to insure the safety of the
animals from beasts of prey when they go to drink.

This “experiment” has an interest of its own, it is true,
but only as reflecting light on the worth of all such experi-
ments. Mr. Thayer knows nothing whatever of the habits
of African game or he would not make such a contention.
Zebra and oryx are beasts of the open country. Neither |
ever goes into reeds except when approaching a drinking-
place, and then rarely. Their habits, and the habits of
other plains game—hartebeests, wildebeests, eland, gazelle,
topi, kob, even impalla—are such that it is absolutely im-
possible for any hunter, beast or man, to avoid seeing them
when they come to water. The animals with striped bodies
and the animals with gray or brown countershaded bodies,
the animals with boldly marked heads and those with nearly
monochrome countershaded heads, all live under the same
conditions, all have precisely the same habits on approach-
ing drinking-places, and at such times none of them are
either helped or hurt by their coloration patterns.

In fact, this particular proposition is merely advanced
as a phase of the theory that the coloration of the various
animals of the open plains, which is obviously of no use in
concealing them while they are feeding or resting, may be
of value in concealing them when they come to drink.
There are no facts advanced to support this theory; and,
on the facts actually observed, there are two conclusive
objections.

74 AFRICAN GAME ANIMALS

The first is that such animals as the wildebeest, topi,
hartebeest, and zebra, which have exactly the same habits
and drink at exactly the same pools, are so differently col-
ored that it is possible to maintain the thesis of concealing
coloration as to one only by insisting that the others are
advertisingly colored; and yet all prosper equally well, and
obviously no one of them, as compared to the others, is in
any way either helped or harmed by its utterly different
coloration, or has altered its life to take advantage of it.
When the dark, nearly monocolored, countershaded harte-
beests, and the nearly monocolored, inversely counter-
shaded topis, and the striped zebras come to water (or
feed or rest) together, or in similar places, it is, of course,
evident either that the coloration of one advertises it and
puts it at a disadvantage as compared with the others, or
else, what is undoubtedly the fact, that neither the so-called
advertising nor the so-called concealing coloration has any
effect one way or the other.

The second and still more conclusive objection to the
theory is that the actual habits of the animals when they
come to drink are such as to render it impossible that their
coloration can have any, even the slightest, effect in con-
cealing them at such times. Motion, unless of an extremely
stealthy and skulking type, at once reveals any animal, with-
out regard to its coloration. We have seen white-tail, and
more rarely black-tail, deer skulk with such stealth as to
tend to escape observation; and in Africa it is possible (we
say possible, not probable) that some of the cover-loving
game—bushbuck, duiker, and dikdik, for instance—may
skulk and take advantage of cover, and even perhaps profit
by their coloration, as they come to drink; of this we can-

COLORATION (6

not speak. But we have carefully observed the habits of
zebra, eland, oryx, wildebeest, hartebeest, topi, kob, impalla,
big gazelle, and little gazelle. All these animals come to
waterin thesame manner. They not only make no attempt
whatever to hide while approaching the water, drinking, or
leaving it, but their methods of approach are such as to make
it impossible for them to escape the observation of the dull-
est beast of prey that has eyes at all. They, if anything,
avoid cover. They advance in the open. They come string-
ing toward the pools or streams in long files, occasionally
bunching together.

Where there is no cover at the drinking-places, they may
come to them alone or in small parties. Where there is
cover, or where there is much fear of lions, the different
species of a given district may come singly or in small
parties, or they may join and come to the drinking-place
in a throng, as has been described by Sir Samuel Baker, in
his “Nile Tributaries,” and by Doctor A. Berger, in “ Afrikas
Wildkammern.”’ In either event, the animals neither hide
nor seek to take advantage of cover during the approach.
They walk forward, halt to look around, and then walk for-
ward again. They are always on the watch. Occasionally
they wheel and gallop off, either in mere panic or perhaps (as
has been suggested by Mr. Hodson, in “The Great Thirst,”
as to roan and sable antelopes) to lure any foe to show him-
self. Their whole concern is themselves to detect the pres-
ence of any lurking enemy; they make no effort to escape the
observation of this lurking enemy. If they are very thirsty,
and yet suspicious of the lion’s presence, they will often
hover about the neighborhood for many minutes at a dis-
tance from the pool. At last they may charge madly down

76 AFRICAN GAME ANIMALS

in a body to drink, gulp a few mouthfuls and retreat at full
speed before their thirst is satisfied, returning again in a
few minutes. Colonel Patterson has described oryx per-
forming this manceuvre, and also once saw it performed by
a mixed herd of zebras and hartebeests. Again and again
we have seen hartebeests and zebra come to water. They
always travelled in the open for at least part of the time,
and halted and stood at intervals as they approached the
pool or stream; sometimes they would suddenly wheel and
gallop off for fifty or a hundred yards; they were very no-
ticeable as they walked down to drink and while drinking;
and they then at once jumped out of the stream bed and
cantered away to some open spot before beginning to graze.

Never once did we see a zebra, oryx, hartebeest, or any
other game animal of the plains approach or stay at water
save under circumstances that made it impossible for any
watching wild beast not to see it. The zebra or antelope
trusted for its safety exclusively to its sharp senses, watch-
fulness, and agility; its coloration was a negligible factor
one way or the other. We have seen game hang about in
the neighborhood of the water for an hour, shifting position
and watching before venturing to drink. Game never re-
mains in the cover near the pool for any time after drinking.
It is well to look at Lord Delamere’s photos of oryx going
to drink, in Lydekker’s “African Game”’; Dugmore’s photos
of hartebeest and waterbuck; Akeley’s photos of impalla;
Schilling’s photos of zebra and gazelle; Rainey’s photos of
baboons, rhino, gazelle, oryx, zebra, and giraffe, some-
times all drinking together. A brief examination of these
photographs shows that when the big plains animals ap-
proach, or are at, the drinking-places no coloration has or

COLORATION (i

can possibly have any effect in concealing them from their
foes. While drinking they are apt to start and look round.
They never lie down near the water; they never stand
motionless in cover near the water save for a few moments
between their forward moves; when they have satisfied
their thirst they move back to the open flats on which they
dwell; and the long tails of the zebra and oryx are perpetu-
ally in motion and would of themselves catch the eye. In
short, the habits of the zebra and oryx are such that at the
only times when they are in the reeds it is an absolute
impossibility for any watching beast of prey to fail to dis-
cover them, and no coloration, whether advertising or con-
cealing, would either help or hurt them. This applies by
day; it also applies by night save when the darkness is
such that no coloration can be seen at all.

But this is not all. Mr. Thayer “proves” in his “ex-
periment” that relatively to the variegated oryx and zebra
a practically unicolored countershaded animal would in the
reeds be revealingly colored. Well, it happens that in
Africa there is an animal, about as large as the zebra or
oryx, which nearly fulfils Mr. Thayer’s definition, which is
of a nearly uniform countershaded color on head and body,
and which does often live in the reeds. This is the water-
- buck. It often spends over half its time in the reeds and
near the water’s edge, standing and lying down. Mr.
Thayer “proves”’ that the zebra and oryx have a coloration
which in the reeds is far more concealing than is the color-
ation of the waterbuck when in the reeds. Yet, as a matter
of fact, the waterbuck often dwells in the reeds and the
zebra and the oryx never do. The actual facts are directly
the reverse of what Mr. Thayer supposes them to be. Not

78 AFRICAN GAME ANIMALS

only is his elaborate theory (based purely on artificial ex-
periment) unsupported by one shred of evidence, but it is
shown to be the direct reverse of the truth by the simple
expedient of studying the facts in the field. Yet it is on
just such theories as this, backed by just such experiments,
that the ultraselectionists rely for support concerning much
of their contention about concealing coloration. If Mr.
Thayer in this instance were sound in his premises he would
have proved the exact reverse of what he announced he has
proved; for he would have proved that natural selection
had developed a revealing coloration in the animal which
lives in the reeds. As a matter of fact, what is really proved
is that the coloration of these animals has no survival value
one way or the other.

Experiments of such an artificial character are usually of
doubtful value; carried on as Mr. Thayer has carried them
on they serve merely to darken understanding. No closet
or laboratory experiment begins to approach in value first-
hand field observations by a competent man. Mr. Charles
Sheldon’s preliminary study of the white and black moun-
tain-sheep, in his “ Wilderness of the Upper Yukon,” affords
a model of what such a study should be, and, when read,
there is no need further to explain why no amount of in-
genious closet guessing can take the place of trained first-
hand observation of animals in their native haunts under-
taken, not to twist facts into the support of a theory, but
with the ability and purpose to find out the truth. His
conclusion concerning the subject with which we are dealing
is that the colors and coloration patterns of these northern
sheep are not caused by natural selection of their protective
values—that they are not survival factors. On the whole,

COLORATION 79

it appears that in both the black and the white species they
represent revealing rather than concealing coloration.

But we must not forget how many men are unable to
observe clearly and understandingly, even when their inten-
tions are excellent. This is true even of savages whose life
is passed in following game, and of white men who make
hunting a business. Among savages, whose very lives de-
pend upon their knowledge of the game, it is extraordinary
to realize the extent to which this real knowledge is mixed
up with not only inaccurate personal observation, but in-
accurate inherited traditions, and the love of pure magic
and mystery. Hunting savages always talk much about
the game round the camp-fires; and the wildest legends are
mixed with acute and accurate observations, all told with
the same implicit belief. Our own Indians always took
this attitude toward deer, antelope, moose, elk, but pre-
eminently toward the buffalo, bear, cougar, and coyote,
animals concerning which the average Indian hunter was
wholly unable to divorce actual observation from mythology.
The same thing is true in Africa even of such acute observers
as the forest-dwelling Ndorobo, while the ordinary savage,
even of the hunting tribes, will believe utter fables, and even
persuade himself that he has witnessed such impossibilities
that his statements must be accepted with extreme caution.
As for the ordinary white hunter, let any one read what
Selous tells of the worthlessness of the views of even
experienced Boer hunters as to such elementary points as
the number of species of rhinoceros and lion, and the dif-
ference in ferocity among these (purely imaginary) species.
We ourselves have had precisely the same experience with
Rocky Mountain hunters in getting them to tell about

<

80 AFRICAN GAME ANIMALS -

grizzly bears and cougars; good, competent men would tell
us about cougars eleven feet long, and about the wide dif-
ference in behavior and ferocity between “blue” and “‘red”’
cougars (merely slight color phases of the same species, as
with red and gray screech-owls) and the similar differences
between ‘‘bald-face,” “roach-back,” and “‘silver-tip”’ griz-
zlies—all of them the same animals. Ina really good book
on the tracks of American game, by a veteran professional
hunter, there is an account of a purely mythical animal, the
“fan-tail deer,’’ of Montana and other parts of the West;
and again and again old hunters insist upon the existence of
the “‘ibex’’ which they have themselves seen and even
killed; yet the smallest scientific inquiry at once and in-
variably proves the fan-tail to be nothing but some small
white-tail, and the ibex to be a young mountain ram.

We need not wonder that unread men with untrained
minds take such positions; for educated men, good observers,
frequently record observations as diametrically opposed to
one another, and draw from one set or the other false con-
clusions. We are not in the least blaming these men;
doubtless we would do just the same ourselves if we had
not been put on our guard. We are all of us feeling our
way and must profit by both the discoveries and the mis-
takes of others. Too often the open-air naturalist, the
out-of-doors observer, backs up his erring closet brother
by observations which really represent nothing but the
tendency to see what the man has been told to look for.
The hunters and explorers unconsciously make observa-
tions in accord with what they have been taught to expect
to find in such a matter as concealing coloration. At one
time they are told that a general uniform tint is concealing,

COLORATION 81

and they see things accordingly. At another time they are
taught the extreme efficacy of stripes, and accordingly any
time a striped animal proves hard to see, the fact is stored
away in support of the theory.

In Sclater and Thomas’s “ Book of Antelopes,’’ Lieuten-
ant-Colonel Olivier says of the lesser koodoos of the Somali-
land region that they are found in “thick, scrubby jungle,”
and he dwells on the way in which the stripes on the slaty
gray of their coats make them “‘harmonize with the foliage”’
and “exactly reproduce the checkered shade of sunshine
through leaves.”” Writing of the gerenuk of the same re-
gions, Donaldson Smith says: “‘Their solid reddish brown
color renders them almost invisible . . . they almost always
keep to the bushes.” The two observations seem incon-
sistent; but they are not; it is only the implied explanations
that are inconsistent. A solid reddish-brown, and a slaty
gray with white stripes, are not in the least alike; yet the
wearers of both coats, in substantially the same country, are
exceedingly and equally difficult to see—as we ourselves dis-
covered on the Guaso Nyiro, where both species are found
under substantially the same conditions. The fact is that
it is neither the stripes nor the absence of stripes, neither
the reddish-brown nor the slaty gray, that is concealing.
Any color, or combination of colors, any solid-colored coat
or striped or spotted or varied coat—save, perhaps, some
highly advertising color like pure black or pure white, or
such a combination as a black back and a white belly—is
concealing im such cover to an animal with such habits.
Cover and habit are the essentials; the exact shade of, or
amount of variety to, the coat is of no consequence. Nat-
ural selection cannot have produced either coat under pres-

82 AFRICAN GAME ANIMALS

ent conditions; the utmost it can have done is to set wide
limits, between which any color or combination of colors
may be developed. Under bright sunlight in such bush
and jungle as that in which gerenuk and lesser koodoo are
found, separately or together, the play of light and shadow
is such, the checkered pattern of sunshine through leaves
on either solid or striped or spotted bodies is such, the bright
colors and deep shades of the accidented landscapes are
such, the vistas are so numerous, so superimposed on one
another, and of such varied length and breadth, and the
amount of interposed and surrounding and backing twiggery
and foliage is so great, that any motionless body of any
fairly neutral tint, whether solid or varied, is so difficult to
make out that the exact shade and the exact character of
the markings are of no consequence whatever. Gerenuk
or lesser koodoo, duiker or bushbuck, leopard or lioness—
all dwelling in the same surroundings, are equally well con-
cealed, whether their coats are striped or spotted or uniform.

It is curious to see how even first-class hunters, if unac-
customed to analyze their observations, have been com-
pletely misled by failing to understand this simple fact;
namely, that normally no animal is conspicuous in an or-
dinary landscape, for such a landscape is broken and acci-
dented, is flooded with light and varied with shadows, con-
tains innumerable vistas and obstructions, and is filled
with a myriad color tones. Naturally in such surround-
ings any animal of any color is difficult to make out;
and when he has recognized this fact, the average ob-
server tends to believe that it is the animal’s coloration,
and not the landscape, that is responsible.

One of the queerest claims of the concealing-coloration

COLORATION 83

theorists is that the giraffe is concealed by its coloration.
(In reality the giraffe is not concealed by anything.) The
advocates of the theory attribute this supposed invisibility
of the giraffe to its varied and mottled coat. They contrast
this “leaf and shadow pattern” with the elephant’s dark
monochrome, which they admit not to be concealing—in-
deed it would puzzle even them to imagine any purpose
which would be served by having elephants concealingly
colored prior to the advent of rifles. They quote Samuel
Baker as testifying to the concealing effect of the giraffe’s
coloration. But in his “Eight Years in Ceylon” Baker says
exactly the same thing about the elephant! All that he
really meant was that in the light and heavy jungles of
India and Africa he had often found even elephants and
giraffes difficult to make out, just as buffaloes and rhi-
noceroses in thick jungle are difficult to make out, just as
the beautifully colored and striped bongo and dark mono-
colored forest hog are equally difficult to see.

It seems impossible to avoid the conclusion that in these
cases the different patterns of coloration have nothing to
do with concealing their wearers; the fact is simply that the
cover, the character of the landscape, the light, are such
that any animal is hard to see; and the coloration pattern
is of no effect one way or the other. Too much light, like
too little light, tends to make all coloration patterns look
alike, and, therefore, to deprive each one of any special
value. The giraffe spends most of its daylight hours in
bright sunlight, as does the zebra, for both live in dry, hot
climates. The glare of light in Africa and India is such
that all patterns of any intricacy or minuteness tend to
disappear under it. This is shown in the case of the Indian

84 AFRICAN GAME ANIMALS

tiger by Mr. Kearton’s interesting moving pictures of a
wild tiger; as the beast comes through the jungle its stripes
are very distinct, but the moment it steps into the sunlight
the stripes disappear as if by magic, and the animal appears
to be of a light monochrome. In Africa, in the same fashion,
a cheetah at a little distance looks like a lioness in color,
and a giraffe appears of a uniform neutral tint.

The stripes of the zebra disappear in like fashion. One
of the amusing contradictions of the ultra concealing-color-
ationists is afforded by Mr. Gregory, the author of “The
Great Rift Valley,’ who considered zebras to be concealingly
colored because (at a distance) he could zot see their stripes;
whereas later it became the fashion for the average writer
of the school to insist that they were concealingly colored
because he could see their stripes. The two best ob-
servers among African hunters of wide experience, Selous
and Stigand, dwell especially on the great conspicuousness
of the giraffe and zebra. As we ourselves have observed,
the zebra often shows either white or black at a distance,
according to the position of the sun. The general tint of the
ordinary giraffe at a distance did not seem to us advertising;
but where the pattern could be seen at all it had a slight
advertising value, and the color of the reticulated giraffe was
always advertising. But the size and shape of the giraffe
are such that, where it can be made out at all in the
landscape, it can be made out at too great a distance for
the pattern to be seen.

Mr. Thayer’s book is a valuable contribution so long as
attention is strictly confined to what it says of colors in
nature from the artist’s view-point. His elaboration of the
importance of countershading in concealing objects under

COLORATION 85

certain circumstances is very valuable indeed as a contri-
bution to coloristic law. But his application of it as being
the prime and well-nigh all-important factor in concealing
coloration, produced through the “omnipotence”’ of natu-
ral selection, is wholly untenable. Nor is there warrant
for his assertion that the brilliant white bellies of so many
animals are the result of natural selection working toward
concealing coloration. Most, although by no means all,
mammals and birds are countershaded, in the sense of being
of lighter color below than above. But this is also gener-
ally true of the most plentiful of all vegetable productions,
leaves, and of many fruits. There is no reason to suppose
that concealment has been the reason for the development
in one case more than in the other. A pure white belly
cannot aid in concealing an animal when that belly is itself
concealed. There are snakes with white or yellow or light-
colored bellies; but as these bellies are themselves con-
cealed they have no effect in concealing the wearer, and
cannot have been developed by selection or otherwise for
such a purpose. The same is true of the myriads of shrews
and meadow mice with legs so short that the bodies, or at
least the hairs on the bodies, practically touch the ground at
all times when the animal is at rest. In these cases the light
belly can have no effect whatever in concealing the ani-
mal; yet when the sides are lighter than the back the
belly is almost always lighter than the sides. If concealing
coloration was the object, and natural selection the agent,
the belly, which is not seen and the coloration of which is
of no concealing service, would not have been lighter than
the sides. When in motion a chipmunk or rabbit is visible
anyhow, and the glimpses occasionally obtained of the color-

86 AFRICAN GAME ANIMALS

ation of the belly in the cases where it is white are of no
consequence; when stationary the chipmunk or rabbit con-
ceals the white, and therefore the white cannot conceal
the animal. Evidently in these very numerous cases the
white color has been developed by some force wholly
diverse from selection working for concealment.

As this is true of multitudes of species, there seems
scant reason to assume that in other species the same type
of coloration 7s assumed through selection working for con-
cealment. Nor is this all. If countershading were pro-
duced by selection working for concealment, it would mani-
fest itself at that time in the animal’s life when it would
be of use; and when animals fail to possess it they would
-be at a disadvantage. Neither supposition is true. In the
case of beasts of prey, the prime use of countershading would
be when they are crouched for the final rush, or crouched
as they lie in wait or slink stealthily forward toward their
prey. But, as a matter of fact, the countershading largely
or entirely disappears when the animal crouches; the lion,
leopard, tiger, or puma when crouched loses all or almost
all of the effect of the countershading. This is just as true
of the beasts that are preyed on. Deer, hares, mice, all
squat when they wish to escape observation; and all or
almost all of the countershading disappears as soon as the
animal crouches flat to the ground. Concealment, and
cryptic coloration as a help to concealment, are of most
value to young animals; the young are far more apt to try
to escape observation than are adults; they trust for safety
to hiding and remaining motionless, and they always crouch
flat, usually with head and neck extended, in such circum-
stances. In other words, the countershading loses practi-

COLORATION 87

cally all value at the very moment when in theory it ought
to be of most use.

Moreover, the so-called “countershading” in very many
animals is not concealing at all, but advertising. Very light
hues on the under part of a bird or mammal, even although
the upper part is dark brown or black, may technically be
called instances of countershading, but they certainly have
not the slightest concealing value. A black squirrel is well-
nigh as conspicuously colored as possible; but when he has
an orange belly or even a mere white belly the addition of
another vivid color does not tend to conceal him. We here
speak of our personal experiences in the woods of the South-
ern States, where the black and orange bellied or white bellied
fox-squirrels are always conspicuous in their native haunts;
and of our personal experience on Buckskin Mountain,
Arizona, where the big black squirrel with a white tail is
of diurnal habits and is always conspicuous—its coloration
being of a highly advertising value. There are very ex-
ceptional circumstances under which the white belly of a
white-tail deer or a white-footed mouse or a weasel may be
concealing; but they are so exceptional that they can be
disregarded; ordinarily, when it can be seen at all (which
on the weasel is almost never, and as regards the mice
very rarely) it has a slight advertising value. When one
can see the white of an impalla or sable it is usually adver-
tising. Herons with white bellies are no more concealed
than those of solid colors.

Finally, there are many animals with no countershading,
living under conditions precisely like those surrounding the
countershaded kinds; and yet no effect one way or the
other is produced by this difference. The weasel family, a

88 AFRICAN GAME ANIMALS

family of finished carnivores, is rich in examples of this.
The mink is not countershaded, the weasel has white under-
parts, the black-footed ferret, thanks to its dark legs, seems
inversely countershaded; but the different color patterns
seem in no way to affect their success in life. The mink is
under natural conditions abundant and wide-spread; it for-
ages on land as successfully as the weasel. Natural selec-
tion has never produced countershading in its coat. The
wolverene is inversely countershaded. The fisher and sable
are not countershaded. All these wide-spread, and under
natural conditions abundant, carnivores are not counter-
shaded and are not concealingly colored; ferocity, hardihood,
keen senses, and (except in the wolverene) agility and power
to take advantage of cover are all so developed that the
quality of the coloration is evidently negligible as affecting
either their prey or the not very numerous animals that
prey on them.

In the deer family there are many of the, under natural
conditions, wide-spread and abundant forms, such as the
moose, wapiti, and sambur, which are not countershaded
or are inversely countershaded; the moose is so black that
it is always conspicuous; the others are no more conspicu-
ous, except for their size, than their countershaded or
spotted neighbors. The wapiti will crouch and hide, as
we have personally seen, precisely like black-tail and
white-tail deer or like rabbits. The wapiti calf is slightly
countershaded and spotted. It therefore appears that the
non-countershaded, solid-colored wapiti has been evolved
from a spotted, countershaded form; and yet the wapiti
still seeks safety in concealment. This means either
that “natural selection” has eliminated the spots and

COLORATION 89

countershading as non-concealing (although wapiti still
dwell much in the forest, so that there has been apparently
no change in the character of the surroundings), or, far
more probably, that the coloration is of no survival conse-
quence. The sambur, according to Doctor Hornaday, is
very difficult to see in its own haunts. Evidently these
solid-colored, non-countershaded creatures are just as con-
cealingly colored as their countershaded or spotted and
varied kinfolk. The coloration is evidently of no conse-
quence so far as their foes are concerned; habit and cover
are the vital factor in avoiding their enemies.

The ordinary red fox, so widely distributed in the Hol-
arctic realm, offers a case where several forms, one counter-
shaded, another solid-colored, another inversely counter-
shaded, all exist in the same species. The black fox is
rare; but the specimens are as fat and in as good condi-
tion as those of other colors, showing that they are as
successful in life. The cross-fox is in places the dominant
form, the development of an inverse countershading having
had no detrimental effect on its welfare. The red fox is in
all its forms very conspicuous, its coloration being reveal-
ing and not concealing. In none of the forms of this species
has natural selection had any effect in securing a conceal-
ing coloration for the highly predatory, much persecuted,
and very successful animal.

Fully as significant is the case of the topi. This animal
is really inversely countershaded, the color being some-
what lightest on the back, while the shoulders and thighs
are very dark. This inverse countershading makes its
coloration more conspicuous than that of its cousin the
hartebeest. The third member of the group, the wilde-

90 AFRICAN GAME ANIMALS

beest, however, although countershaded, as is the harte-
beest, is so much darker than the topi that it is more
conspicuous. The three animals have substantially the
same habits. They never try to hide; they are always in
the open; when they approach a pool they are always in
motion. The countershading of one and the inverse counter-
shading of another make not the slightest difference; neither
does the lesser or greater degree of conspicuousness. Their
habits are such, and the habits of their foes are such, that
the particular quality of their coloration has no effect what-
ever of survival value.

All these instances tend to show that the non-counter-
shaded beasts are in no way hurt by the absence of coun-
tershading. In the great majority of cases countershading
is certainly not a factor of survival value. It is of service
only (if at all) when the difference in tint above from below
is small, and the general color in strict harmony with the
surroundings. If the color is not thus in harmony, the
countershading has no effect whatever. A wildebeest is
countershaded, a bear is not; but each can be seen, if on an
open plain, a mile or two off, because the dark color is so
conspicuous that the question of countershading is of no
consequence one way or another. Innumerable familiar
birds—grackles, cow-buntings, red-winged blackbirds, crows,
red-headed woodpeckers, etc.—bear witness to the same
facts

Rabbits when motionless are either crouched flat, or
else with the fore parts raised, but always with the hind
parts pressed against the ground. Seen from the side, there
is a slight, a very slight, countershading effect, at this, the
only time when the rabbit’s coloration conceals it at all;

COLORATION 91

but from behind there is no countershading effect whatever,
the part of the back which is pressed against the ground
being as dark as the rest. If natural selection had de-
veloped countershading as of survival value on the rabbit’s
fore quarters, where it can be of use only when the rabbit
sits partially raised on its forelegs, it would certainly have
developed it for the back and hind quarters, where it would
be of use both when the rabbit crouches flat and when it
sits with its fore quarters raised. But, as a matter of fact,
the countershading probably has no effect—certainly no
appreciable or survival effect—in concealing the rabbit,
which is just as difficult to make out when seen from the
rear as when seen from the side. Rabbits, like mice,
gophers, and the immense majority of small mammals are
hard to make out merely because their general tint is incon-
spicuous, wholly without regard to the countershading or
the details of the pattern, so that the animal escapes detec-
tion because it is colored so that it does not strike the eye,
and sits motionless like a clod or stump, amid such sur-
roundings that the eye overlooks it because of the great
abundance of the inanimate objects round about. Cover
and immobility primarily, and secondarily a dull color, with-
out regard to minute pattern or countershading, are the
factors in the animal’s concealment. On a perfectly bare,
flat, unbroken plain a rabbit or a gopher is seen at once, at
a long distance; even a mouse is visible under such condi-
tions, and, therefore, meadow mice are most reluctant to
venture beyond the protection of the grass, under the cover
of which they pass their lives.

The zebra has an unholy fascination for the entire school
of ultra selectionists, ultra concealing-colorationists—prob-

92 AFRICAN GAME ANIMALS

ably because it presents a problem the solution of which
needs such infinite (and perverse) ingenuity on their part.
Recently, a gentleman who observed a zebra in a zoo
standing motionless after nightfall has pointed out that it
gradually became invisible as the light faded. Of course it
did. From actual observation in the field we can testify
that on the average an eland or a roan becomes invisible
a little sooner as the light fades; and unquestionably this
would be true of the unstriped wild ass as compared with
the striped zebra. Selous has commented on the same fact
in comparing the coloration effects of zebra and oryx (or
eland) at night in their native haunts. All these animals
are countershaded. The topi, however, is not counter-
shaded. It becomes visible farther off than any of them,
and remains visible longer as the light grows dull. The
wildebeests and sables and the adult male white-withered
lechwis and white-eared kobs, because of their dark hues,
and in spite of being countershaded, are in their turn a little
more conspicuous than the topi under such circumstances.
The natural foes of all these animals hunt chiefly by scent,
although their great foe, the lion, has very keen sight; and
their habits, so far as escaping from their foes is concerned,
are substantially the same under like conditions, neither the
countershading nor the absence of countershading, neither
the conspicuousness nor inconspicuousness of the colora-
tion, seeming to have any effect in revealing or concealing
them or enabling them to escape from their foes.

As regards big game, which, when adult, have nothing to
fear from birds, and whose mammalian foes are no taller
than, or not so tall as, themselves, the beast of prey looks
on his quarry from about the same level or from a lower

COLORATION ier

level; that is, from practically the level of the hunter, who
in a stalk is usually crouching or creeping, so that he sees
the game with level glance at a distance, and looking up
when near by. But the immense majority of small mam-
mals and small birds are far more often seen by their foes
from above than from below; and from above the effect of
the countershading is at a minimum and is usually nil.
Mr. Thayer, when he observes birds or squirrels, naturally,
like other observers, looks up more often than down; and
unconsciously he ascribes to beasts and birds of prey this
up-looker attitude. But most birds of prey have habit-
ually the down-looker attitude toward both the birds and
mammals on which they prey, and most beasts of prey have
the down-looker attitude as regards the majority of the
mammals on which they prey. Mice and shrews make up
the immense majority of mammals, considered as indi-
viduals, not as species. Almost all their foes look down on
these small mammals from above. Hawks, owls, shrikes,
crows, foxes, bears, skunks, coyotes, cannot look up at
mice. They look down on them. Weasels follow them
purely by scent; and when they do see them it is always
from above or on a level, and usually from a little above.
Rabbits are looked down at, not up at, by eagles, big owls,
big hawks, wolves, cougars, and lynxes; foxes generally
look at them from a level or from above; weasels look up at
them, but, as a matter of fact, practically always hunt them
by scent. Comparatively few species of birds, when adult,
have mammals or reptiles as their chief foes. The great
majority of adult birds have to fear primarily other birds
—eagles, hawks, and owls. These birds of prey usually
approach them from above, frequently from one side,

94 AFRICAN GAME ANIMALS

almost never from below. This is true of thrushes, war-
blers, sparrows, snipe, and ducks. Now, birds in tree tops,
notably in a giant tropical forest, but also even in our own
woods, are so difficult for us to see as we stand (usually
with the light in our eyes) on the ground, even when bril-
liantly colored, that hasty observers are apt to think that
these brilliant colors have some concealing value. There is
no warrant whatever for such belief. The foes of these
birds are up among them or above them; and from above
or one side these birds at once become visible; although in
the flood of light and color, and in the immense confusion
of vistas and obstacles, among the tree tops, they are all,
whether plainly or brightly colored, infinitely more difficult
to make out than when in the dark uniform monotony of
the landscape underneath the trees. Let any man look
out from the second story of a Virginia house in the woods
at the summer redbirds in the trees on a level with the
window, or in similar fashion at the Baltimore orioles in
the elm from the third story of a Long Island house—we
are speaking from actual experience—and contrast the
ease of seeing these brilliant birds from above or from a
level with the difficulty of making them out when staring
up at them in the tree tops with the light in one’s eyes.
But even from above or from a level they are far more diffi-
cult to make out in the tree tops than they are on the rare
occasions when they come to the ground; a Baltimore oriole
on the lawn, and still more a scarlet tanager on a low bush
in the woods, glows like a live coal. Asa rule, birds which
live on or near the ground in forests are sombrely colored;
they are hard to see because they are not brightly colored;
countershading ordinarily plays no part in their conceal-

COLORATION 95

ment; a wood-thrush or hermit-thrush is more difficult to
see (as is the case with a meadow-lark in the open) when
it sits with the nearly uniform dulness of its back toward
the observer—although there is then no effect of counter-
shading—than when it sits sideways, when there is an effect
of countershading but when this effect is more than offset
by the brightness of the white under-parts. As regards
these birds it is possible that natural selection has played
a certain part, not in producing a given pattern, but by
eliminating the birds that were brilliantly or advertisingly
colored.

In the infinitely varied and brilliantly lighted landscape
of the tree tops, the coloration of the bird becomes so
much less important that selection ceases to eliminate the
brilliant hues. That selection does not in these circum-
stances produce the brilliant hues, as Mr. Thayer fancies,
is evident from the fact that many of the birds, especially
the females, which most need protection, remain dull-col-
ored; the females of the scarlet tanager and redbird are
no more conspicuous than the thrushes which dwell in the
comparatively sombre region beneath them. Of course, if
the brilliant hues of the males were developed by natural
selection for concealing purposes, the females would also
have developed them. The fact simply is that the bril-
liant colors have been developed in spite of their adver-
tising quality.

What happens is analogous to what Mr. Reighard has
described in his ‘Experimental Study of Warning Colora-
tion in Coral Reef Fishes’’; he shows that certain fishes,
whose habits and habitat render concealing coloration of
no consequence to them, develop brilliant, highly advertis-

96 AFRICAN GAME ANIMALS

ing coloration patterns, unhampered by selection, in conse-
quence of some tendency or internal force having no refer-
ence to utility. Immense groups of birds, African bee-eaters
and sunbirds, for instance, American humming-birds, al-
most all kingfishers everywhere, gulls (in the adult form),
pelicans, most herons, darters, and cormorants (of all ages
and sizes), most adult male ducks, all swans, the whole
crow family, the whole grackle family, and multitudes of
others, have developed a highly advertising coloration with
no seeming relation to utility; and neither in these birds
nor in the great majority of the ordinary dull-colored birds
does countershading play any part in concealing them.
The effort to explain all or almost all patterns of coloration
in birds by natural selection working either for concealing
or directive markings represents nothing but the darken-
ing of wisdom; it serves merely to bemuddle an already
sufficiently difficult problem. What is needed is the kind
of study advocated in, and instanced by, Witmer Stone’s
“Phylogenetic Value of Color Characters in Birds.”
Among mammals, squirrels are very apt to have an
advertising coloration. Countershading practically never
helps them. Among mice and shrews the species in which
countershading can have any survival effect in concealing
them are very, very few—and probably there are none.
In sum, it appears certain that although the discovery
of the effect of countershading has a very important artistic
bearing, it is of practically no value as affecting the question
of concealing coloration in the higher vertebrates, the mam-
mals, and birds. Among mammals it can by no possibility
be of consequence save in a very few cases, and it is by no
means certain that it is of consequence in these. Among

COLORATION 97

birds it may affect a slightly larger proportion of species,
but the number remains but an insignificant fraction of
the whole. In the great majority of species both of birds
and of mammals countershading is certainly not a factor
of survival value, such as could in any way affect or be
affected by natural selection. Even in the comparatively
few remaining cases, where it is possible that it has a small
effect of solid value in concealing the species, it is very
doubtful if it has been produced by natural selection, or in
any other way, because of its concealing value; it seems far
more probable that it is produced by some such cause as
that which makes the under side of most leaves lighter than
the upper, and that its trivial influence in increasing the
concealing value of the coloration in these few species rep-
resents nothing but an accident.

One reason why the study of the coloration of big game
is of prime scientific importance is that it enables us to
visualize to ourselves something of the effect which the
coloration of small animals has on their prey or their ene-
mies. Nobody in his senses regards a cock ostrich as con-
cealingly colored; yet it is exactly as concealingly colored
as a cock bobolink, or tanager, or magpie, or lark-bunting,
or yellow-headed blackbird, or red-winged blackbird, or
cow-bunting, or purple grackle, or oyster-catcher, or black
and white spur-winged plover, or European blackbird, or
black and white chat. An ostrich looms to our eyes pre-
cisely as a coot does to the eyes of a mink or a prairie-hen
to the eyes of a black-throated ferret; an ordinary antelope
appears to the eyes of a hunter from the same angle that
a deer appears to a cougar, or a zebra or buffalo to a lion.

Another thing always to keep in mind is that in nature

98 AFRICAN GAME ANIMALS

very, very few animals are seen in simple surroundings.
The ordinary man thinks of a zebra or a tanager or a tiger
or a bird of paradise as he sees it in a museum or as he
sees its picture in a book, when it jumps to the eye; and
when in nature it does not jump to the eye he immediately
ascribes the concealment to the animal instead of to the
landscape. There are in nature landscapes so flat and
uniform that every living thing on them is thrown into
relief; and others, not quite so uniform, but where anything
conspicuous is easily seen. All the mammals and birds in
the first type of landscape, and most of those in the second,
are readily seen. Antelope on the bare, flat plains, and
smaller animals on these plains if the grass is very short,
sea-birds on or over the ocean, orioles and blackbirds and
robins on lawns, loons and ducks on lakes or wide rivers,
cormorants, auks, puffins, guillemots on cliffs and rocky
islands, herons and pelicans and ibis along beaches and
bare shores—all show as distinctly as they would under like
conditions of light and distance in a museum. If in these
places any birds are concealed as young gulls and stilts
or some grouse and night-hawks are concealed, the conceal-
ment is due to coloration and not to the landscape.

But most landscapes are not of this simple character.
Animals of all colors and patterns that live in grass below
the level of the tops, or in forests, or even in bare country
which is much broken and accidented, become extremely
difficult to see. The inexperienced or unthinking man has
no idea of the extreme complexity of the average landscape.
The myriad of varying lights and shadows, the countless
gradations of color, the innumerable twigs and leaves, the
hummocks and irregularities and infinitely varied details

COLORATION 99

of the accidented landscape, the multitude of varying vistas,
interlaced and superimposed, and of every length and size,
make it well-nigh impossible for the eye of man or of beast
or bird of prey to pick out anything motionless. Unless
of strikingly conspicuous coloration, any animal that is mo-
tionless escapes notice. Hawk and owl and hunting man
are all alike—ever on the watch for movement; and this is
also true of the things that are hunted. This is why any
animal that “freezes”? (as that keen observer, Mr. Seton,
calls it), from an elephant, or rhino, or buffalo to a grouse
or meadow-mouse or vireo, is so hard to make out. The
immobility and the cover and surroundings count for infi-
nitely more than the coloration. A first-class naturalist
and observer, Mr. Pyecraft, in his “Infancy of Animals,”
speaks of the pattern of cover-haunting animals as “form-
ing a more or less perfect obliterative coloration, causing
the solid body to vanish, as it were, into thin air.” In
reality it is the landscape, and not the coloration, which
thus obliterates the animal. The spotted, the striped, and
the unicolored animals are all obliterated in precisely the
same manner. The landscape, not the coloration, is the
obliterator in the great majority of cases. All that is de-
manded of the coloration in these cases is that it shall not
be of such a highly advertising type as, say, black or white
—and nature does not always insist even on this modest
requirement.

The ultra natural-selection and concealing-coloration ex-
tremists, who treat concealing coloration as being of primary
importance to the welfare of the great majority of mam-
mals, including the big mammals, and as being almost uni-
versally present, should carefully study the big bovines,

100 AFRICAN GAME ANIMALS

cervines, and other grass-eaters, and the big felines, canines,
and other flesh-eaters, in their native haunts. As regards
big mammals the theory is certainly untenable. Not only
are most big mammals not concealingly colored, but in most
of the cases where the concealing coloration exists it confers
no advantage on the possessor, because of the overwhelm-
ingly superior importance of such matters as bodily power
and habit.

Probably the most effectual concealing-coloration pat-
terns among big mammals, under the average conditions of
actual environment, and at the crises of their lives, when
they prey or are preyed on, are the countershaded khaki or
tawny brown of the lion and cougar and the countershaded
gray of the ass, wild or tame. The lion and cougar we have
studied with care when wild; we have never seen the wild
ass; but the tame ass and feral ass, colored in substantially
the same manner, we observed under natural conditions on
the Western plains. Among the horses, cattle, and asses
out on the range, observed under all weather conditions at
all seasons, and by day and night, the gray countershaded
donkeys were on the average the most difficult to see. Next
came the dun or claybank horses, which were not counter-
shaded; usually such a horse was at night almost or quite
as inconspicuous as a donkey, except for the size. Then
came the bay, brown, and red horses and cattle; and finally
the black and white, which, under almost all circumstances,
by day and by night, were the most conspicuous. At cer-
tain times at night the donkeys were practically invisible,
even when very close, until they moved.

In Africa, of the big game of the plains and the thin
forest or scrub which we observed, the eland, big gazelle,

COLORATION 101

roan, and oryx, in that order, most resembled in their body
patterns the coloration of the wild ass, and were, on the
whole, the most difficult of the big game to see when feed-
ing or resting. The face markings of the oryx and roan
and the long horns of the oryx possessed some advertising
value; and both eland and oryx, when not too distant, were
advertised by the continual switching of their long, black-
tipped tails; again and again, if the animals were among
bushes or trees, it was this continual tail-switching which
first betrayed their presence. Doubtless the same thing is
true of the roan, but we happened not to observe it. When
at a distance such that the various black and white mark-
ings and the horns were invisible or indistinct, the general
body color of all three animals harmonized well with the
ordinarily almost leafless or brown-leafed scrub and dry
grass; if in company with hartebeest, wildebeest, or zebra,
any of these three animals were almost always seen before
the eland, oryx, or roan. In the flood of bright sunlight
which was usual the wildebeest was most conspicuous,
near or far; at night it was, on the average, most conspicu-
ous, although on moonlight nights the zebra sometimes
seemed white, and was then the most conspicuous animal
of all. The topi, perhaps because of its inversely counter-
shaded coloration, came next to the wildebeest in conspicu-
ousness under average conditions, while of the hartebeest
and zebra one was the more conspicuous under certain con-
ditions of light and shade, and the other when the condi-
tions were reversed, the coloration of both being normally,
and at the most important crises of the animals’ lives,
advertising and not concealing. The bull sable, buck
white-eared kob, and buck white-withered lechwi possessed

102 AFRICAN GAME ANIMALS

coloration patterns of a strikingly advertising quality.
The stripes of the eland had no effect, one way or the other,
in advertising or concealing it, when compared with the
unstriped roan. The black lateral and leg markings of
the oryx were advertising, as was the case with the black
lateral lines on the Tommy gazelles and on the female and
young Grant gazelles. The male Grant gazelle was as
concealingly colored as the roan and eland and, because of
its smaller size, less conspicuous. The dark-gray water-
buck and reddish impalla were more conspicuous when
away from cover, and when thus away from cover never
sought safety in trying to escape observation; nor did the
reddish kobs. But these animals often sought the con-
cealment of cover, unlike the regular game of the open
plains. The waterbuck, eland, roan, oryx, and big gazelle
seemed a trifle the least conspicuous on moonless nights;
if the nights were dark all the animals were practically
invisible, and yet these seemed to be the nights when the
lion was boldest and most successful. The wildebeest,
topi, and hartebeest are advertisingly colored compared to
the eland, oryx, and roan of the same habitat. We saw the
former more readily and at a greater distance than the
latter, but careful study on the ground failed to show that
this was of any disadvantage to the former or produced any
change in habits.

The big game of the plains live substantially similar lives,
and trust exclusively to their senses, their watchfulness, and
their ability to run away. Coloration plays no part in
shielding them from their foes; nor do they seek to take
advantage of cover; habit is everything. The lion’s eye-
sight is so good that probably none of these animals ever

COLORATION 103

escape his notice unless at a distance that renders it im-
material whether he sees them or not; he trusts mostly to
his sense of smell; he ordinarily hunts at night, and is most
successful on dark nights, when the coloration is of little
consequence; and when near enough to be dangerous,
whether by day or night, he sees so clearly that the color
of the quarry is a negligible factor—an eland or a roan is
evidently seen as clearly as a sable, a wildebeest, or a zebra.
Hartebeests in the aggregate far outnumber oryxes, yet
relatively to oryxes they are conspicuous, and their habits
are identical so far as escape from their foes is concerned.
Evidently the advertising color of the hartebeest’s coat has
not been of the slightest disadvantage to him in the struggle
for life. As is the case with all other plains game, his
habits are such that his coloration is from this standpoint
of no consequence. The hartebeests in a state of nature
are probably the most successful and widely distributed of
all African animals of their size, yet they are not conceal-
ingly colored and carry no recognition marks.

The ostrich offers a very instructive case in point. The
black and white cock ostrich is, except the elephant, and
possibly the giraffe and rhino, the most conspicuous creature
of the plains. The soberly colored hen cannot be seen at
all at a distance when the cock is already plainly visible.
She may fairly be said to be concealingly colored. He is a
striking example of advertising coloration. But both have
precisely the same habits. Save when brooding—and both
brood alternately—the great birds never seek to escape
observation, never do escape observation, and trust for
safety solely to their wariness, their eyesight, and their
speed. The nest is placed on the bare sand, usually with

104 AFRICAN GAME ANIMALS

bushes round about, and, when on it, both birds lie flat, with
outstretched neck, if they suspect danger. It is said that
the hen broods by day and the cock by night, but twice we
came on cocks brooding by day. Moreover, by moonlight
the cock’s coloration is almost as conspicuous as by day;
and on all nights, if the birds can be seen at all, the cock
is most conspicuous. At all times, therefore, including the
most vital moments and crises of their lives, the cock’s
coloration is boldly advertising and that of the hen some-
what concealing. Yet the habits of the two are identical.
Both have the same foes, the lion and very occasionally
the leopard, while the young are in danger from hyenas,
jackals, eagles, and big hawks. The birds are monoga-
mous, so that the numbers of the sexes are substantially
equal; which means that the highly advertising coloration
of the cock is of no disadvantage to him in the struggle for
existence. Evidently his coloration has not been developed
by natural selection or because of any utilitarian purpose,
and, equally evidently, the relatively concealing coloration
of the hen has no survival value as compared with the cock’s
coloration.

The same statement applies to various antelope. The
female and young Grant gazelle and the Tommies of all
sexes and ages have a lateral black band which is dis-
tinctly advertising. The adult male and occasionally the
old female Grant gazelle lack the stripe, and their coloration
becomes concealing. Yet there is absolutely no difference
in habits, and no benefit or detriment is caused by either the
presence or the absence of the black lateral band. The adult
males of the sable, white-withered lechwi and white-eared
kob are advertisingly colored compared with the females;

COLORATION 105

but this has not made any difference in their habits, and the
advertising coloration of one sex and concealing coloration
of the other apparently confer neither any advantage nor
any disadvantage in the struggle for life. Habit and cover
are the vital factors. The old bucks of the white-withered
lechwis are inversely countershaded, being white on top
(like one species of serow). We are often asked by the con-
cealing-colorationists to consider how the coloration of the
tiger would reveal it if it were inverted; well, in these ante-
lopes the coloration 7s inverted! and no detrimental results
follow, although they, also, are cover-hunting species. This
fact alone is sufficient to cast grave doubt on the assertion
that the tiger’s stripes and countershading and white belly
have been developed by natural selection for concealing
purposes; probably they have no survival value. ‘The sit-
atunga dwells in reeds on water-soaked ground like the
lechwi; their colorations are so totally different that they
cannot both be concealing. As a matter of fact, it is prob-
able that neither is; but that of the adult buck lechwi is
certainly revealing.

In the deep, moist forests the true big-game animals are
the beautifully striped and richly tinted bongo and the
dark monochrome forest hog. Both have the same foes
and the same habits so far as their relations to these foes
are concerned. The colorations are utterly unlike, and if
one is concealing the other must be revealing. As a mat-
ter of fact, neither has any effect in either concealing or
revealing the wearer; habit and cover are all-important.

The bushbuck extends over vast areas of Africa, and its
various forms exhibit the widest variety of coloring. In
some regions the coat is a network of white stripes and

106 AFRICAN GAME ANIMALS

spots (the patterns, however, being very different in dif-
ferent forms); elsewhere these are reduced in the old males
almost to the point of total disappearance. Their foes are
everywhere the same. Their conditions of life vary as
much concerning certain of the harnessed forms—the form
we found in the dry, open, thin forest of the Lado, for
instance, compared with the form found in the dense wet
jungle of Uganda—as they do between the harnessed forms
and the nearly solid-colored forms. Solid-colored duikers
are found in the haunts of both the solid-colored and the
patterned bushbucks. Under these circumstances it is im-
possible to regard the pattern or lack of pattern as having
any survival value in concealing the animal from its foes;
some factor other than natural selection working toward a
concealing coloration has been at work in securing the re-
tention of the pattern in some forms and its reduction or
elimination in others. The great explorer and naturalist
Dr. Schweinfurth regarded the harnessed bushbuck as
conspicuous because of its striped and spotted coat. I am
inclined to agree with him; but in any event it is quite
impossible that one form of bushbuck is concealingly col-
ored because it does possess, and the other because it does
not possess, the stripes and spots.

Of the other bush game the reedbuck is colored much
like the white-tail deer. It trusts to hiding for conceal-
ment. So does the oribi, where the grass is long; but where
the grass is short its habits are like those of the gazelles.
The duiker is, of all these antelope, the one that can most
properly be called concealingly colored; and we are inclined
to think that its dull gray, slightly countershaded coat
does have a concealing value. But we are not certain, for

COLORATION 107

its stealthy, skulking ways, and its habit of clinging to
and taking advantage of cover, are such that it would be
difficult to detect anyhow. The same observations apply
to the dikdiks. The Chanler reedbuck behaves much like
the klipspringer; its coloration is more revealing.

The steinbok offers an interesting problem when com-
pared with other buck of the same size but different habits,
such as the klipspringer and Tommy gazelle. The klip-
springer lives among rocks; its coloration tends to har-
monize with its surroundings; and with the possible excep-
tion of the duiker it is the only one of all the African
antelopes in which it seems as if the countershading may
add to the concealing quality of the coloration. But, com-
pared with the steinbok, the klipspringer does not hide at
all; it is a noisy little buck, continually uttering a shrill
whistle of alarm or curiosity; it trusts for safety to its
vigilance and its ability as a rock-climber. The Tommy
never tries to hide. It lives on the bare, open flats, and trusts
to its vigilance and speed for protection. The steinbok, on
the other hand, which lives on the grassy flats and among
bushes, both on the plains and along the hillsides, is a
skulker and hider; yet its coloration is not concealing.
Its bright-red coat was advertising under all the condi-
tions under which we observed it; nor did we anywhere see
a normal background with which its coloration would have
harmonized. Nevertheless, it trusted almost exclusively to
concealment for safety. Nor did it merely use as a screen
thick cover so impenetrable that through it no coloration
could be made out. It continually endeavored to hide in
very scanty cover; we often found it practically concealed
in tufts of rank grass or behind or alongside of some small

108 AFRICAN GAME ANIMALS

bush. As soon as it saw a man at a distance its tendency
was to lie down; it treated the recumbent posture as its
natural position when threatened; whereas the Tommy
always felt that it was at a disadvantage when lying down,
was uneasy, and immediately sprang to its feet when it
saw what it regarded as a possible, but distant, menace.
The steinbok lay down when its foe was at a distance in
hopes to escape observation, but kept its head raised watch-
fully. If the foe came nearer and it still hoped to escape
notice it laid its outstretched neck and head on the ground,
and sprang up only when it deemed it hopeless to expect its
enemy to pass it unseeing. When it ran it would often stop
behind a bush, after going a few hundred yards, and lie
down again; and then it might be readily stalked. If under
or beside a bush, the shade, or, if the foliage was thin, the
checkered play of sunlight and shadow, made it easy for
it to escape the eye unless we knew just where to look for
it; but so far as, and whenever, its coloration had any effect
it was an advertising effect. I am somewhat puzzled how
to account for these facts in the compared cases of the stein-
bok and klipspringer. The latter has a much more con-
cealing coloration than the former, but is noisy and alert,
and trusts hardly at all to concealment; whereas the animal
that lives under conditions that would seemingly make a
concealing coloration helpful, and whose habits also would
seemingly make it helpful, has no concealing coloration.

If we trusted only to such cases as this we would be
inclined to agree with those ornithologists who, like Mr.
Outram Bangs, doubt whether concealing coloration is cor-
related with the hiding instinct, and are inclined to believe
that all animals, no matter what their coloration, alike tend

COLORATION 109

to try to escape observation by remaining motionless.
There are, however, plenty of instances to the contrary. A
good example, among closely related species, is the case of
the desert chat compared with the black and white chat.
Both birds are found along the edge of the Egyptian desert.
The black and white chat is a boldly colored bird, with
actions which correspond; it is alert, wary, active; its col-
oration advertises it at a long distance, and it makes no
attempt to hide, dodging round rocks and through crev-
ices, and trusting to its agility and watchfulness for safety.
The desert chat is colored above much like the desert itself;
when it crouches it offers an excellent example of concealing
coloration (non-countershaded), and it makes use of this
fact in a way that tends to show that the coloration is, in
this case, a factor of survival value; for its main reliance,
when danger threatens, is in crouching motionless to avoid
observation. As regards the black and white chat, it is
self-evident that neither natural selection nor any other
tendency toward the production of a concealing coloration
has been at work; evolution has brought about a highly
advertising coloration of a kind which can hardly serve any
purpose of utility, and habit is practically the sole factor
in securing the bird’s safety from its foes. (Mr. Thayer’s
theory that their coloration has any effect, as regards any
of these insect-eating birds, in helping them secure their
prey need not be taken seriously.) As regards the desert
chat, the coloration may have been developed by natural
selection for purposes of concealment; but as it is of sub-
stantially the same type as the coloration of many other
desert animals which do not trust to, and are not benefited
by, concealment, it seems more likely that it is due to

110 AFRICAN GAME ANIMALS

other forces generally, although not universally, at work as
regards desert animals. If this is true, the habit of acting
so as to benefit by the coloration has become ingrained,
while no such habit has arisen, or has persisted, in the case
of the black and white chat, where the coloration is such
that no action would make it beneficial for concealing pur-
poses. The steinbok, on the other hand—although it does
not possess a concealing coloration—does not possess an
extreme and striking type of advertising coloration, and
in its case the habit of endeavoring to escape notice by
immobility, so wide-spread among living creatures, has per-
sisted, and, to judge by the fact that the species is common
over a wide range, has been successful as a survival factor.

Among the African carnivores we came across two or
three of the lesser species with a strikingly advertising
coloration; the ratel, for instance, and the white-tailed
mongoose, a highly predatory creature. What “warning”
function this coloration can serve in these two cases it
would be hard to imagine. The black back of the black-
backed jackal is revealing; but the animal thrives as well
as its more concealingly colored kinsfolk. Most of the
smaller carnivores have colorations that if they do not
conceal at least do not reveal them. The hyenas include
three forms—a brown, a spotted, and a striped. Their
habits, their lack of enemies, and the circumstances under
which they make their rare attacks on living things make it
difficult to believe that they have special need of any one
of these three coloration patterns; doubtless the coloration
of no one of them serves any concealing or other useful pur-
pose to the animal at present; and it certainly seems as if
the three totally distinct patterns had been developed for

COLORATION ie

some non-useful reason, without any reference to natural
selection. The habits of the hunting-dog forbid the belief
that its conspicuous coloration has any effect one way or
the other on its success in life, or that it has any survival
value such as would give a chance for the working of natu-
ral selection.

There remain the lion, the leopard, and the cheetah. On
the whole, as we have said, the lion’s coloration, like the
cougar’s, represents pretty nearly the maximum conceal-
ing power for a beast which—unlike the nearly stationary
animals or those of narrowly limited habitat—necessarily
roves through many and varied kinds of landscape. But
there are features of the coloration which are not concealing
—the lightish and black-edged patch on the rump, for in-
stance—while the black tips of the ears and tail are often
the first things to attract attention. Much more impor-
tant, however, is the mane. This is always of a somewhat
advertising value, and when it is black it has a strongly
advertising value. Yet black-maned lions are very fine
and powerful beasts, at least up to the general level of their
kind, as well-fed, and as strong. The fact that a conspicu-
ous black-maned male and an inconspicuous maneless male
or female do equally well in killing game rather shakes
our belief in the survival value of the lion’s general conceal-
ing coloration. We are inclined, having this fact in view,
seriously to doubt whether the lion’s coloration is a sur-
vival factor, although it is probably, among all the col-
orations of all the big cats, bar only the puma, the most
effective concealer. Young lions are dimly spotted and
striped; probably the lion comes from a spotted and striped
ancestral form. The loss of the spots is ascribed by natural-

112 AFRICAN GAME ANIMALS

selectionists to the fact that the lion has changed its hab-
its and become ‘“‘a dweller in the open,” while the tiger
(so closely kin to the lion that the species interbreed, their
skeletons being almost indistinguishable) dwells “in dense
jungle, where his stripes, harmonizing with the tall reed
stems, make him invisible, and enable him to stalk his
prey without being seen.” This explanation is probably
opposed to the actual facts. The tiger is very rarely found
among reed stems; reed-dwellers like the lechwi are rarely
striped, and, as we have already said, Kearton’s photo-
graphs show that the stripes are much more conspicuous in
the shadow of a jungle than in the open under the sunlight.
The sambur is one of the animals on which the tiger habitu-
ally preys; the two dwell in the same dense jungles; the
sambur is solidly colored, a dark unicolor without counter-
shading, and yet Hornaday has shown that it is peculiarly
hard to see, harder than the tiger. Moreover, the tigers
of Mongolia and Manchuria live in surroundings wholly
distinct from those of Hither and Farther India. We do
not believe that the tiger’s coloration, as being of any use to
it in eluding the vigilance of its prey, is a factor of survival
value, or has been developed by natural selection for this
particular utilitarian purpose.

The cheetah is a diurnal animal, and lives far more
completely in the open than the lion. Yet the cheetah has
retained its spots. The spotted coat of the leopard is often
advanced as “‘the most perfect imaginable concealer,” hav-
ing “the flecked leaf and sunlight pattern” best suited for
a dweller in woodland. There is advantage in consid-
ering together these four big cats, the lion, tiger, leopard,
and cheetah, with the two big American cats, the puma (or

COLORATION ~~. 113

cougar) and the jaguar; four of the six we have ourselves
carefully studied in their native haunts. As soon as this
study is seriously undertaken it will be seen that it is infi-
nitely harder to draw sound conclusions than the extreme
partisans of selection and concealing coloration admit. In
the first place, as to the loss of spots by the lion and puma,
alleged to be because of their taking to a life in the open:
The cheetah lives more in the open than either, and is more
diurnal in its habits, yet it has retained its spots. At a
short distance in the African sunlight these spots disappear;
probably in view of the habits of the cheetah it would be
of no slightest consequence to it if the coloration of the
lion, leopard, or tiger were substituted for the coloration it
actually possesses.

Moreover, the spotted cats, the leopard and jaguar, and
the unicolored cats, the lion and puma, have for ages pos-
sessed overlapping ranges, in which both types of coloration
have persisted side by side practically unchanged and with
seemingly equal benefit to the wearers. The New-World
and Old-World couples, each of one spotted and one uni-
colored cat, reverse conditions of distribution. In the Old
World the spotted leopard is far more widely distributed
than the unicolored lion and occupies most of the lion’s
range and habitat. In the New World, on the contrary,
the unicolored cat, the puma, occupies a far wider range,
which includes most of the jaguar’s range and habitat. In
most of the open country in which the lion is found, from
Somaliland and the Lado and East Africa to the regions
south of the Zambesi, the leopard is also found; and every-
where within this vast range they are most often found in
the same cover of thorn-trees, bushes, reeds, and grass; al-

114 AFRICAN GAME ANIMALS

though the leopard also penetrates the deep forests in which
the lion does not dwell. In the reeds, the very cover sup-
posed to be peculiarly fitted to the tiger’s coloration, both
lion and leopard are practically invisible; for in such cover
it is the physical screen of the cover and the habits of the
big cats that are practically the sole causes of their invisi-
bility. The lion’s coat is supposed to have been developed
to suit the open country; but a leopard in the grass, under
the flood of bright light, is at least as difficult to see as a
small lioness and more difficult to see than a black-maned
lion. This we can testify from our own observations in the
field. In the sun-flecked or uniformly dark shade of the
bush or tree cover there was again no difference perceptible
in the average visibility of the coloration; if anything, the
leopard’s coat was just a trifle less concealing. But the
difference, one way or the other, was wholly unimportant.
The leopard was smaller than the lion, was even more cau-
tious and wary, and was an even greater adept in stealthy,
sinuous crawling and hiding; and therefore it was, on the
whole, harder to see, whether in grass-land or bush.

The contrast between the two big cats and the game on
which they preyed, under like conditions, showed the utter
unimportance of color compared to habit. There were in
places enormous stretches of flat country covered with grass
a couple of feet tall more or less, either entirely treeless or
very thinly studded with occasional thorn-trees. Every
zebra, eland, wildebeest, hartebeest, topi, roan, oryx, kob,
waterbuck, or big gazelle, and, of course, every giraffe and
rhinoceros, in such a stretch, was visible; the animal made
no attempt to avoid observation and no attempt to crouch
and hide when it discerned danger. But if, as often hap-

COLORATION 115

pened, there were lions and leopards in such a tract, it was
difficult to see or find the former, and far more difficult to
see or find the latter, because both species were always
slouching and crawling, or lying down in some thicket or
tall bunch of grass; and either one of them when it first
saw danger, if it thought itself unobserved, promptly
crouched and lay motionless or else slunk rapidly and
stealthily off below the level of the grass tops. Neither the
lion’s nor the leopard’s coloration was in any degree more
concealing than the coloration of some or most of the grass-
eaters; if the big cats had been mounted on higher legs and
had possessed the habits of the grass-eaters, their coats
would have shown no more concealing quality than the coats
of the hartebeest, eland, and oryx. Habit, and especially
the ability to take advantage of cover, were the vital factors,
the survival factors, in securing concealment; the exact
shade or pattern of the coat was a wholly negligible factor.

The case of the puma is very instructive in this regard.
It is as plentiful as the jaguar in the great tropical forests,
and is found in many great temperate forests which the
jaguar never reaches. Mr. Thayer gives an ingenious pic-
ture to show how the jaguar’s spotted hide absolutely con-
ceals him among the foliage (not natural foliage, but foliage
put in by Mr. Thayer to prove his point), and points out
that if it were unicolored the animal would be revealed;
he forgets that the puma, equally common in exactly the
same surroundings, is unicolored and is not revealed. Now,
our experience, in accord with the experience of most hunt-
ers and out-of-door naturalists, is that the cougar is, of all
American big animals, the most difficult to see and the
most rarely seen. Its neutral-tinted, nearly unicolored,

116 AFRICAN GAME ANIMALS

slightly countershaded coat unquestionably has a conceal-
ing quality in the woods and among clay banks and rocks
under ordinary conditions, and for long we accepted this
as the chief element in explaining its invisibility. But
when we came to think out the matter we realized that a
totally different theory must be invoked to explain the
cougar’s invisibility during winter in that large part of its
range which at that time is snow-covered. We have spent
some time in country where cougars were common, on the
Little Missouri and in the Rockies. In places the land was
thickly forested; in other places it was open, the bad lands,
hills, and mountains being sparsely clad here and there
with pion, cedar, buckbrush, and the like. The wapiti
and blacktail that lived in such places were easily found and
seen—unlike what was true in the dense forests. But the
cougars were invisible, and this was just as true in winter,
when the landscape was white, as in summer. They could
be trapped, and they were readily killed with hounds; but
even the oldest and most skilful hunters hardly ever saw
them on other occasions. This means that the cougar’s
coloration was really an insignificant and practically negli-
gible factor in its concealment. The prime factors were
its nocturnal habits, its sharp senses, its wonderful ability
to take advantage of even the scantiest cover, and its power
of lying indefinitely motionless and of advancing with in-
conceivably noiseless stealth. The cougar is found in boreal
regions of heavy snowfall and in steaming tropical forests,
on bare broken plains and in the Rockies and the Andes, in
the great hardwood and coniferous forests of the United
States and on the barren Patagonian grass-land, in thick
cover and in scanty cover. Everywhere it is equally invisi-

COLORATION sb hg

ble. It would be as invisible in the homes of the tiger and
leopard as it is in the home of the jaguar. The black
leopards and jaguars really are as invisible in their homes
as are their spotted kinsfolk. The conclusion seems inev-
itable that no one of the coloration patterns, that of lion,
tiger, cougar, cheetah, leopard, or jaguar, has any survival
value as a factor in the life-history of the animal; it cannot
have been produced by natural selection working for con-
cealing coloration. We believe that the same conclusion
applies to the ounce, lynx, ocelot, clouded tiger, jaguarondi,
eyra, wildcat, and all the other spotted, mottled, and uni-
colored cats, big and little. It is greatly to be wished that
some trained scientific observer and investigator would
study the cat family—and for the matter of that, such dis-
tinctively colored groups-as the tragelaphs and the deer—
along the lines laid down by Witmer Stone in his pamphlet
on the “Phylogenetic Value of Color Characters in Birds,”
elsewhere referred to.

The giraffe, because of its size and shape, is always
recognizable at such a distance that its coloration is of no
consequence; the reticulated giraffe has a bright-reddish
coat of a highly advertising quality; but this does not hurt
it any more than does the common giraffe’s less conspic-
uous—but, when close, conspicuous—coat. ‘The coloration
is a negligible factor. If among trees, in the flood of the
African sunlight, the giraffe may escape the notice of an
ordinary native or white man; but it practically never es-
capes the notice of the trained eyes of the native hunting
tribes. Good observers have expressed their surprise at
its being difficult to see. Our experience was that on the
average it was seen at a longer distance than any animal

118 AFRICAN GAME ANIMALS

except the elephant. It never sought to hide or to escape
notice. The black, monochrome buffalo was not nearly as
conspicuous, in spite of its coloration, because the giraffe’s
size and shape rendered it visible at such a distance that
the coloration was of no consequence. Even the bulky
monochrome rhinoceros was usually made out at a distance
less easily than the giraffe; for the giraffe rarely lies down
and the rhino lies down as freely as a pig. Of course, under
certain circumstances every animal, no matter of what
shape, size, or color, even the elephant itself, becomes hard
to see if it remains absolutely motionless. The coloration
of the giraffe might, seemingly, be of benefit to the buffalo
on account of the buffalo’s size and habits; yet, as a matter
of fact, the buffalo has a dark, highly advertising coloration
—intensity of color being of far more consequence in either
revealing or concealing an animal than either pattern or
countershading. But the buffalo is not seemingly harmed
by its coloration.

The study of American big game shows that coloration
plays an unimportant part as a factor of survival value in
their life-histories. The bison and peccary are neither
countershaded nor concealingly colored. The color of the
white goat is highly advertising, especially at the most
critical period of its life, when the kid is young and is in
danger from the eagle. The prongbuck, although counter-
shaded to the extent of having a white belly, possesses a
highly advertising coloration. It never hides or seeks to
escape or does escape observation when adult; the very
young, as with the young of all these animals, do try to
hide, lying flat on the ground with outstretched neck, in
which position the countershading disappears. Moose and

COLORATION 119

wapiti are not countershaded and are revealingly colored;
but both seek to escape notice by immobility and stealth,
and the wapiti crouches flat like a deer. The Rocky Moun-
tain and coast blacktails are, at least in some of their forms,
not countershaded; often, owing to the black hair on the
chest, the fore parts of their bodies are inversely counter-
shaded; but they crouch and skulk, and the general tint of
their coloration is not revealing, and under certain condi-
tions is concealing, although the white rump of the Rocky
Mountain blacktail is always revealing. The mountain-
sheep has a revealing white rump; its general coloration in
the United States and Mexico is concealing rather than re-
vealing; but the black and white sheep of the boreal moun-
tains are advertisingly colored, and yet no difference in
their habits is thereby caused; evidently no one of the col-
oration schemes is a factor of survival value.

The case of the white-tail deer is interesting when it is
compared with the fallow deer of Europe and the axis of
India. In its winter coat, unless when snow is on the
ground, the coloration is not revealing—although its col-
oration is hardly concealing in the sense that is true of the
blacktail and the southern form of the mountain-sheep.
But in spring and summer the bright-reddish coat is of a
very revealing quality against the bright green of its home;
and this at the very time that the breeding or nursing does
and velvet-horned bucks are most in need of protection.
Nevertheless this advertising quality of the coloration is of
no detriment, because the animal is such a finished skulker,
and lives in such dense cover that in summer it need fear
only foes which hunt it by scent. The flaunting flag is very
conspicuous; and the white belly, although to a less degree,

120 AFRICAN GAME ANIMALS

is also advertising. [he young fawn, however, is spotted;
the spots are lost in the adult. In the fallow deer the spots
persist in the adult in its summer coat; and in the axis they
are permanent at all seasons.

The view taken of these phenomena by some moderate
and sensible natural-selectionists and concealing-coloration-
ists, men whose judgment is generally sound, and whose
position must be treated with careful and respectful consid-
eration, is well put by Mr. Pycraft. He says, “‘The fallow
deer is a forest-haunting species,’ and during the summer
months lives “where the play of the sunlight through the
foliage scatters spots and shafts of golden light on every
side. Thus the spotted hides of the deer blend insensibly
with their surroundings.” But in the fall the spots are lost
and the unicolored coat then “no less perfectly harmo-
nizes”’ with the dull landscape. “In the axis deer, also a
forest animal, the spotted coat is worn the year round, and
this because it is happy in living in regions where the
sombre pall of winter is never spread.” Mr. Pycraft ex-
plains that the spotted (and striped) patterns are normal
for species which dwell amid the cover afforded by vegeta-
tion, where they form “a more or less perfect obliterative
coloration, causing the solid body to vanish, as it were, into
thin air,” but “‘that so soon as these patterns fail to serve
any useful end—their purpose is to cut up the solid appear-
ance of the body and so destroy its contour, hence they are
called secant patterns—they begin to disintegrate,” while
they are retained so long “as they confer benefits,” their
“change of form” being ‘‘determined” by “that sumptuary
law which is part of the machinery of natural selection,” for
the pattern degenerates when “the need for a spotted

COLORATION 121

livery is past,’ “because natural selection no longer re-
wards those individuals in which the livery is best de-
veloped.”

This is the best presentation of the theory of the natural-
selectionists. If only the fallow deer and axis were con-
sidered, it would seem convincing. But it breaks down
completely when other deer, the majority of deer, are con-
sidered; for although they still live in the cover afforded
by vegetation, and are descended from spotted forms, the
adults, in the large majority of the species, have lost their
spots. Take the abundant and widely spread white-tailed
deer of America, which, in its various forms, extends from
the northern isotherm marking the northern range of the
fallow deer to the tropics, between the isotherms in which
the axis dwells. The fawns are spotted; doubtless the
adult ancestral whitetails were spotted; the whitetails
live now in just such cover as do the fallow deer and axis;
and yet they have lost their spots and are solid-colored
above. It seems incredible that natural selection can be
responsible for both of two such diametrically opposite re-
sults; and, of course, if being spotted tends to conceal the
deer, then the loss of the spots cannot have been due to
natural selection making for a concealing coloration. This
is self-evident. The red deer, which lives in the same
country as the fallow deer, and the sambur, which lives in
the same country as the axis, have also both lost their
spots in the adult forms. All these deer have substantially
the same foes; wolves or wild dogs, and the big cats. If
a spotted coat really is concealing, then surely natural selec-
tion ought not to have eliminated it in the great majority
of the deer, as it has actually done.

122 AFRICAN GAME ANIMALS

Mr. Pycraft further illustrates his point by the tapirs.
The young tapirs of America and Malaysia are spotted
and striped, while the adult Malayan tapir is boldly pied
black and white; and the spotted coloration of the young
and the pied coloration of the adult are both stated by Mr.
Pycraft to be “of a highly protective character.’’ But the
adult American tapirs, of several species, are of a dark
monochrome, which Mr. Pycraft does not claim to be pro-
tective, and which most concealing-colorationists treat as
highly advertising. The adult tapirs of the present day
are held to be descended from spotted and striped ancestors
and, therefore, to have changed their coloration. But all
the evidence indicates that they have not changed their sur-
roundings, their habitat. They dwell in forest and jungle
now, exactly as their striped and spotted ancestors did.
There has been, as far as we know, no change of environ-
ment to alter the survival value of the pattern. Yet it
has disappeared as completely as in the case of any plains-
dweller. The argument Mr. Pycraft makes as to the reason
for the disappearance of spots among the descendants of
forest-dwellers which now live in the open prairies does not
apply to, and, indeed, seems to be controverted by, the facts
regarding those creatures which continue to dwell in forests
exactly as their ancestors did; for the loss of spots seems
to have gone on in one set of cases much as in the other.
The cheetah lives on the plains, but has retained its spots;
the tapirs and most deer and pigs live in the forests, but have
lost their spots. In the case of the tapirs, the Malayan
animal is said to have developed a concealing coloration by
being given a livery of white, and blackish; yet the more
plentiful American tapirs have developed a uniform, solid,

COLORATION 123

blackish color. The variety of habitats among the Ameri-
can tapirs is wider than is the difference in character of
environment between one or two of the species and the
Malayan tapir. All have foes of the same kind, the big
cats; the tiger and leopard in Malaysia, the jaguar and puma
in America. The coloration of the Malayan tapir can only
be called concealing if the coloration of the American tapir
is admitted to be revealing. As a matter of fact, we do
not for a moment believe that the coloration of the former
is really as concealing as the coloration of the latter; the
observers who so read it really only mean that in their sur-
prise at finding that almost all animals, of all colorations,
are inconspicuous in forest and jungle they attribute to the
animal’s coat an invisibility that is really due to the land-
scape. The observer whom Mr. Pycraft quotes states that
the Malayan tapir, when lying down in the daytime, re-
sembles a gray bowlder bathed in sunlight, the implication
being that natural selection has picked out its coloration so
as to resemble bowlders in the sun. Well, in Africa we
have seen rhinoceros when lying down in the sun mistaken
for ant-hills; and there is no question whatever that an
American tapir would, under similar circumstances, be mis-
taken for a darker-colored bowlder. The facts in the case of
the tapirs do not prove that one coloration pattern is more
effective than another as a concealer of survival value under
the play of natural selection. They prove the exact re-
verse. The facts quoted by Mr. Pycraft, if the interpre-
tation of the natural-selectionists were correct, would show
(1) that tapirs were formerly concealingly colored; (2) that,
without as far as is known change of environment, one of
them has developed a totally different type of concealing

124 AFRICAN GAME ANIMALS

coloration, while all the others, under practically similar
conditions of habitat and of enemies, have developed a
revealing type of coloration. We do not believe this inter-
pretation to be correct. What such facts really show, in
our judgment, is either that there has been a steady trend
away from a concealing and toward a revealing coloration,
or else that, as regards these forest and jungle dwellers,
habit and cover are of such overwhelming importance in the
lives of the creatures that the pattern of coloration ceases
to have any survival value, and is determined by some tend-
ency, or cause, or tissue of causation, entirely distinct from
the operation of natural selection to secure the survival of
the fittest by shielding them from the observation of their
foes. The same conclusion applies to the deer of the for-
est and jungle. Nineteen-twentieths of them are now
solid-colored. The tendency, evidently, is to lose the
spotted pattern, even where the environment is unchanged
so far as conditions telling for concealment are concerned;
which necessarily shows that this pattern does not possess
survival value.

Thus all the facts tend to show the fundamental error
of the entire body of conclusions advanced by the natural-
selectionists as to the present survival value of the spotted,
striped, and pied patterns of coloration among forest-dwell-
ing animals. These are found much more frequently and
much better developed among the young than among the
adults; so that the tendency among the adults has evi-
dently been away from the patterned ancestral type.
Already the tendency has gone so far that the older pat-
terned types are far outnumbered by the monochrome
types among the bigger carnivores and grass-eaters in the

COLORATION 125

thick forests of the temperate and torrid zones. The
jaguar is spotted; but the other big cat of the same region,
the puma, and the big dogs, and all the big animals upon
which they prey—the tapirs, peccaries, and deer—are prac-
tically monochrome, and many of them are not counter-
shaded. The tiger, leopard, and axis, which are striped or
spotted, are far outnumbered by the monochrome or prac-
tically monochrome lions, dholes, wolves, boars, deer, ante-
lopes, and oxen of the same forests and jungles. There is
one pied tapir; there are several monochrome tapirs. The
wild oxen are practically monochrome, except for one or
two species which have an advertising white rump; nine-
tenths of the species are monochrome. In Africa the same
tendencies are manifest. The nearly monochrome or nearly
unicolored species of the forest, the jungle, and the reed
beds much outnumber the species which are appreciably
spotted or striped, and among these the tendency evidently
is for the spots or stripes to disappear; the elands, evidently
descended from a highly striped type, have in some forms
almost or completely lost their stripes, and the same is true
of the bushbucks. In the arctogeal forests, boreal and
temperate, the elimination of the patterned types has gone
much further. The conclusion seems inevitable that the
patterned types in the world of to-day have no survival
value, and that their coloration at present (using “ present”’
in a geologic sense) confers no benefit upon these spotted
and striped animals when compared with the monochrome
or nearly unicolor animals, whether these latter are or are
not countershaded.

This statement about present-day conditions does not
justify us in dogmatizing about conditions in the immemorial

126 AFRICAN GAME ANIMALS

past. Mr. Pycraft (unlike Professor Poulton, and especially
unlike Mr. Thayer) fully recognizes, and insists upon, the
fact that there is no warrant for believing that these pat-
terns are due to existing factors of selection or to the action
of selective forces now at work; he explicitly states that
they are due only to the action of these transforming factors
on the generations of a remote past—although we believe
he lays unwarranted stress upon their having survived only
to the degree that they still continue to be adaptatively
perfect and efficient, for we believe that they no longer
have survival value. As to whether they formerly were
survival factors, we are in doubt; for their elimination, even
where the environment seems substantially unchanged, cer-
tainly warrants such doubt. Spencer Trotter, in his studies
in the Auk and the Journal of the Academy of Natural
Sciences, of Philadelphia, dwells on the evanescent nature
of what we speak of as “faunas,” pointing out that “a
fauna”’ is merely an expression of the temporary adjust-
ment of any group of animals to given conditions of en-
vironment—vegetation, moisture, temperature, and many
other factors combining to determine or condition the en-
vironment—so that the species which embody a fauna are
mobile elements, not hard-and-fast fixtures in their en-
vironment, their present phase of distribution being a se-
quence to their past history.

A glance at the animals of desert, or arid and semiarid
regions, is interesting from this standpoint. The general
color tone of the animals of arid regions is pallid and neu-
tral, just as the general color tone of arctic animals is white
and that of animals of temperate humid regions, such as
the neighborhood of Puget Sound, is dark. There are, of

COLORATION 127

course, exceptions to these general rules, such as the musk-
ox, raven, and wolverene in the boreal realm, or the male
ostrich, black and white chat, lark-bunting, and various
black or advertisingly colored lizards, like the Gila monster,
in arid or semiarid lands. The rule is general, however.
We do not here discuss the interesting problem of the col-
oration of arctic animals. But in the arid and humid re-
gions referred to the prevalence and repetition of pale and
dark types is almost certainly due to the varying degrees of
sunlight and moisture. A consideration of certain forms
only would make it possible to contend that the coloration
was developed by natural selection for concealing purposes.
But there are other forms, to which it is not possible to
apply this theory, which nevertheless are also colored in the
same general manner. The pale desert antelopes, such as
the desert oryx, show what is simply a more extreme de-
velopment of the coloration already tending to develop in
their several kinds on the semiarid plains; they never hide;
and their lives are led under conditions which, as we have
shown in the case of the ordinary plains game, make it
absolutely impossible that any type of coloration can have
any survival value for them. In similar fashion, the black
duck-hawk of the Puget Sound region and the pale eagle-
owl of the Western American plains show each the general
type of coloration of the faunas of their respective localities,
and yet lead lives such that this coloration can in no way
help them to secure their prey. It may, therefore, be ac-
cepted as certain that the pale desert coloration is not pro-
duced by natural selection directed toward a concealing
coloration. It is possible that there exist obscure relations
between pigmentation and bodily health, which have se-

128 AFRICAN GAME ANIMALS

cured the elimination of the darker individuals. It is
infinitely more probable that the general tone of color is
due to the action of the climate—sunlight, moisture, and
temperature—on all the individuals of a great number of
species, generation after generation, and to the inheritance
of the characteristics thus chemically or mechanically ac-
quired. We believe that this is probably a clear case of the
inheritance of acquired characteristics. We have never felt
that the demonstrations (for instance of Cope, in his “ Fac-
tors of Organic Evolution’’) of the occasional and limited
inheritance of acquired characteristics have been success-
fully refuted,* although we entirely agree with Dewar and
Finn in their masterly book on bionomics to which we have
already referred, ““The Making of Species,’ that all the
facts tend to prove that the inheritance of acquired char-
acteristics is too rare and limited a phenomenon to make
it possible to regard it, any more than natural selection,
as a primary factor in the making of species.

We saw in Africa one curious instance of the intrusion
into arid regions of a form normally belonging to an utterly
different environment. In East Africa we found the black
and white colobus monkeys in the depths of the cool, tall,
matted mountain forests. They lived in the tops of the
high trees, and came to the ground with much reluctance.
They never wandered out into the hot, dry country, where
plains of grass were dotted by scattered acacias, and palms

* Recent experiments have demonstrated conclusively the inheritance in
guinea-pigs of acquired alcoholic characteristics; subtle Weismannic theorizing as
to germ-cells and plasma-cells, and hypothetical suppositions as to the infection
of the former, whether these theories and hypotheses have a scientific or merely
a mystical value, do not and cannot alter the proved facts (1) that these char-
acteristics were acquired, and (2) that they were inherited.

COLORATION 129

and thorn-trees fringed the rare streams; although the small
greenish or yellowish monkeys were common in these trees
near the streams of this dry, hot, open country. But in
the Lado, where the country was precisely like that into
which in East Africa they never ventured, and although
there were no forests such as elsewhere constitute their
dwelling-place, we found a form of the colobus, with the
black increased relatively to the white in the pelage, living
among the thorn-trees, precisely like the ordinary monkeys,
and likewise trusting for safety to galloping off over the
ground. Even in the great forests the colobus was far more
revealingly colored than the ordinary monkey; but in this
new habitat its color was an almost startling advertisement.
Neither from the standpoint of cover nor of food, nor of
climate nor of enemies, was it to have been expected that
the colobus would so completely change its environment
and life habits.

It is necessary always to remember that it is impossible
to lay down laws regarding concealing coloration which
shall apply both to the higher and the lower animals. |The
conditions are wholly different. A caterpillar which passes
its whole larval life on green leaves may be permanently
concealed by a green color. But a sharp-shinned hawk
hunts through woods and across open fields, over snow and
Over green grass, in summer and in winter, through every
kind of landscape; and when it attacks it normally darts
with such velocity that its tint and pattern must be a mere
blur to its prey. It is evident to any serious thinker that
coloration may be all-important to the caterpillar, whereas
it must be of minor or negligible consequence to the hawk.
Again, a wood-frog or a tree-toad lives almost all its life in

130 AFRICAN GAME ANIMALS

surroundings with which its color harmonizes well-nigh
perfectly; and moreover passes much the larger portion of
its time—ninety-nine per cent of it—sitting motionless in
one position. Some frogs, if their legs are extended, will
seem to have five separate blotches of the same color; one
on each lower leg, one on each thigh, one across the back.
But when in the normal attitude there will appear merely a
broad, uninterrupted stripe across the frog’s back; it passes
so much of its time motionless, with its legs drawn up, that
the pigmentation is affected precisely as if these legs were
merely lateral developments of the body. Concealing col-
oration in such a case may really conceal. Contrast this
with the wolf, which ranges over the old and new worlds,
and, in its various subspecific forms (using the term
“species” with Linnean largeness), extends over more than
half of the land area of the globe, from the polar regions to
the tropics. We have studied the wolf carefully in the
Rockies and on the great plains; and also its smaller brother,
the coyote. A wolf is a wide-ranging animal. It hunts in
the white winter, the green spring, the gray fall; on the
open plains and in the thick woods; and amid all kinds of
surroundings of every conceivable color. Perhaps, on the
average, under such conditions a slightly countershaded
khaki or drab or gray would be most concealing; and there
are wolves of such coloration; but in company with them in
different places are white and black and red and brown
wolves, and wolves of a conspicuous iron-gray; and all are
equally successful. Evidently the coloration is to them, in
their varied lives, a factor of no survival value, and there-
fore negligible from the standpoint of natural selection;
the survival factors are cunning, ferocity, endurance, dash,

COLORATION 131

speed, the keenness of their senses, and the formidable
nature of their weapons. This also applies to the coyote,
although this small wolf does not show such advertising
color phases as its big brother. If we had only the coyote
to consider, indeed, we might be tempted to think of its
coloration as possibly due to the survival value of its vari-
ous coloration phases to the extent that natural selection
may have defined the wide limits within which other forces
can develop the various tints and patterns; but having in
view the big wolf, with its white and black color phases, it
is doubtful whether it would be correct to assign even this
small role to concealing coloration working through natural
selection. The coyote is one of the very few mammals
which it is possible that the countershading may in rare
instances help very slightly toward concealment; but it cer-
tainly is not a factor of survival value. The black-backed
jackal, when compared with the common jackal, also shows
that even an advertising coloration is of no detriment to a
beast with its habits.

The foregoing remarks are based largely on a somewhat
extended series of observations, in their native haunts, of
the following big game and carnivores:

NortH AMERICA

Cougar. White goat.

Lynx. Prongbuck.

Wolf. Moose.

Coyote. Caribou.

Red fox. Wapiti.

Gray fox. White-tail deer.

Grizzly bear. Rocky Mountain blacktail.
Black bear. Coast blacktail.

Bison. Peccary.

Bighorn.

132 AFRICAN GAME ANIMALS

Lion.

Leopard.
Cheetah.
Serval.
Spotted hyena.
Striped hyena.
Hunting dog.
Aardwolf.

Black-backed jackal.
Silver-backed jackal.

Elephant.

White rhinoceros.
Black rhinoceros.
Grévy zebra.
Common zebra.
Hippopotamus.
Wart-hog.

Bush pig.

Forest pig.
Common giraffe.
Reticulated giraffe.

East African buffalo.

Abyssinian buffalo.
Giant eland.
Common eland.
Bongo.

Bushbuck (several forms).

Koodoo.
Sitatunga.
Sable antelope.

AFRICA

Roan antelope (two subspecies).

Oryx.

Grant gazelle (three forms).

Thomson gazelle.

Gerenuk.

Impalla.

Chanler rock reedbuck.

Reedbuck (two subspecies).

Common kob.

White-eared kob.

Common waterbuck.

Defassa waterbuck (two sub-
species).

White-withered lechwi.

Abyssinian duiker (two sub-
species).

Red duiker.

Steinbok.

Oribi (two species or sub-
species).

Dikdik.

Klipspringer.

Wildebeest.

Topi.

Kongoni hartebeest.

Neumann hartebeest.

Jackson hartebeest.

Uganda hartebeest.

Nile hartebeest.

This makes a total of about eighty animals which,
because of their size, must on the whole be looked at by
hunters from about the standpoint that the big cats and
the wolves look at their prey; from the standpoint that

COLORATION 133

bobcats, foxes, minks, and weasels look at rabbits, wood-
chucks, gophers, and mice. The animals observed were
sufficiently numerous in species and individuals—of some
we must, all told, have seen many thousands under every
conceivable circumstance and surrounding—to give some
warrant in generalizing about them. We have touched
above on the coloration characters of some of them; the
characters of those omitted are similar and teach iden-
tically the same lessons.

Among these animals a fair proportion are not merely
not concealingly colored, but have a strikingly advertising
or revealing coloration. The prongbuck, white goat, black
bear, the wolf when either in the white or the black coat,
the giraffes, zebras, sable antelope, wildebeest, topi, white-
eared kob, and white-withered lechwi are so colored that
unless screened by cover it is almost impossible for them
to avoid attracting attention under normal conditions.
Many of the other animals, although not so glaringly con-
spicuous, nevertheless possess a coloration sufficiently con-
spicuous to insure their being seen by any brute or human
foe that trusts to eyesight at all—for instance, the pallah,
bongo, bushbuck, white-tail deer, and all the plains antelopes
of Africa—not one of which, from the eland, roan, and oryx
to the gazelles, ever tries to escape observation or lives
under conditions which would enable it to escape observa-
tion. There remains a minority of the grass-eating ani-
mals, which live in forests or swamps, whose coloration, if
not exactly concealing, is at least not conspicuous. These,
however, do not seem to be any better able to shift for
themselves than such of their neighbors as happen to be
advertisingly colored; the little antelopes that are foxy-red

134 AFRICAN GAME ANIMALS

do just as well as their neutral-tinted neighbors; and the
same is true of their big kinsfolk. The forest and swamp
dwellers of dull and uniform coloration and those of bright
and varied coloration get on equally well, and are equally
hard to see or kill; it is evidently the cover and the beasts’
shy, wary watchfulness, and not the coloration, that count.
Countershading has no effect at all of an appreciable quality
as regards these animals. It may very doubtfully be of
some slight help in the case of two or three of them; but
even in these it is quite impossible that it is a survival factor
operating through natural selection for purposes of con-
cealment. Ina dozen cases, ranging from the buffalo down
to the peccary, there is no countershading, and in two cases,
at least, the topi and one form of the red fox, there is inverse
countershading.

We have observed and more or less carefully studied
hundreds of species of the smaller mammals in both North
America and Eastern Africa. We have likewise observed
the birds of North America with some care, and among
African birds the waders, the birds of prey, the bigger land-
birds, and some of the more conspicuous smaller birds,
including hornbills, night-hawks, swifts, swallows, sun-
birds, bee-eaters, kingfishers, barbets, bulbuls, plantain-
eaters, parrots, weaver-birds, and whydah finches. Some
facts concerning the revealing and concealing coloration of
these species are set forth by Colonel Roosevelt in the
Bulletin of the American Museum of Natural History, vol.
XXX, pages I19 to 231.

Thousands of species of birds which are abundant and
highly successful in life, such as red-headed woodpeckers,
white-headed woodpeckers, red-winged blackbirds, yellow-

COLORATION 135

headed blackbirds, grackles generally, black and white
storks, oyster-catchers, black-bellied plovers, white egrets,
auks, puffins, guillemots, cormorants, orioles, tanagers, scar-
let ibises, flamingoes, vermilion and scissor-tailed fly-
catchers, and countless others, from the biggest to the
smallest, on land and on water, have highly advertising
colorations, and never gain any benefit from concealment.
Many of the smaller mammals, including the immense
majority of mice and shrews, lead their lives under such
circumstances as to make their different coloration pat-
terns of no consequence one way or the other in conceal-
ing them, and this although their prime hope of safety
from their foes lies in escaping observation. Habit and
cover are all-essential in their cases; and if their color-
ation in any case has been affected by natural selection,
it has merely been done by setting wide limits, beyond
which coloration cannot transgress as regards advertising
quality.

We wish to emphasize the fact that we are not now dis-
cussing the lower vertebrates and insects. From what we
have read it appears clear there are certain insects which
are protectively colored and which show that this is a sur-
vival factor by the care with which they light on objects
with which their coloration harmonizes. What is needed
to reach conclusions as regards these insects are such studies
as Doctor Henry Skinner’s “ Mimicry in Boreal American
Rhopalocera” (The Journal of the Academy of Natural
Sciences of Philadelphia, vol. XV, second series, March 21,
fanz), Dhese studies. make it evident that the extreme
positions taken in this matter by gentlemen like Professor
Poulton are not tenable.

136 AFRICAN GAME ANIMALS

As regards birds and mammals, we are in a position to
draw from our studies certain tentative conclusions.

The first and most important is that both the ultra
natural-selectionists and the ultra adherents of the doctrine
of the inheritance of acquired characteristics are almost
completely in error in their claims that they satisfactorily ex-
plain the origin of species. Mere closet theorizing is almost
worthless, and artificial experiments are insufficient. What
is needed in addition to such experiments is observation of
animals in a state of nature, carried on by trained, compe-
tent, and open-minded men, desirous of finding out the
truth. A book like Mr. Thayer’s (in spite of the artistic
value of some of his theories) 1s, we are compelled to say, in
its sum a hinderance and not a help to earnest students.
His first main contention is that concealing coloration is
universal, or nearly so, and sufficient to account as a present-
day factor for all present-day coloration. This is in flat
contradiction of the facts as regards the immense majority
of birds and mammals. His second main contention is
that the counter-gradation of shadings is the main factor in
producing concealing coloration. As a matter of fact, the
countershading of a mammal’s or bird’s coat, when it
exists, is never more than a minor factor in concealing it,
and in the immense majority of cases is a negligible factor.
It is almost certain that it is not a survival factor, at least
with regard to the higher vertebrates, and has not been
developed by natural selection working toward the produc-
tion of concealing coloration, but is due to some principle
or law, such as that at work in the vegetable world, which
tends to make leaves darker on their upper than their un-
der surfaces, and similarly affects fruits. Under certain cir-

COLORATION 137

cumstances this countershading tends toward the conceal-
ment of a few mammals and birds, but as regards the
immense majority of mammals and birds so many other
elements enter into the problem that the working of the
law is negligible from the standpoint of practical effect.
There remains the general question of concealing color-
ation as a survival factor produced through natural selec-
tion in the case of birds and mammals. The first and most
important point to remember is that, as regards the im-
mense majority of birds and mammals, the prime factors in
securing their safety are habit, if they do not trust to con-
cealment, and habit and cover if they do trust to conceal-
ment. Most landscapes in nature are so varied and ac-
cidented and contain such myriads of differently shaped
objects, such a multitude of vistas superimposed and inter-
lacing, and such a well-nigh infinite variety of colors, lights,
and shades, as to make it extremely difficult to pick out
any one object of any color. This is why a thick and high
forest, especially a tropical forest, offers such extraordinary
protection to all its inhabitants that no coloration scheme
among the adults of these inhabitants seems to be of any
consequence. This is why it is so hard in a varied moun-
tain or hill landscape to pick out a pika in a mass of slide-
rock, even though he can be heard uttering his querulous
note; or to see a marmot feeding; or a mountain-sheep ly-
ing down; or a klipspringer going about his daily business.
As regards the big game of Africa and America, the
effect of the coloration, whether concealing or revealing, is
almost or quite negligible as far as the animals’ enemies or
the other individuals of the species are concerned. Some
of them have a coloration which may legitimately be called

138 AFRICAN GAME ANIMALS ‘

concealing, others a coloration which is revealing—often
strikingly advertising—but neither seems to have any effect
upon their lives or their actions. If in cover they all try to
escape observation at times by remaining motionless, and
most of them in addition at such times crouch flat to the
ground. If on the plains, and especially when coming to
drink, they make no effort to escape observation; and all
are easily visible. The young, almost all of them, crouch
flat and motionless to escape observation, and seem to suc-
ceed equally well whether, as is generally true, they are
monotinted substantially like the parents or are spotted
in the way that it is possible the adult ancestral form was
spotted. The prongbuck uses the white hairs on the rump
in a way that looks as if designed for advertisement, and
the white tails and white rumps of some of the animals have
a marked advertising effect; it is possible that they may
serve as signal marks for the young; but if so it is to be
remarked that the young of the kinds without these signal
marks seem to thrive equally well. We believe that certain
general tints common to most of the animals of a given type
of country—the pallidity of desert forms, the lustreless
darkness of forms of a cold, humid region—are probably
due to the inheritance of characteristics acquired by the
animals because of the effect of the climatic conditions on
all the individuals, generation after generation. Aside from
this, we have found no proof that the colors and markings
are due to the inheritance of acquired characteristics. In
no case do we believe that any particular pattern or color
of the big game is due to its survival value, and, therefore,
to its production for concealing or revealing purposes by
natural selection. As regards most of these animals, the

COLORATION 139

evidence is overwhelming that their coloration, whatever
it may be, has no effect whatever as regards their foes or
prey. As regards a very few of them, it may have some
slight effect, but only to the extent that natural selection
may have set wide bounds to the variation, having no effect
on the pattern or color developed within these bounds. It
can safely be said, therefore, that natural selection has had
no effect in producing the present coloration of any of these
animals. We wish we were able to give an opinion based on
actual observations as to what the cause of the coloration
is, but at present all we can say is that no satisfactory
explanation has been given.

Among the big-game animals the countershading has no
effect; the animals without it thrive as well as those with it;
and those that seek to escape notice almost always crouch,
and then the effect of the countershading practically dis-
appears. When the white is confined to the belly of any of
these big animals it has little more effect than on the belly
of a frog or snake—where it has absolutely none and is
never seen, and therefore cannot have been developed for
concealing purposes.

We have not in this chapter more than touched on birds
and small mammals, both because we have not studied
them as we have studied the big game and because of a lack
of space. We allude to them now because the conclusions
we have drawn as to big game do not apply in anything
like such general fashion to birds and smaller mammals.
Among small mammals they apply to most members of the
squirrel family, for instance; among species with identical
habits, and often in the same species, there are found in-
stances of markedly advertising coloration and instances

140 AFRICAN GAME ANIMALS

of a coloration which, at least relatively, is concealing; and
neither type of coloration seems to have any effect on the
habits or success in life of the wearer. Evidently natural
selection has failed to find any one of these colors or pat-
terns a survival factor so far as squirrels are concerned.
The same statement applies to various other families; to
most weasels for instance. But rabbits, although when in
motion they have a highly advertising rear-end coloration,
have a concealing coloration when they crouch motionless;
and the way they lay their ears back and sink to the ground
shows that they desire to take advantage of every circum-
stance that will enable them to elude observation.

As regards all these species, the particular pattern is in
all probability not produced by natural selection; but in
most cases it certainly appears probable that natural selec-
tion has eliminated all variants of an advertising type and
all animals whose habits render them unable to take ad-
vantage of the natural coloration. The case of the little
spotted skunk, the spilogale, as observed by Doctor Hart
Merriam, is even more interesting. He writes me as
follows:

VANISHING PowER OF THE LITTLE SPOTTED SKUNKS (Spilogale)

One night in September, 1889, when sleeping deep down in
the Grand Canyon of the Colorado, in northern Arizona, near the
place where the Hance Trail was afterward built, I was awakened
at midnight by a sniffing noise near my head. Sitting up, I real-
ized that a small animal was hurrying away. After running about
twenty feet it stopped, but I could not see it until it moved; then
its color appeared to be grayish and I mistook it for a ring-tailed
civet (Bassariscus). I fired, and was surprised to find that I had
killed a little spotted skunk (Spilogale). It was so near that I
could not understand why I had not seen its white markings.

COLORATION 141

Five years later (October 15, 1904), my sister, Mrs. Florence
Merriam Bailey, and I, accompanied by Henry Gannett, of the
Geological Survey, spent a night in the Grand Canyon, near the
Cameron Trail. We had no water, and no food except some
canned corned beef, which made us so thirsty that we ate very
little. At dusk a family of little spotted skunks appeared. They
showed little fear of us, and none at all of the small fire we had
kindled. I tried the experiment of tossing them pieces of the
corned beef, which they seized greedily. They were on grayish
rocks, against which background their black bodies were easily
seen; their white markings also were rather conspicuous.

While shuffling about, searching for morsels of the corned
beef, they showed no fear so long as we stood still, but if we took
a quick step or made any sudden move they instantly vanished.
At first I thought they had gone in among the rocks, but soon saw
that they had not moved away. This struck me as so extraor-
dinary that I tried the experiment of frightening them. This I
repeated a number of times, and each time the animal or animals
nearest my feet immediately became invisible. All traces of
black and white had disappeared, and even the outline of the ani-
mal could not be made out against the uniform grayish back-
ground of rocks and sand. The sudden transformation was most
puzzling. The moon was full, but my notes are not clear as to
whether it had risen high enough to shine down into the part of
the canyon where we were; probably it had not. At all events, I
was utterly unable to see the spilogales in the act of changing
color, and could not be positive as to how they did it. I was
convinced, however, that the change from conspicuousness to in-
visibility was brought about, under the influence of excitement
or fear, by the erection of the body hairs in such manner that the
white and black markings blended to form a neutral, grayish drab,
rendering the animal completely invisible. The deception was
doubtless heightened by the solid black of the top of the head
and basal half of the tail, for at night black areas pass for shadows
or hollows—thus tending to mask the body form by cutting out
the dark parts.

In most kinds of open country, as every field naturalist knows,
grays and drabs are “obliterative” both by day and by night.

142 AFRICAN GAME ANIMALS

In camping in various parts of the West I have repeatedly ob-
served that at night a gray horse, and even more strikingly a
donkey, cannot ordinarily be seen. More than once, when stand-
ing guard or going for water, I have actually run into a donkey,
utterly unaware of its presence until I came in contact with the
body.

[t is highly probable that an owl about to pounce upon a
spilogale would be completely baffled by the transformation.

This is exceedingly interesting, from several different
standpoints. At my ranch-house on the Little Missouri,
the big black and white, unspotted skunks were unpleas-
antly common nocturnal visitors. It was always easy to
see them, and they never disappeared from sight as the little
spotted skunks do. On most nights they were the only
animals we could see at all. A badger, for instance, once
came to the ranch-house; we heard it, and thought it a
skunk; but we could not see it at all until we got a lantern,
as the neutral grayish-drab coloration was invisible by star-
light. It therefore appears that the big skunks have not
developed a concealing but, on the contrary, a highly adver-
tising coloration at night, and have not developed any way
of neutralizing its advertising character. But the evolu-
tion of the little skunks has apparently produced two con-
tradictory results: (1) a coloration pattern which is reveal-
ing at night unless the animal is excited or alarmed; (2) the
power of altering the coloration pattern under the influence
of excitement or alarm, so as to neutralize its advertising
character and make the vividly marked animal as incon-
spicuous as an ordinary animal, like a badger. Of course,
these facts, if the above interpretation of them is correct,
tend to show that in this case the coloration developed is, as

COLORATION 143

a survival factor, detrimental to the animal’s existence, and
has had to be partially neutralized by the development of a
well-nigh unique bodily habit. Equally, of course, the neu-
tralization is only partial, and apparently the animal would
have been far better protected against owls and other foes
if it had never developed a coloration which needed such
elaborate neutralization. Owls are birds of noiseless flight,
and in the vast majority of cases they must pounce on the
little skunk before it has any warning of their presence;
surely it would be far better protected if it did not have to
wait until it believed it was about to be attacked before
losing its advertising coloration. An animal like a badger,
which does not first advertise itself and then, if advised of
an emergency, neutralize the advertisement ( I do not mean
voluntarily in either case) must be in better shape than one
with the complicated characteristics and needs of the
spilogale.

Among birds there are many thousands of species, includ-
ing whole orders and families, where the coloration is usually
highly advertising, and where, nevertheless, those having
such a coloration and those, fewer in number, with a con-
cealing coloration have precisely the same habits and suc-
cess in life; so that in these cases it is evident that the
coloration is in no sense a survival factor, and has been pro-
duced without regard to natural selection. The young of
many of these species, cormorants, pelicans, and egrets, for
instance, are as advertisingly colored as the adults. But
there are a few species in which the coloration is seemingly
a survival factor as regards the adults of both sexes, and
there are very many species in which it is seemingly a survi-
val factor as regards the young, often as regards the adult

144 AFRICAN GAME ANIMALS

females, and sometimes as regards the adult males at cer-
tain seasons. There are some birds with what at first
glance does not appear to be a concealing coloration, which
nevertheless does really help in their concealment. The
hoopoo, for instance, crouching flat, with crest laid back
and wings partly outspread, obviously gains help from its
coloration in escaping the observation of hawks. Night-
hawks squatted on the ground in the daytime, whippoor-
wills on limbs, hen-pheasants, quail, and grouse on their
nests, many bay-birds or shore-snipe crouched on the beach,
desert-larks, and chats on the sand, ducklings and young
plovers, gulls, skimmers, avocets and stilts, female ducks,
and male ducks at moulting time, are among the many
birds which profit by, and by their habits show that either
deliberately or instinctively they purposely profit by, their
coloration.

It would at first seem as if in these cases the conceal-
ing coloration must have arisen through natural selection,
especially when we see the concealingly colored individ-
uals taking advantage of their coloration by crouching
motionless, and the individuals of an advertising color-
ation making no effort to hide; compare, for instance,
desert-grouse, larks, and chats, and young skimmers, and
stilts, and moulting moor-fowl, with black and white spurred
plover, black and white chats, adult skimmers and stilts,
and black cock in the fall. But more careful study and
consideration make it appear probable that the minute
patterns in these cases have not been produced by natural
selection; that natural selection has merely perpetuated
the general scheme of a useful pattern, which has been
produced by some other cause.

COLORATION 145

We must also remember those animals which, although
not concealingly colored, yet crouch and hide precisely as if
they were; the steinbok, for instance, and the white-tail
deer with its red coat showing against the green foliage of
spring and summer, crouch motionless and take the utmost
advantage of cover. It is probable that the race experi-
ence through immemorial ages has instilled into the major-
ity of the higher vertebrates the prime value as survival
factors of immobility, and of taking advantage of cover, in
order to escape observation. When other and more power-
ful factors intervene these two may, of course, lose all im-
portance and be abandoned; many animals which have not
developed concealing coloration, such as adult~ prongbuck
and white goat, wildebeest and zebra, never seek to es-
cape observation, whether by immobility or by taking ad-
vantage of cover. Nevertheless, other animals which have
not developed a genuine concealing coloration, such as the
bongo, and the wapiti with its pale, advertising rump, do
crouch behind cover and seek by immobility to escape ob-
servation. Yet other animals have developed a concealing
coloration and take advantage of it.

As regards the immense majority of birds and mammals,
habit and cover remain of such overwhelming importance
that coloration is either not a survival factor at all or else
becomes such—that is, becomes a factor telling against
survival—only if very, very striking hues are developed;
and probably only rarely under these circumstances. To
mice and shrews, immobility, night, and the actual physical
screen of earth, grass, or snow are all-important; the colora-
tion may be of any one of many tints or patterns without
producing the smallest survival effect. This is equally true

146 AFRICAN GAME ANIMALS

of small birds. In temperate North America two of the most
abundant small birds are the song-sparrow and the junco,
each in various forms; their ranges are largely the same;
their habits are alike as regards taking cover, etc.; one is
streaked and spotted, one is of a slate-like unicolor, with a
white belly; they are strikingly contrasted in coloration—as
much as are the cock and hen ostrich; and yet they are
equally successful in life. The same is true as regards in-
digo buntings and field-sparrows, seaside and sharp-tail
finches, towhees, catbirds, mockers, and thrushes. As
regards the great majority of mammals and birds, it is evi-
dent that natural selection, using concealing coloration as a
survival factor, has had nothing whatever to do with pro-
ducing their patterns and tints; and where it has worked
at all—and in many, probably in most, species it has had
no effect whatever—it has only been by setting wide limits
beyond which the variation cannot go. In all these cases
countershading has but little effect and cannot possibly be
a survival factor.

There remain the numerous cases, especially numerous
among young mammals and birds, where the coloration
has an undoubted concealing value, probably sufficient to
make it a survival factor. Although numerous, these cases
are far outnumbered by the others already discussed. In
most of these cases it is probable that natural selection has
kept the coloration concealing, but it is improbable that it
has produced either the exact pattern or the exact tint. In
a few of these cases countershading helps the concealing
quality of the coloration; but it usually ceases to act when
the animals have most need of concealment, and crouch to
avoid notice. The probabilities are overwhelming that it

COLORATION 147

is never a survival factor which operates in natural selection,
or else so very rarely that it can in practice be disregarded.
In very many cases where the young are concealingly
colored, the adults have a different and relatively a revealing
coloration; and in very many cases where the female is
concealingly colored, the male has a revealing coloration.
This means, in all probability, that the evolution has been
from a concealingly colored form to a form in which both
sexes when adults, or else the adult males, are revealingly
colored; there could be no more striking proof of the un-
importance of coloration as a survival factor, and of the fact
that it is quite impossible that in most cases among the
higher vertebrates their present adult coloration can have
been developed by natural selection. Evidently in the
varied lives led by most adult birds and mammals the im-
portance of the animal’s senses, and its truculence, wari-
ness, prowess, speed, agility, and ability to take advan-
tage of cover so far outweigh coloration that the latter
becomes negligible as a survival factor. In some cases,
however, it is a survival factor; this is especially true as
regards many young birds and brooding mothers, which of
necessity must trust to immobility for escape at critical
periods; but it is also true of old birds of both sexes in
various species of night-hawks, grouse, woodcock, etc.
Spotted fawns and young tapirs, and striped and spotted
chicks of various birds, probably indicate ancestors of such
coloration in a very remote past, although there is no in-
dication of a change of habitat. On the supposition that
these striped and spotted patterns have a concealing value,
this may mean that these ancestors were more immobile
than their present-day descendants; it may mean that the

148 AFRICAN GAME ANIMALS

increase in the brain power or sharpness of senses of the
adults has caused the coloration to become a negligible factor
in the animal’s success in life. But it must mean that the
coloration of the adults of the present day has not been pro-
duced by natural selection for purposes of concealment.

In sum, natural selection, operating with concealing
coloration as a survival factor, has played a wholly minor
part in developing the coloration of the higher vertebrates.
In the vast majority of cases it has certainly had no part in
originating the coloration; in a very large number, probably
a considerable majority, of cases it has certainly had no
effect even in preserving the coloration when produced.
It is possible, but not certain, nor even probable, that in a
very few cases it has produced the precise existing colora-
tion; it is probable that in a few cases it has preserved the
precise existing coloration after other factors have produced
it. It is well-nigh certain that in a number of cases, much
less than a majority, it has had a very real effect, not by
producing or preserving certain definite patterns or tints,
but by setting wide limits of variation which these patterns
and tints may not transgress.

As far as our observations go, in the very limited field
they cover, they support Mivart’s contention that the chief
part played by natural selection in organic evolution is the
part of a pruning-knife. As for asserting that as yet there
is warrant in dogmatizing about the origin of species, and
about evolution itself—why we are hardly on the threshold
of the matter.

CHAPTER V
GAME PRESERVES

One of the features in the psychical growth of the last
quarter of a century has been the gradual recognition by
civilized nations of their duty to protect from destruction
and extermination the various forms, including the nobler
and more imposing forms, of animal life. Thirty years ago
_ there was no effective sentiment against the destruction of
the most interesting, imposing, or beautiful birds and mam-
mals which our world contains. The bison and passenger-
pigeon in America, the white rhinoceros and quagga in South
Africa, were allowed to go the way of almost or complete
extinction; and sea-elephants, and fur-seals, gorgeous pheas-
ants and birds of paradise, beautiful egrets, ibises, and fla-
mingoes, and many, many other strange and lovely crea-
tures, were treated as the legitimate prey of every brutal
and squalid spoiler who wished to gratify his own greed or
the still baser greed of his employers and their customers by
robbing the earth of beauty and destroying what never
could be replaced. Too late to save many creatures from
destruction, but in time to save many others, the civilized
peoples waked up to what was needed. One of the first
steps taken was the establishment of the Yellowstone Park
as a game preserve by the Government of the United States.
Many other preserves now dot the American continent north

‘of the Rio Grande. The German Government has also
149

150 AFRICAN GAME ANIMALS

taken certain important steps. But it is the British Govern-
ment which has done most. Major Stevenson Hamilton’s
delightful and noteworthy book on the subject gives an idea
of the immense good that has been accomplished in South
Africa by the establishment of game preserves. In East
Africa the establishment of the game preserve between the
coast and the capital of the colony has made the railroad ride
through it an experience literally unique—an experience such
as has never been posssible for men to enjoy before our own
day. Moreover, the game laws generally in these and
other African colonies, such as Uganda and the White Nile
region, have been so observed that for the last decade there
has been no sensible diminution of the game. ‘The elephant,
buffalo, hippopotamus, eland, and the like are in the aggre-
gate as plentiful now as they were ten years back, the chief
diminution in the elephant being among the big tuskers,
while the buffalo have increased in numbers. Too much
credit cannot be given to the officials responsible for this
condition of affairs, and to men like Schillings, Sir Harry
Johnston, Buxton, and the others who by their writings and
preachings have created the public opinion which rendered
possible such action as the officials in question have taken.
Schillings’s books, and those of Harry Johnston also, are
the most effective pleas imaginable against waste and
destruction of wild life. William T. Hornaday’s volume
entitled “Our Vanishing Wild Life” should be in the hands
of every American legislator, and indeed could be read with
profit by the legislators of most other civilized countries.

It is necessary to make one point, however, in connec-
tion with the movement for the preservation of game. Our
chief task so far has been to awaken people to the need of

GAME PRESERVES 151

preserving the game, to the need of preventing the slaughter
of birds for millinery purposes and of stopping all wanton
and useless shooting of song and other birds. This is the
prime need. We need to put a stop to the entire feather
trade, excepting as regards the feathers of domestic or semi-
domestic birds, such as barnyard fowls, common _pheas-
ants, and ostriches. We need to prevent the sale of game
and to put a stop in most places to market shooting and, of
course, to every species of butchery. But the partial suc-
cess of the movement has developed another need; because
this very success developed within its ranks an enemy who
masqueraded as a friend. This enemy is the mushy emo-
tionalist, the purely hysterical creature who is under the
impression that his or her heart is soft, whereas the softness
is really merely of the head. These people need to remem-
ber that in nature animals outbreed their subsistence unless
they are kept down by natural foes. If the protection for
them is perfect, the result is that there must be some killing
off or else they will die miserably by disease and starvation.
In most places, and as regards most species, our whole
anxiety as yet is to see the animals re-establish themselves
and increase in numbers. But as regards certain species in
certain places, this result has already been achieved, and the
chief trouble is now their overincrease. The prime need at
the moment as regards the wapiti of the Yellowstone Park
and the deer of Vermont is to provide for a big additional
killing. At present, the former perish by thousands of cold
and starvation, and the latter are becoming a serious prob-
lem to the farmers. The men who protest against this kill-
ing occupy a position of unpardonable folly. In the same
way, in East Africa, it has been necessary to provide for

152 AFRICAN GAME ANIMALS

additional killing of zebras, because under protection they
became too numerous, and in Uganda, the buffalo had to
be taken off the protected list, and in various places the
hippopotamus. Moreover, the assault on collecting ani-
mals for museums is so unspeakably silly that it is difficult
to accept it as being in good faith—or, at least, it would
be, if human capacity for folly were not infinite. There
should be no collecting excepting for an adequate and pub-
lic purpose, and if species are on the verge of extinction
there should usually be no collecting at all; and purpose-
less slaughter committed under the pretense of “collect-
ing’’ should be rigorously punished. But, if these conditions
are fulfilled, it is as necessary to collect animals for museum
specimens as to kill sheep and chickens for food. It is, of
course, just as silly to object to killing wild game for
food or for scientific purposes, within reasonable limits,
as to object to killing tame animals for food or for
clothing.

There are always wicked people eager and adroit in
turning the folly of foolish people to their own base pur-
poses. When we started out on our African trip, the edi-
tors of a foolish little philanthropic publication in Boston,
nominally devoted to the welfare of dumb creatures, started
an agitation which finally took the form of petitions in the
public schools against our going. The children who signed
these petitions knew nothing, and of course appreciated
nothing, of what they were doing; probably they would
have as readily signed a petition for the hanging either of the
editors of the paper in question or of ourselves. As regards
ourselves, the action had hardly the effect of exciting even
amusement. But it had a distinctly regrettable effect in

GAME PRESERVES 153

tending to make people, in their contempt for such folly,
include in that contempt the whole movement for the pro-
tection of wild life. It is a misfortune for any cause to have
action taken which, in the public mind, identifies it with the
vagaries of mere folly. It seems incredible that grown
people, with sufficient intelligence to edit any paper, should
fail to understand that ours was merely one of the many
scientific exploring expeditions which are opening to civ-
ilized mankind a wealth of knowledge concerning the dark
continent. Practically every specimen we secured and ab-
solutely every specimen excepting a very few animals
used purely for food was preserved. With the exception
of about a dozen trophies kept for the private collections
of the members of the expedition, and with the exception
of an elephant given to the University of California, and of
a white rhinoceros group given the American Museum of
Natural History in New York, all of the many thousands of
specimens went to the National Museum at Washington.
Probably (although of this we cannot be certain) the editors
of the paper in question, who started the petition, would not
have started petitions to abolish all our museums; yet even
a moderately intelligent child of six ought to be able to see
that museums cannot exist unless collections are made for
them, and that if it is wrong to collect the specimens that
are put in a museum, it can’t possibly be right to permit
the museum to exist at all. We have no sympathy with
mere unintelligent private “collecting,”
lections”

or with any “col-
of forms of wild life as if they were postage-
stamps; and there are some animals becoming so rare that,
even for the great museums, collecting should be done with
such caution and care as not to harm the species. In

154 AFRICAN GAME ANIMALS

Africa we collected series of the white rhinoceros and whale-
headed storks, because it was of great importance to have
them well represented, for purpose of study as well as for
general interest, in our National Museum; but nothing
could have persuaded us to kill an individual of either
species wantonly or merely for the sake of a personal
trophy. The only warrant, but the ample warrant, for
collecting them was to use them in great public museums,
where they were indispensably necessary. In other words,
we collected them precisely as Mr. Hornaday, some twenty-
four years previously, had collected a score of American
bison; just as he later collected white goats, and at an
earlier date fine series of the orang-outang in Borneo,
and of the elephant and gaur in India. We mention Mr.
Hornaday, because no man in the United States has done
more than he has done in battling for the preservation of
our wild life, and because there is no man living who would
be less capable of killing any wild creature wantonly, or
without full warrant. He rendered a greatly needed serv-
ice to science when he collected the bison above men-
tioned, and when he collected his specimens in India and
Borneo; and we did the same kind of work in Africa.

We mention the attack of this little paper, the very name
of which we have forgotten, for two reasons: In the first
place, it represents the kind of folly which tends to discredit
a good cause, and therefore does real harm; in the next place,
it encourages similar attacks by sinister creatures who can-
not even give the poor excuse of good intentions for their
actions. Doubtless, the paper in question was merely silly.
But well across the continent, in Denver, there was another
paper nominally devoted, in part, to the same cause, which

GAME PRESERVES 155

was not silly at all, but which, when we came back from
Africa, repeated the same kind of attack, in the political
interest of certain representatives of what was unhealthy in
both politics and in business. The paper in question was
nominally published in the interest of worthy causes, like
protection of children and of dumb animals. In reality, it
was published in the interest of certain big financiers and
big politicians, who were violently opposed to Judge Ben
Lindsey, the judge of the Children’s Court, because Judge
Lindsey has been one of the most fearless and effective oppo-
nents of the commercialization of politics to be found in
all the United States. Mr. Roosevelt heartily championed
Judge Lindsey; thereupon the paper in question assailed
Mr. Roosevelt for having killed a cock and a hen ostrich,
and preserved their skins as well as the eggs and young birds
for the Smithsonian Institution. The paper denounced this
as an outrage on motherhood! The editor was guilty of a
similar “outrage on motherhood” whenever he ate a hen’s
egg or a spring chicken—and, moreover, in such event, he
was merely gratifying his own appetite, whereas the
ostrich group in the National Museum is for the gratifica-
tion of literally hundreds of thousands of people. It will
doubtless be hard for sensible people to believe that this
incident really occurred; but it did; and heated partisan
sheets throughout the country actually copied the article in
question.

But we must not let our contempt for the silly or sinis-
ter people who seek to twist the movement into a wrong
course cause us to forget how fundamentally necessary
the movement is. Above all, the people, as a whole, should
keep steadily in mind the fact that the preservation of both

156 AFRICAN GAME ANIMALS

game and lesser wild life—by wise general laws, by the pro-
hibition of the commercialism which destroys whole species
for the profit of a few individuals, and by the creation of
national reserves for wild life—is essentially a democratic
movement. It is a movement in the interest of the aver-
age citizen, and especially in the interest of the man of small
means. Wealthy men can keep private game preserves and
private parks in which they can see all kinds of strange and
beautiful creatures; but the ordinary men and women, and
especially those of small means, can enjoy the loveliness and
the wonder of nature, and can revel in the sight of beautiful
birds, only on terms that will permit their fellow-citizens
the like enjoyment. In other words, the people as a whole,
through the government, must protect wild life, if the peo-
ple as a whole are to enjoy it. This applies to game also.
If there are intelligent game laws honestly and efficiently
administered, then any man of sufficient skill and hardi-
hood, who can once a year get a holiday of a week or two,
will be able to enjoy healthy sport during that week or
two and kill one wapiti or one white-tailed deer or a dozen
or score of quail, or duck, or prairie-chicken, accordingly
as the legal game-bag limit permits, and accordingly as his
skill enables him to take advantage of this permission.
The ever-growing numbers of those who do not wish to
shoot, but who do love to see the wild creatures, can study
them in the most fascinating of all zoological gardens, that
is, in wild nature, wherever there are good game preserves.
Many thousands of people can travel through the Yellow-
stone Park and there see deer and antelope, sheep, elk, and
bear, in the most interesting fashion; in time they will be
able to repeat this experience along the edges of the Grand

GAME PRESERVES 157

Canyon of the Colorado; and, as long as conditions in
Africa are kept as they are now, the railway journey from
Mombasa to Nairobi will enable a man to see before his
eyes the splendid fauna which has made wild Africa through-
out historic times the marvel of the earth, and can get a
glimpse of just what a world it was upon which our beast-
like ancestors gazed in Pleistocene times.

Surely, every one ought to prize the song-birds—indeed,
all the birds of the garden, the field, and the woodland.
They are the allies of man in his fight against destructive
insects, and a knowledge of how to appreciate them adds
incalculably to the pleasure of living. Public opinion,
backing up law, should make it impossible to destroy our
bird life. Moreover, as we grow really civilized we will de-
cline to permit the existence of that base commercialism,
and the base vanity to which it panders, which would totally
destroy beautiful forms of life, whether at home or abroad,
to gratify either the epicure or the devotee of fashion. The
skins of humming-birds and the plumes of herons should
never be used for ornament, and the sale of these skins, and
the sale of any game bird or game animal which is becom-
ing rare for the purpose of food, should be forbidden by
law. There are plenty of birds with ornamental plumage
and plenty of species of game animals, just as there are
plenty of fur-bearing animals, which can be preserved or
bred under conditions that render them all alike proper
objects for consumption.

In Africa commercial hunting is the cause of the de-
struction of the ordinary game. The skin hunters, if per-
mitted to have their way, will kill every zebra and antelope,
every giraffe and buffalo in the land, and turn what was

158 AFRICAN GAME ANIMALS

a paradise into a mournful and lifeless waste. Unfortu-
nately, the settlers are often so short-sighted and improvi-
dent that they wish to go into this butchery, although their
own children would suffer from its effect more than would
any one else. The game laws are especially in their inter-
ests. I believe that in Africa this will more and more be
recognized as time goes on, just as it is now recognized in
various States in our country. In Maine, for instance,
there are more moose and deer than there were fifty years
ago; and, indeed, throughout New York and New England
the deer are far more plentiful than fifty years ago. This
is due to wise laws honestly enforced, and the citizens
living in the neighborhood are, in each case, the chief bene-
ficiaries.

Game-preserving in Africa has been so well treated by
Major Stevenson Hamilton that it is needless to go into the
subject minutely. What he says of South Africa applies
just as well to game-preserving in East Africa. As we
have said, it must not be carried to the extent of interfering
with settlement; and in many places, as the lions are killed
off, it is necessary to permit the killing of large bags of the
common game animals, such as zebras and hartebeest, in
order to prevent their excessive and detrimental multiplica-
tion. Of course, where, as sometimes happens, elephants,
buffalo, hippopotamus, and rhinoceros have grown so nu-
merous and bold as to be a menace not only to the crops
but to the lives of the inhabitants, their numbers must be
thinned. But these are exceptions. Speaking generally,
the one need is to protect the game. The game laws in
British Africa have been most wise and their enforcement
most beneficial to everybody. It is, however, very unde-

GAME PRESERVES 159

sirable to establish merely nominal game reserves so far
distant from civilization that it is impossible to enforce
the law in them. It is always bad to have a law which
honest men and good sportsmen regard as binding, while it
offers no check to the game butcher and to unconscientious
men generally. Game reserves should be established only
where there is such opportunity of supervision over them
as to offer a reasonable guarantee of genuine protection.
Moreover, the natives should not be given European weap-
ons. These weapons do them no permanent good what-
ever, and simply put a premium on the ruthless slaughter
of all forms of animal life, especially of the females and
young.

We wish to insist upon one point, which is, that the day
of the mere collector, if not past, has, at least, become of
much less relative importance than formerly. On the whole,
better work can now be done with the camera than with the
rifle. Better work still can be done by the competent and
patient observer who studies the life-histories of the great
and strange creatures of the wilderness. The joy of hunting
is entirely legitimate, and we sympathize warmly with the
man who kills dangerous game or by fair chase gets a
trophy worth having. Yet, after all, the exact size of
this trophy is a matter of unimportant personal vanity.
Whether a good pair of waterbuck or koodoo or oryx horns
measures half an inch more or less than a similar pair shot
by somebody else is supremely unimportant, whereas a bit
of first-hand observation concerning the life-history of any
species of big game is very important indeed. We sympa-
thize with and admire the big-game hunter, but the time
has passed when he can be taken very seriously if he is

160 AFRICAN GAME ANIMALS

nothing but a big-game hunter. He ought to be in addition
an out-of-doors naturalist, a faunal naturalist, or, at least,
a trustworthy and interested observer and recorder of what

he sees while in the wilderness.

CHAPTER VI
THE LION

INTRODUCTION TO FAMILY Felide (Cats)

THE existing cats form a well-circumscribed group or
family, familiar to every one as the Felide. They are very
highly specialized and farther removed from the original
carnivore stem than any of the other groups, or, putting it
more idiomatically, they are the last word in carnivorous
mammals. ‘Their specialization concerns chiefly the short-
ening of the jaws, the enlarging of the orbits to accommo-
date the large nocturnal eyes, the development of retrac-
tile claws, and the loss of the molar teeth, the function of
which has been usurped by the development of the pre-
molars into enlarged cutting teeth. The dentition has been
so reduced that the cat may be said to have but eight teeth,
the four dagger-like, tearing canine teeth and the four long,
knife-edged carnassial or last premolar teeth. The cat with
these eight teeth and its retractile claws has battled very
successfully for a place on the earth and has established itself
in every corner of the world with the exception of the polar
regions, Australia, and its neighboring islands. “Two genera
are represented in Africa, the familiar Felis and the aber-
rant genus, Acinonyx, of which the cheetah is typical.
Acinonyx is characterized by its non-retractile, dog-like
claws, abbreviated snout, high, dome-shaped skull, which

gives the head a peculiarly spherical shape, and the slender,
161

162 AFRICAN GAME ANIMALS

hound-like form of the body. The genus Felis, of which
the lion and the leopard are typical representatives, has
short, curved, and sharply pointed, retractile claws, a more
elongated head, the skull being much lower in dorsal out-
line and the snout longer and more dog-like.

THe Lion
Felis leo

The lion is quite as familiar to most persons as our com-
mon domestic animals and needs no description. We can-
not, however, include the various geographical races in this
last statement. Much diversity of opinion exists among
sportsmen as to the actual existence of the differences as-
signed by naturalists to the various described races. Some
of the hunters who have had the widest experience with
lions have observed such great color and pelage variation
among them that they refuse to accept the differences
which naturalists have pointed out in their diagnoses.
The well-known African hunter Selous is apparently one of
the leading sceptics. This lack of confidence in the exist-
ence of geographical races is quite in keeping with the broad
methods of comparison employed by even careful field ob-
servers, and has been shared by us in the hunting field. The
comparison of a large number of skins and skulls preserved
in various museums has, however, shown us the error of such
ideas. ‘The racial differences upon which the races of lions
are based are so fine that they can only be detected and ap-
preciated on the actual comparison of specimens. Our field
observations on differences usually lead us astray. The
extremes only of color, mane, size, and other characters are
kept in mind, and the average differences are wholly ob-
literated or undiscoverable to us. Age characters which
are dependent upon the examination of the skull sutures
are seldom made in the field, and much confusion results on
this account. Nothing, for instance, is more common in
the literature than the statement of the occurrence of adult
maneless lions. A careful examination of many museum
specimens, however, has failed to find really old lions,

THE LION 163

having skulls with the occipital-sphenoidal sutures anchy-
losed, associated with maneless skins. The so-called mane-
less lions are really immature specimens of adult size which
-have every appearance of being fully adult and are on that
account considered so by sportsmen. Adult size of skull
or body is not a reliable character of maturity; imma-
ture animals not only equal but occasionally exceed the
mature ones. The average observations on the coloration
of the mane are similarly misleading. Owing to the occur-
rence of black-maned lions occasionally in all the districts
where lions occur, it is assumed that the color of the mane
is of no racial value. We must not lose sight of the fact
that in geographical races we are dealing with average
characters and not absolutely distinctive ones, such as are
possessed by species. The Cape and the North African
lions are usually black-maned, while the East African is
decidedly a tawny or yellow-maned race. Black-maned lions
are occasionally seen in East Africa, but they occur in the
proportion of about one to fifty, and are of such rare occur-
rence that we are quite justified in calling the East African
a yellow-maned race.

It is a singular fact that the lion, though so widely sep-
arated by color, mane, and habits from the tiger, is almost
indistinguishable from it in skull characters. The skulls of
the two great cats are quite identical in size, shape, and
dentition, although readily distinguishable in these char-
acters from those of other members of the Felide. The
skull of the lion may be distinguished from that of the tiger
usually by the character of the nasal bones, which do not
project posteriorly beyond the end of the maxillary bones,
and by the flatter or less convex outline to the under-
surface of the lower jaw-bone. But these differences are
slight and at most only average affairs. Frequently paleon-
tologists refer fossil skulls to the lion, a proceeding in which
they are seldom justified. Fossil species based on skulls and
obviously with unknown color characters cannot be assigned
definitely to either the lion or the tiger. Large cats related
to the lion or tiger were formerly much more widely dis-
tributed than at present. They are known as far back as
the Pliocene in North America, Europe, and Asia. The
lion at the present time occurs from western India westward

164 AFRICAN GAME ANIMALS

through Persia and Arabia to Africa, where it is found
throughout the whole continent. In addition to this
range, the lion was found within historic times in Asia >
Minor and southeastern Europe. The lion has recently
been exterminated in Cape Colony, south of the Orange
River, and in Algeria and along the Mediterranean region
and Egypt generally. It is now of rare occurrence in India
and Persia.

The lions inhabiting Africa proper or Ethiopia may be
divided into some six or seven races, on differences in gen-
eral body size, relative size of teeth and skulls, and differ-
ences in tone of coloration and color of manes. At least
two of these races occur in the region covered by the pres-
ent treatise, a light-colored, yellow-maned race in British
East Africa, and a more tawny, shorter-maned race in
Uganda and the Nile Valley. Just outside of this territory
a small-toothed race, senegalensts, occurs in the Congo basin,
and another in Abyssinia, roosevelt1, having a very broad
skull and black mane. The lion occurs everywhere except
in forest or alpine meadows. He is found throughout the
dry, hot deserts wherever water sufficient to quench his
thirst can be obtained.

The lion is common throughout all the portions of East
Africa which we visited except on the high, wet plateaux and
in the dense forests; we did not come across it in Uganda;
but it was found on the Lado and less commonly along the
White Nile to the Sobat. There are geographical varieties;
but the presence or absence of the mane, and its color—
black, tawny, or mixed—represent individual and not spe-
cific or subspecific variation; black and yellow-maned lions
come from the same litter, and the fullness of the mane may
vary greatly among males from the same litter, although
it is apt to be heaviest where the climate is cold.

The litters are certainly born at various times. Judging
by the cubs we saw, one litter must have been produced by

THE LION 165

a lioness on the Kapiti Plains in January, and another on
the upper Guaso Nyiro of the north about the first of June;
and in each there were in the immediate neighborhood
of the litters of comparatively young cubs—three or four
months old—other young lions probably three or four
months older. This must mean that in East Africa litters
may be born at almost any season of the year. The lying-in
place of the lioness is sometimes in a cave, sometimes in
thick brush or long grass. Normally the cubs remain where
they were born for a few weeks, the mother leaving them to
hunt, and returning sometimes after an absence of forty-
eight hours; but they make no noise even when left thus
long. If game is abundant they may keep to the original
lair for several months, but if game is scarce, or for other
reasons, the lioness may shift her quarters when her young
ones are not much bigger than tom-cats, and the family
may then be seen travelling long distances until another
suitable place for a lair is reached. y When the cubs are three
months or so old, they habitually travel with the mother;
then, instead of eating her fill at a kill and afterward return-
ing to the cubs, the latter run up to the kill and feed at it
with their mother. We found flesh and hair in the stom-
achs of two cubs; for they begin to eat flesh long before they
stop suckling. While still very young they try, in clumsy
fashion, to kill birds and small animals. By the time they
are four or five months old they sometimes endeavor to
assist the mother when she has pulled down some game
which is not formidable, but has not killed it outright be-
fore they come up; and soon afterward they begin to try
regularly to help her in killing, and they speedily begin to
help her in hunting and to attempt to hunt for themselves.

166 AFRICAN GAME ANIMALS

Evidently in their first attempts they claw and bite their
prey everywhere; for we found carcasses of zebra and harte-
beest thus killed by family parties which were scarred all
Over.

Lions are sometimes monogamous and sometimes polyg-
amous, and there is much variety in the way they conduct
their family life. It is a common thing for an old male
to be found alone, and it is no less common for two adult
males to be found in company, living and hunting together;
the two famous man-eaters of Tsavo, which for a time put a
complete stop to the building of the Uganda Railroad, were
in the latter category. A lion and a lioness are often found
together, and in such case a strong attachment may be
shown between them, and the union be apparently perma-
nent; at least this would seem to be the case from the fact
that such pairs will often remain together just before the
birth of the cubs and while the latter are very little, the
lion lying up during the day in the neighborhood of his mate
and her litter. But it is a frequent thing to find a party of
lions consisting of one old male, of two or three or four
females, and of the cubs of some of the latter; and these
parties are well known to the Ukamba and ’Ndorobo hunt-
ers, and their association is permanent, so that these cases
evidently afford instances of polygamy. ‘Two or three lion-
esses sometimes live in companionship, with perhaps the
cubs of one or more of them; and a single lioness may be
found either by herself or with the cubs of one litter, or of
two litters. On one occasion we found a lioness associating
with a young male, not yet quite fully grown but already
much bigger than she was, and a couple of young cubs per-
haps two or three months old; now, from information given

HEAD OF THE LION IN THE GROUP BELOW
From a photograph by Dr. R. W. Shufeldt

GROUP OF EAST AFRICAN LIONS

Shot by Theodore Roosevelt on the Loita Plains, B. E. A
Mounted by G. B. Turner in the United States National Museum
From a photograph by Dr. R. W. Shufeldt

LION GROUP IN THE UNITED STATES NATIONAL MUSEUM

THE | LION 167

us by the natives, we are inclined to think (although, of
course, we are not certain) that the young male was one of
her cubs of a former litter, and the father of the cubs that
were with them. Finally, it may happen that lions join
temporarily in larger parties, which may contain two or
three adult males, several females, and young animals of
various ages; but we are inclined to believe that these asso-
ciations are short-lived, being due to peculiar conditions,
such as great local abundance of game—for lions often hunt
together in order to profit by mutual support.

Lions are noisy animals where they have not been much
molested; but, for some reason or other, if they are so
hunted that their numbers are much thinned, the survivors
seem to roar less frequently than formerly. The roaring is
done at night; but once in the Lado we heard a lion roar
after sunrise. There is no grander sound in nature than the
roaring of a troop of lions. The old male begins, and the
others chime in, at first with low moans that grow louder
and louder until the full-lunged roaring can literally be
heard for miles; then the roars gradually die away into
gasping grunts. The volume of sound is extraordinary and
cannot possibly be mistaken for any other noise if reason-
ably close; but, of course, if far enough distant it becomes
only partially audible, and may then resemble the booming
of an ostrich heard near by; and in thick cover the grunt or
growl of a lion, indistinctly heard, may be mistaken for the
grunt of a buffalo or the occasional growl—I know no other
word to describe the sound—of an elephant, a beast which
sometimes utters the queerest and most unexpected noises.
It has been asserted that the lion never roars when hungry,
because to do so would frighten his prey, and that this roar-

168 AFRICAN GAME ANIMALS

ing is a sign that he is full-fed. This sounds plausible; and
yet, as a matter of fact, we doubt if it is true. Undoubtedly,
after a successful chase lions roar freely; we have most often
heard them between midnight and morning. But we have
also heard regular roaring—not mere moaning, or the pant-
ing noise occasionally indulged in by a hungry, questing
beast—soon after dark, and this was persevered in at inter-
vals for an hour or so. We are inclined to think that gen-
erally lions are silent until they have killed, but that
occasionally, whether as signals to one another or from mere
pride and overbearing insolence, they roar at intervals on
their way through the darkness from their resting-place to
their hunting field. Of course, when they reach the actual
place where they are to hunt they become quiet; unless they
deliberately try to stampede the animals by roaring, or
unless several are hunting together, spread out around a
herd of zebra or antelope, when one may roar or grunt to
scare the animals toward the others. Ordinarily lions make
no sound that can alarm their prey; yet even when actually
hunting an occasional hungry lion may utter a kind of sigh
or moan, an eerie sound when heard close by in the pitchy
darkness. On rare occasions a lioness deprived of her cubs
or one of a pair of lions whose mate has been shot will roar
savagely after nightfall, perhaps in the neighborhood where
the loss occurred, or perhaps while travelling about. Old
males may roar again and again in answer to one another
as if challenging; and if one party begins to roar it will
often bring an answer from any lion within hearing. At
bay a lion utters a continuous growling, broken by muttered
roars; and he grunts loudly as he charges. When disturbed
a lion grunts as he gallops away.

THE LION 169

Lions do not go into heavy forests, although they make
their day lairs along the edges. They like to lie up for the
day in patches of jungle which border on open plains; in
bushes in open scrub; in clumps of reeds; in any thick bit
of cover in the open thorn forests which are so plentiful in
much of the game country; and perhaps especially in a
strip of cover along a river, or one of the dense masses of
brush and trees, of small extent, which are found along
the watercourses. They also lie in tall grass. Occasionally
they lie, throughout the day, right out in the open, on a
mound or the side of an ant-hill, or under a low bush or tree
that does not shield them from sight. If the grass is very
tall they find it easy to get close to their prey and to evade
human observation; and where the brush is thick or the
open forest fairly continuous it is almost a chance if one
comes on them. If much molested they become strictly
nocturnal; otherwise, under more natural conditions, al-
though they spend most of the day sleeping, they may some-
times be seen leisurely strolling in the open, and they often
return to their resting-places after sunrise, and leave them
before sunset—although even under such circumstances it is
only exceptionally that they hunt except under cover of
darkness. Once we came on a big male lion in mid-after-
noon walking back across the open plain to a zebra he had
killed on the previous night; and once, at the same time of
day, we came on a lioness leading her cubs back to the
carcass of a wildebeest, also slain overnight. On another
afternoon we came across a lion and lioness gazing intently
at an old bull wildebeest which was returning their stare,
very much on the alert, at a distance of sixty yards.

Except when resting and in the breeding-season, the

170 AFRICAN GAME ANIMALS

whole career of a lion may be summed up in the single
word rapine. For all the creatures of the wilderness, save
the full-grown elephant, rhinoceros, and hippopotamus, he
is the terror that stalks by night. His prowess is extraor-
dinary. His tactics are stealth, surprise, and sudden, over-
whelming fury of attack. Occasionally he hunts by day,
but in the great majority of cases by night; and the darker
the night the bolder he is and the more to be feared. If an
animal passes close to his resting-place in the daytime he
will often attack it; and in wild regions he may, if hungry,
begin to hunt early in the afternoon or continue to hunt
late in the morning; but that this is not common seems to
be shown by the fact that if lions are abroad in the
daytime, the game does not seem especially disturbed by
their proximity; hartebeests, zebras, and gazelles will keep a
watch on a lion thus moving up, and will not go very near
it, but show no special alarm or excitement. Where game
swarms and beasts of prey are abundant, and therefore
often seen, the animals that are preyed on are so constantly
exposed to assault that, although always on the watch and
often very nervous if they suspect the presence of a lion or
leopard without being able to place it exactly, they yet
grow to reckon their chances with coolness if the creature
they dread can be seen, and show a curious indifference to
the presence of the marauders if they believe themselves
safe; their moments of mad and panic terror are only when
the foe actually charges, especially if he has been hitherto
unseen. Animals of the open plain, which trust in their
speed, seem unconcerned about the presence of a lion if far
enough from him to avoid his first rush. Animals of the
bush are even more confident in his presence, or at least this

THE TION 171

is true of the smaller ones, which are adepts at dodging and
twisting through the bushes and among the tree trunks.
Once we found a reedbuck lying up in a large patch of reeds
which also contained a lioness; the two animals were spend-
ing the day in peaceful rest not fifty yards apart. On an-
other occasion we found a bushbuck at home in a thick bit
of jungle, by a small river, which jungle contained a quantity
of lion dens, although only one lion was at home at the time.
This lion made off along a dim trail, passing by the bush-
buck within ten yards; but these ten yards were filled with
small tree trunks, tough, close-growing bushes, and vines,
and the bushbuck, although much on the alert, evidently
did not think it worth while to move.

The lion’s coloration, taken as a whole, is undoubtedly
concealing. Considering all conditions, white is probably
the most conspicuous of all colors, and next to white among
mammals comes black; while a countershaded yellow dun or
dull gray is probably the least conspicuous, the most con-
cealing. Town dwellers, or unobservant persons, are some-
times surprised to learn that even at night a black animal is
ordinarily (although not always) more easily made out than
a dull-gray or khaki-colored one; but all Western cowboys
know that on the average a white horse is most conspicuous
at night, a pied horse next, and then a black horse; while
the clay-banks or yellow duns, or the dull grays, are the
hardest to see. In the old days while night herding on the
Western cattle ranches there would, of course, be nights
when I could see nothing, or when all the animals looked
alike until I was within arm’s length of them; but on the
average the colors of the horses and cattle were conspicuous
in the order above given. Donkeys, gray and counter-

172 AFRICAN GAME ANIMALS

shaded, were the hardest of all animals to see, even harder
than the only less invisible dun horses; at the time I attrib-
uted their greater invisibility purely to their inferior size,
my attention not having been drawn to the question of
countershading, which may have had some effect.

The lion’s general coloration, then, is concealing, as con-
cealing as the general body color of an eland, oryx, roan an-
telope, or buck of the big gazelle. The body coats of all
these animals have a concealing value in their ordinary sur-
roundings; and the habitual attitudes of the antelopes are
such that the countershading would have an effect were it
not that they never seek to conceal themselves and never
profit by concealment. But unlike the big grass-eaters of
the open plains, which always stand upright, the lion in-
variably squats and crouches when seeking to elude obser-
vation, so that in its case the effect of the countershading
disappears at all critical moments, and is doubtless entirely
negligible as an element in the beast’s concealment. But
this is not all. Even with a lioness the black-tipped ears
are revealing, and so is the seemingly involuntary waving
of the black-tipped tail. The male lion has some strongly
revealing bodily attributes. His mane is conspicuous, and
when it is black it has a highly revealing quality. Yet the
black-maned lions are generally beasts in high condition;
apparently neither the presence of this highly revealing
black mane in some males, nor the absence of all mane in the
females, has any effect one way or the other in helping or
hampering the animal against its prey. It is therefore evi-
dent, neither the revealing quality of the black mane nor the
concealing quality caused by its absence has any effect as a
survival factor. The slightest reflection will convince every

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EAST AFRICAN LION AND LIONESS FROM NAIROBI, B. E. A.
Presented by W. N. McMillan to the National Zoological Park

SAME MALE LION AS SHOWN ABOVE

EAST AFRICAN LIONS AT NATIONAL ZOOLOGICAL PARK

THE LION 173

one of the truth of this statement; but very few seem to
perceive its apparent meaning; for it is difficult to account
for this evident fact except by the admission that the lion’s
coloration is really a wholly minor, and probably a wholly
negligible, element in enabling it to approach its prey unper-
ceived—in other words, that the undoubtedly concealing
quality of the lion’s coloration is of interest chiefly as a
coloristic fact, and plays little real part, and probably no
part at all, in the animal’s success as a hunter, and has
not been developed by natural selection or otherwise for this
particular utilitarian purpose.

The lion sometimes lies in wait at a drinking-place, espe-
cially in seasons of drought when the water-holes are few in
number, and when the game is obliged by thirst to come to
each of them. But of the numerous kills we came across,
several hundred in number all told, only a few were by the
drinking-places. The great majority were out on the plains.
Evidently the lion far more frequently kills his game by
stalking, still-hunting, or driving on the plains than by ly-
ing in wait at a watering-place. Unquestionably a party
of lions will sometimes drive game; they spread out, and
those to one side, by grunting or merely by their smell,
stampede the game so that those on the other side may
catch it. Ordinarily, however, the lion crouches motionless
as his prey grazes toward him, or himself crawls toward it
with almost inconceivable noiselessness and stealth. The
darker the night the bolder the lion; under the bright moon-
light a lion is apt to be somewhat cautious, whereas there
is almost no limit to its daring in black, stormy weather.
No matter how pitch-dark the night, the lion seems to have
no difficulty in seizing his prey in such manner as to insure

174 AFRICAN GAME ANIMALS

its well-nigh instant death. Except full-grown elephant,
rhinoceros, and hippopotamus, there is no animal in Africa
which the lion does not attack, and it preys on the young
of all three creatures; and in altogether exceptional in-
stances parties of lions have been known to attack and
master nearly full-grown cows or half-grown young bulls
of all three of them. The giraffe is occasionally killed. In
parts of Africa the buffalo is a common prey; but where other
game is plentiful, lions prefer to avoid combat with such
formidable quarry, and they rarely attack a buffalo bull in
full vigor unless several of them are together. On Heatly’s
ranch near Nairobi lions sometimes laid up in a big papyrus
bed which sheltered a herd of buffalo, but zebra and harte-
beest and other buck swarmed near by, and during our stay
the lions never meddled with the buffalo except on one oc-
casion when a lion and a lioness together killed a young cow
which they found by itself. At Meru Boma I met a visiting
district commissioner, Mr. Pigott, who a few months pre-
viously had found the remains of a big buffalo bull which
had been attacked and overpowered by a party of lions;
the struggle had been terrific; and near by lay the body of a
lioness, her flank ripped open by one of the buffalo’s horns.
A full-grown male lion, however, will kill a cow buffalo sin-
gle-handed, and when sharp set by hunger has even been
known to kill a full-grown bull, usually after a hard struggle.
Of course, in such a case the lion owes his success to surprise,
the attack being delivered with terrific rapidity, and the
quarry taken completely unawares. Even a cow buffalo, if
on her guard, would have a good chance of beating off a
lion, and a bull would almost certainly do so. But if the
lion can bound on his victim, fixing the claws of one fore-

THE LION 175

paw on its face or muzzle, while the other holds it by the
shoulder and the great fangs tear at the neck, feeling for the
bone, he is very apt to win. In sucha case the buffalo is so
hampered that it cannot exert its full strength and, with its
head twisted to one side, there is a fair chance of its break-
ing its neck in one of its headlong plunges; and unless it
can shake off the lion sooner or later the latter’s teeth meet
through the spinal marrow and the fight is over. When
several lions attack jointly they apparently interfere with
one another, or else embolden one another so much that the
quarry is less scientifically seized, and is usually clawed and
bitten all over. Probably lions occasionally strike heavy
blows with their massive, powerful forearms, but this is
certainly not common; ordinarily the claws are merely used
to hold the animal, and the killing is done with the teeth.
Thick brush, and, to an even greater degree, long grass,
favor a lion’s attack, enabling him to make his rush so close
up that the prey has little chance of escape; but on a bare
plain the game may get just the second’s time necessary to
escape, and if it is a big, powerful, even though unwarlike,
animal, like an eland, it may wrench itself free from a bad
hold where its head or neck has not been seized, and escape.

The great majority of the kills that we saw were zebras
and hartebeests; but we also came on the carcasses of eland,
wildebeest, oryx, waterbuck, wart-hog, kob, impalla, and
gazelle, which had fallen victims. Usually it was impos-
sible to tell just how the killing had been done; twice we
found zebras with the big fang marks on the back of the
neck; we found a hartebeest which had been seized by the
throat; several animals showed claw marks on their faces;
a young waterbuck cow had been bitten through the head

176 AFRICAN GAME ANIMALS

—we think, but of course cannot be certain, that this was
the way she was killed. With none of these game, not even
the plucky oryx and wildebeest, did we find the slightest
evidence of resistance or of anything in the nature of a fight.
The oryx is said at times to defend itself against the lion;
but although along the Guaso Nyiro of the north we came
on a number of oryx kills, we never saw one instance in which
there had been more than the usual scuffle. Evidently the
surprise had been complete, and the sudden rush and im-
mense strength and ferocity of the assailant had left no
time for resistance. Young and clumsy lions, or very old
and weak lions with poor teeth, may occasionally meet with
misadventures in tackling a big zebra or antelope; but in
East and Middle Africa, of the animals habitually preyed on
by full-grown lions in their prime, only the buffalo are dan-
gerous to them. Lions kill camels, horses, donkeys, cattle,
sheep, goat, and tame ostriches; but where zebra and harte-
beest swarm, as in much of East Africa, they find it so easy
to catch them that, as a rule, they do not seriously interfere
with the flocks and herds of the natives and settlers. But
even under these circumstances an occasional lion will take
to cattle-killing and to ravaging the flocks of tame ostriches.
Ordinarily in killing their prey, lions keep their hind feet on
the ground; but in one case where a horse was killed the lion
leaped on its back; the hoof-prints showed that the horse had
run for sixty yards or so before falling, and during this dis-
tance not a paw of the lion had touched earth. The marks
on the dead horse showed that the lion’s hind claws had dug
deep into the haunches, while the fore-paws grasped the
shoulders, and the teeth bit into the neck until the bone was
crushed.

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Vudaz V NO MOVILV S.NOIT V 40 GOHLAW

THE LION Lit

Man-eating lions have always been fairly common in
East Africa. The most noted, but far from exceptional,
case was that of the two man-eaters which for a time
stopped the building of the Uganda Railroad by their rav-
ages among the workmen, until they were finally shot by
the engineer in charge, Mr. Patterson. Another lion, after
killing several men around a station on the railroad, carried
off and ate the superintendent of the division; the latter had
come down in his private car, which was run on a siding,
and he sat up at a window that night to watch for the lion;
but he fell asleep and the lion climbed on the platform,
entered the car by the door, and carried off his would-be
slayer through the window. In the summer of I909 a
couple of man-eating lions took to infesting the Masai vil-
lages on the plain around the headwaters of the Guaso
Nyiro, west of Kenia, and by their ravages forced the Masai
to abandon the district, and the native travel routes across
it were also temporarily closed. A few weeks later we were
hunting in the district; we kept the thorn boma round our
camp closed at night, with a fire burning and askaris on
guard, and were not molested. Near Machakos Boma a
white traveller was taken out of his tent by a man-eater one
night a good many years ago; a grewsome feature of the
incident was that on its first attempt the lion was driven
off, after having seized and wounded its victim; the wounds
of the latter were dressed, and he was again put to bed, but
soon after he had been left alone the lion again forced its
way into the tent and this time carried the man off and ate
him. Every year in East Africa natives are carried off from
their villages or from hunting camps by man-eating lions.
Occasionally one hears of man-eating leopards, which usu-

178 AFRICAN GAME ANIMALS

ally confine themselves to women and children, and there
are man-eating hyenas; but the true man-eaters of Africa are
lions and crocodiles. As has long been known, man-eating
lions are frequently very old individuals, males or females,
which have lost many teeth, and are growing too feeble to
catch game, whereas they find it easy to master man, who
is the feeblest of all animals of his size, and the one whose
senses are dullest, and who has no natural weapons. But
it is a mistake to think that all man-eaters are old and feeble
animals. Where lions are much hunted it is doubtless true
that they grow so wary of man that only the dire want pro-
duced by utter feebleness could make them think of preying
on him; but where they are less molested, their natural
ferocity and boldness make it always possible that under
favorable conditions a hungry lion, not hitherto a man-
eater, will be tempted to kill and devour a man, and will
then take to man-killing as a steady pursuit. Many noted
man-eaters—those killed by Mr. Patterson, for instance—
have been full-grown male lions in the prime of life and
vigor. It is worth noting, by the way, that Mr. Patterson
states that in advancing to the attack on the railway camps
the lions always roared loudly to one another until within a
mile or so, but were absolutely silent during the hour that
preceded the actual seizure of some unfortunate man. The
attack of a man-eater is always delivered at night, and is
practically always by surprise; but if the first attack fails, a
bold lion will sometimes persevere and do his best to seize
another victim. Even a man-eater thoroughly realizes that
men are dangerous; he is no more apt to make a hard fight
when himself hunted than is any other lion of like vigor;
and when he is the hunter he always retires with his vic-

THRE: LION 179

tim, as soon as he has caught it, out of reach of vengeance,
although he may only go for a distance of a few hundred
yards, being confident in the shelter yielded by a dark night.
This is entirely unlike the lion’s conduct with other prey; if
a zebra or hartebeest is killed, the lion stays on the spot with
his victim, and may eat it where it has fallen or drag it a
few yards to a more convenient spot. Save in very wild
places the lion leaves its prey at or before dawn, and may
then travel some miles to its resting-place; it will probably
come back the second night, unless it has been molested or
has had its suspicions aroused. Lions feast on any dead
animal they find, from an elephant to a dikdik, and even
eat carrion. When they kill game of any size they first
neatly disembowel the body, usually burying the entrails,
and then either eat the heart, lungs, and brisket, or else
begin straightway at the hind quarters. Sometimes, how-
ever, they do not disembowel the prey. They feed greedily,
bolting strips of the hairy hide with their meat, but in one
case I heard from an eye-witness of a lion’s striking a dead
zebra’s body with its spread paws, and clawing off big
patches of hide before beginning its feast.

Except man, the lion has few or no regular enemies in
his prime. He will get out of the way of either an elephant
or rhinoceros, and a herd of cow elephants with calves will
‘charge any lion they find lurking in their neighborhood, and
would undoubtedly kill it if it could not get into cover.
Very probably a buffalo herd might, under exceptional cir-
cumstances, behave in the same way. Hyenas hang around
lions to get the offal of any beasts the latter slay; and it is
no uncommon thing to find the body of a hyena which has
itself been slain by a lion when it has incautiously or over-

180 AFRICAN GAME ANIMALS

boldly ventured too near the killer’s feast. Ordinarily a
vigorous lion has nothing to fear from hyenas, and pays no
heed to them; but hyenas are powerful brutes and, in spite
of being normally abject cowards, they possess a latent
ferocity which when they are in sufficient numbers at times
renders them foes to be feared. Once Donaldson Smith,
watching from a thorn boma at night, saw a regular pitched
battle between several lions and a big mob of hyenas, in
which the latter got rather the best of it. On one occasion,
while lying near an elephant carcass, with Carl Akeley, I
heard the hyenas which had been feeding on it throughout
the night become roused to a fury of noisy defiance by a
lion which approached the dead elephant a little before
dawn, uttering the moaning sighs so characteristic of a hun-
gry lion; the hyenas yelled, screeched, growled, laughed, and
cackled, and apparently actually bluffed the lion, which did
not venture to come to close quarters. Moreover, I am in-
clined to think that very old and feeble or badly wounded
lions find their normal ends in the maws of hyenas. On
one occasion Lord Delamere and one of his Somalis were
desperately wounded by a lion he had attacked. They had
to camp directly where the accident had occurred, and the
lion, also very badly wounded, lay in the bushes but a couple
of hundred yards away. Soon after nightfall the hyenas
began to gather round the wounded lion, and eventually
attacked it; the lion roared and fought fiercely, and a long
battle ensued, but in the end he was overcome and eaten.
The hunting-dogs or wild hounds also attack lions, and are
much bolder in doing so than are the hyenas. Mr. Paul
Rainey saw a small pack of them harrying a big lion, which
was skulking off in much alarm; whether the wild hounds

THE: LION 181

ever actually kill a lion we cannot say, but they certainly
sometimes harass and cow them and force them to slink off
in terror. Lions have been known to eat leopards. With
the other man-eaters, the crocodiles, relations are more
equal. Unquestionably, lions have on rare occasions been
seized, dragged under, and drowned by big crocodiles; a big,
hungry crocodile accustomed to seizing animals while they
are drinking at the water’s edge would seize a lion as soon
as an ox. But the lion also at times kills crocodiles, catch-
ing them while they are lying on the shore. A trustworthy
elephant hunter whom we know writes that in the neighbor-
hood of Lake Rudolf he found a number of skeletons of croco-
diles which had evidently been killed by lions; and the na-
tives assured him that at this point lions frequently pounced
on and ate crocodiles. Doubtless the lion, like other ani-
mals, varies in character and habits from place to place; and
if by any chance a single lion in some particular locality
learns how to prey on an animal not ordinarily attacked,
other lions may readily learn to follow his example. At any
rate, it sometimes happens that lions in one district as com-
pared with those in another district have entirely different
customs as to what game they prey on; as I| have said above,
in some places they make buffalo their especial quarry,
whereas in others they never molest the formidable wild cat-
tle if zebra and hartebeest are to be obtained. This is like
the American grizzly bears, which in some localities feast
on carrion, and in others attack cattle at times and kill
game; whereas in yet other localities they never kill any-
thing larger than a gopher, and pay no attention to the car-
cass of a dead animal.

It is unsafe with any animal, and especially with an ani-

182 AFRICAN GAME ANIMALS

mal of such high and varied development as a lion, to try to
lay down invariable rules. Speaking generally, lions are bold
and dangerous to men at night, and exactly in proportion to
the darkness of the night; on black and stormy nights a lion’s
daring is sometimes almost incredible. Even where there
are no confirmed man-eaters a solitary man is never safe
after nightfall if walking through a country where lions are
numerous; if one is encountered it will probably get out of
the way, but there is always the chance that it will attack.
For instance, in photographing lions with the flash-light at
night, there is always danger in going out to reset the flash-
light; if a lion is in the immediate neighborhood, it is always
possible that it may charge. Mr. Coolidge, of Boston, is
one among several men we have known who have had such
experiences. In daytime there is usually no danger whatever
in meeting unwounded and unharassed lions, unless they are
actually stumbled on in thick cover; they may growl or
stand and stare ominously, but if unmolested they will
almost always retire. But two of the friends we met in
Africa, Mr. Percival, the game ranger, and Mr. Harold Hill,
have had uncomfortable times with lions which they met in
broad day. Mr. Percival was riding through some fairly tall
grass when a lion charged him and soon overtook the gallop-
ing, frightened horse; Percival leaned over and yelled in the
lion’s face; thereupon it drew up, but again came on; the horse
was now at full speed, and reached the open plain of short
grass ahead of the lion, which then halted. Probably this
lion at first mistook the horse for a zebra, and it halted when
it discovered it was pursuing a man; yet it again came on,
and would probably have overthrown both horse and rider
if the stretch of tall grass had not come to an end. Mr. Hill,

THE LION 183

while unarmed and walking along the wagon trail to his
house, in the middle of the day, suddenly came on several
lions in the middle of the road; they lay watching him and
declined to move; so he walked off to one side and round
them; whereupon they rose and followed him nearly to his
house, slinking along through the patches of thin cover.
They did not attack him; but they were certainly not fol-
lowing him with friendly motives; and the walk home with
such attendants was distinctly uncanny.

When hunted, the lion certainly stands high in the cate-
gory of dangerous game. There has been endless discussion
and endless variety of opinion among experienced hunters
as to which animal is the most dangerous. Nor is this dis-
cussion confined to African game; the same diversity of
opinion as to the relative danger of hunting on foot the tiger,
the Indian elephant, and the various wild oxen, such as the
buffalo, obtains among men who have hunted in Asia; and
this is likewise true when we consider the grizzly bear and
jaguar. Enough attention is not paid to the wide differ-
ences of character among individual animals of the same
species—just as among individual men; and moreover, the
surroundings under which one kind of animal is dangerous
may be the very surroundings under which another, nor-
mally more dangerous, would be less dangerous. Ona level,
open plain a fighting buffalo bull, or even a fighting tusker
—elephant—would be rather more dangerous than a lion or
leopard, because harder to stop—at least the buffalo would
certainly be harder to kill, turn, or stop than either of the
big cats, and the tusker very much harder to kill or cripple,
although perhaps easier to stop or turn. On the other
hand, the buffalo would be less apt to charge than the ele-

184 AFRICAN GAME ANIMALS

phant, and very much less apt to charge than either the
lion or leopard. In most kinds of cover, however, the lion
or leopard would, from every standpoint, be more dangerous
than the buffalo or elephant, unless it was cover that would
not hamper either of these big animals and would hamper
anything smaller. There is thus every allowance to be
made for changed conditions and for the wide variation of
temperament among individuals. Moreover, there can be
no doubt that in certain localities all the animals of a given
species show an unusual development of certain traits com-
pared to the aggregate of individuals dwelling in another
locality; and so in one locality the buffaloes, elephants,
rhinos, leopards, or lions may be much more dangerous
than in another locality.

Taking the average of individuals and the average of
surrounding circumstances, we consider the lion more dan-
gerous to the hunter than any other game. The leopard is
an even greater adept at hiding, is even quicker and more
reckless in its charge, and is smaller and more difficult
to hit; therefore, we consider that a fighting leopard is a
trifle more likely than a lion to get home when it charges;
but it is so much smaller and less powerful that it is far less
likely to kill its antagonist—very few hunters have been
killed, although many have been mauled, by leopards—and
a few good dogs will not only stop but themselves kill a
leopard, so that with dogs it can be hunted with entire
impunity. Under many, perhaps most, conditions, a fight-
ing bull buffalo offers a rather harder proposition than a
fighting lion, because the huge, massive creature is far more
difficult to stop when he does charge, and because he usually
shows the most vindictive and ferocious resolution when,

THE LION 185

after being wounded, he finally makes up his mind to fight;
but a buffalo is not nearly as apt to charge as a lion, and is
far more easily seen and far less quick in his movements,
and in most cover is more easily evaded; so that on an aver-
age we do not consider that the buffalo is as dangerous.
Elephants are much more apt to charge than buffaloes; a
herd will charge under circumstances when even lions would
slink quietly off; and in consequence to hunt them seems to
us about as dangerous as to hunt buffalo; but the actual
charge of a single elephant does not seem as dangerous as the
actual charge of a lion or buffalo. The rhinoceros we regard
as on the average much less dangerous than lion, leopard, ele-
phant, or buffalo; in fact, as only about as dangerous as,
or very little more dangerous than, the grizzly bear. But
it must be remembered that these are only our personal
views; and while we believe that they are shared by the
majority of the big-game hunters most competent to speak
on the subject, we are well aware that many equally experi-
enced and observing hunters are on record as expressing
widely different views. Good authority can be produced
for the statement that the buffalo, or the elephant, or even
the rhinoceros, is the most dangerous, and the lion or the
leopard the least dangerous; and different observers have
arranged the five animals in every conceivable order, as ad-
versaries. The truth is that in this matter there is such
wide, individual variation, both among the hunters and the
hunted, that every general statement must be made with
full knowledge of the many exceptions that exist thereto.
There are circumstances or occasions under which the chase
of each of the five animals—six including the white rhi-
noceros—may be very dangerous; while under other con-

186 AFRICAN GAME ANIMALS

ditions a number of each kind may be killed with the mini-
mum of risk. They are the only kinds of African game the
chase of which can ever be properly described as dangerous.
A hippopotamus, however, will sometimes charge a boat,
both unprovoked and when it has been wounded; and of
course many other animals in Africa, as elsewhere, will show
fight if cornered and rashly approached too close. The
roan antelope when wounded will charge savagely from
some little distance, and is then more dangerous than any
American animal except the grizzly bear, and more dan-
gerous than any European animal whatever, with the
possible exception of the brown bear; the sable is almost
as dangerous; then comes the oryx; then the wildebeest
or gnu. Probably none of the deer of northern lands are
as dangerous as any of these. The bushbuck, in spite
of its small size, and the koodoo, waterbuck, hartebeest,
and also the zebra and wart-hog, will turn at bay, but we
consider them as formidable only in the sense that a bull
moose or wapiti is formidable; that is, under all normal con-
ditions the element of danger in their chase is entirely neg-
ligible. Eland and giraffe are exceptionally mild-tempered;
yet the giraffe will drive both the big zebra and the oryx
from a water-hole, although giving way to rhinoceros and
elephants.

The chase of the lion, if fairly followed, is an enthralling,
but certainly a dangerous pastime. Of course, a few lions
may be killed under such circumstances that the hunter is
practically in no danger whatever; and in certain forms of
hunting lions, such as sitting in a tree or a high thorn zariba
and waiting for them to approach a carcass or a tethered
domestic animal, the element of danger is eliminated.

THE LION 187

Moreover, the experience of Mr. Paul Rainey, who, in com-
pany with Mr. Heller, took to Africa a pack of American
bearhounds and fighting dogs, and by their aid killed about
sixty lions, shows that with thoroughly trained dogs of the
right temper lions and leopards can be pursued by good
hunters with the minimum of risk and almost the certainty
of success. But the man who on foot or on horseback, with-
out dogs, follows lion hunting as a steady pursuit will not
kill many of them without being viciously and resolutely
charged, and if he persists long enough his life will be put
in jeopardy. During the last twenty years scores of white
hunters have been killed by lions, elephants, buffaloes, and
rhinos in East and Middle Africa; and the lions have killed
much more than half of the total number. Except Mr.
Rainey, who worked with dogs, Lord Delamere has killed
more lions than any other man we know—fifty-three; he
was badly mauled on one occasion, and has now given up
hunting them, stating that no man can count on killing more
than fifty lions without himself being killed or fatally in-
jured. Mr. Roosevelt and Kermit killed only seventeen
lions between them, two being cubs and two not much
more than half-grown; and thirteen full-grown lions are too
few to permit of free generalization as to their fighting
capacity. Three of these thirteen lions—two big-maned
males and a lioness—charged with the utmost resolution,
from a distance of nearly two hundred yards when wounded
and brought to bay by the pursuing horsemen; three others
(all male lions) were at bay and were about to charge—one
had begun to trot forward—when killed; five were killed or
disabled under circumstances that gave them no opportu-
nity to charge; two (both lionesses) were killed close up, after

188 AFRICAN GAME ANIMALS

being wounded, under circumstances which seemed to invite
a charge, yet they made no effort to charge. Only one other
lion was shot by any other member of our party—a lioness
killed by Alden Loring; she charged with the utmost reso-
lution when mortally wounded, and died while still charging.
This short experience, taken by itself, would tend to show
that a full-grown male lion is rather more apt to charge,
and is rather more resolute in charging, than is likely to be
the case with a full-grown lioness (save where the latter has
young cubs); but it is impossible to generalize on such in-
sufficient data. Tarlton, who was with us, a noted lion
hunter of long experience, was inclined to think that on the
average the lioness was a little more apt than the lion to
charge. Probably the difference in this respect between the
sexes is not great, while the amount of variations among the
individuals of each sex is very great indeed. A beginner °
might readily kill three or four lions without danger; and he
might be charged and killed by the first one he attacked.
If the sport is persevered in, the man who achieves success
must possess coolness, wariness, resolution, and reasonable
skill with the rifle; and now and then he will need to show
all these qualities.

Lions can be hunted in many ways, aside from lying in
wait for them at night and from hunting them with dogs.
The three most satisfactory ways are to trail them in the
early morning, just after they have begun their return to
their day lairs, to drive them out of thick cover by a line of
beaters toward the previously stationed hunter, and to gal-
lop them down on horseback, usually after having roused
them from the shelter of some patch of bush, trees, reeds, or

THE LION 189

long grass in which they have been lying. Often, moreover,
especially just after sunrise or before sunset, alert and ener-
getic hunters will stumble on them by accident or run across
them when they are just ending or beginning their night
rambles. There is luck in all hunting; but more in lion hunt-
ing without dogs than in the chase of any other big animal
of the regions where it dwells, for the lion is far harder to
find and see than any of the big grass-eaters that live under
similar conditions. One man may never seea lion ina year’s
hunting; another in the same time may come across a couple
of score.

In both tracking and driving the hunter is, of course, on
foot. Tracking can only occasionally be employed with
lions, as their spoor is so much more difficult to follow than
that of hoofed animals, and as they are experts at hiding and
skulking. But it can occasionally be employed in the early
morning when the dew is heavy on the grass; and in long
grass it often pays to follow a band of lions for some time,
as they make fairly distinct trails, and frequently move
slowly off before the hunter, grunting now and then. A
shot at an unwounded animal under these conditions is
not very dangerous, for the animal has not been harassed or
injured, and rarely charges until hit; while the shot is usu-
ally taken at a distance that ought to enable the shooter
to kill or cripple his game. Driving is followed in the usual
fashion; a rocky hill, a valley of long grass, a reed bed, a
fringe of trees, or.a patch of scrub being beaten through by
a line of shouting natives. The lion rarely charges back at
the beaters, although it is always well to have a man with a
rifle accompany them. The hunter himself in these cases
is very rarely charged before shooting, for the lion has

190 AFRICAN GAME ANIMALS

not been baited and does not feel that it is cornered. But,
of course, as the hunter is on the ground, perhaps seventy or
eighty yards from the lion, it behooves him to use straight
powder.

Riding lions, as it is termed—that is, running them on
horseback until they turn to bay, and then shooting them
on foot—is a more exhilarating, a less fatiguing, and, on the
whole, a more dangerous sport. Usually two or more
mounted hunters go out together. They may spy a lion in
the open, or they may rouse him from his day lair. In
either event, they run him hard over the open plains until he
comes to bay—either on the bare open or ina bush. When
he is thus brought to bay he is far more apt to charge than
is the case with an unwounded lion shot by a man on foot.
In any such case, and of course especially in the case of a
wounded lion or of one suddenly surprised at very close
quarters, if the lion is approached too close it may charge
without any warning. If a man on horseback gallops too
close behind a fleeing lion, it may whip round and charge
him without a moment’s pause. But when brought to bay,
and when the hunter is some distance off, the lion usually
spends some little time in threatening and in working itself
up to the final pitch of fury. It stands erect, the head held
lower than the shoulders, the tail lashing from side to side,
and all the time it growls hoarsely, the lips drawn down
over the teeth like those of an angry bear, or, more rarely,
drawn back in a prodigious snarl. When just on the point
of charging, the tail is usually thrown stiffly up two or three
times, and if it changes its course during the charge the tail
is slued to one side like a rudder. The animal may break
into a gallop at once, or it may begin by trotting with the

THE LION 191

tail erect. One of the lionesses that charged us came on
with occasional great bounds; but all the other lions gal-
loped like huge dogs. The pace is very rapid for one or
two hundred yards. A horse which is standing but a hun-
dred yards distant may be caught before it has time to get
into a full gallop. Usually the lion, when it does charge,
charges with the utmost determination; and, as we have
said, it is more apt to charge when brought to bay on horse-
back than in other kinds of hunting; but it is also compara-
tively easy to kill under these conditions, for it advances
from some distance in open country upon a man fully pre-
pared and expectant. Moreover, it is much more easily
killed or crippled than is the case with the bigger kinds
of dangerous game. A fair shot who is cool-headed and
has a good rifle ought under these conditions to be reason-
ably certain of stopping the lion before it can get to close
quarters. Occasionally, if a man stands stock-still, even with
an empty gun, a lion after running straight at him will at
the last moment swerve; this is not ordinarily true, however;
but if of two men together one runs when the lion is close,
it will usually seize the runner. When it comes to close
quarters it may rear and strike with its fore-paws, but far
more often runs in on all fours, like a dog, knocks the man
down as it seizes him, and then lies on him, using the claws
to hold him, and doing the killing with its great fangs. If it
seizes him by the throat or head he is killed instantly; but
in the hurly-burly of the mellay the beast seems to lose
somewhat of its “‘instinct for the jugular” and bites re-
peatedly at any part that is nearest—arm, leg, side, or chest.
In consequence, if help is at hand, the lion can usually be
killed or driven off before it has killed the man; although the

192 AFRICAN GAME ANIMALS

latter may die of his wounds later. While in East Africa
we met many more men who had been badly bitten and
clawed by lions, but who had recovered, than we heard of
men who had been killed by them, or who had died of their
wounds. The wounds should be cauterized at once, to
avoid blood-poisoning, as the lion’s teeth and claws often
seem to contain some poisonous element, perhaps minute
particles of dead matter. Moreover, the lion often bites
deep, and with closed jaws pulls the muscle loose from the
bone, thus causing the deep-seated tears which become
sources of corruption. Finally, the shock of the bite is tre-
mendous, the full muscular power of the great jaws being
brought into play. Many men die from the shock. On the
other hand, it so numbs them that at the moment little
pain is felt. We sawa dozen men who had been bitten. One,
on whom the lion had lain for some time while biting him,
had suffered much; all the others assured us that at the time
and for several minutes afterward they did not suffer at
all. The process of healing is long and painful. Most of the
men who die from lion wounds die from blood-poisoning
several days after being mauled, and not directly from the
mauling.

The hunter should never go near a lion until it is dead,
and even when it is on the point of death he should not
stand near, nor approach, its head from in front; for a lion
at its last gasp will summon all its energies for one final
attack, flinging itself on the man who has thus incautiously
approached it, especially if it can see him, and spending its
last dying moments in biting him. Necessarily, lion hunt-
ing has elements of danger if legitimately followed; for a
slight deviation in aim—possible enough to any marksman

THE LION 193

when the target is coming on ata gallop—may mean a
mauling; but most accidents occur because of some bit of
carelessness or recklessness, some lack of caution or prepa-
ration, of the kind that ordinarily brings no retribution, but
which it is always possible will invite disaster.

The fault may lie in the overeagerness and ignorance
of a beginner; it may be due to lack of nerve or timidity—
and sometimes vanity will induce timid men to venture into
a field for which they are totally unfit; or the mischance
may occur to a keen, experienced hunter whose skill, hardi-
hood, and prowess have finally led him to feel an unwar-
ranted contempt for the game. ‘This last was the cause of
the lamented death of George Grey, the brother of Sir Ed-
ward Grey, the British Minister of Foreign Affairs, who
was killed by a lion the year after we left East Africa, while
out hunting with Sir Alfred Pease and Mr. Harold Hill,
the two men in whose company I killed my first six lions,
and not more than half a mile from where my first big
male lion was killed. We had met George Grey in Africa
and again in London; he was a singularly fine type of
man, very modest, utterly fearless, as hard as nails, and
probably as good a man in a serious emergency as was to be
found in all Africa. Shortly after Kermit killed his two
bongo, a cow and a yearling, Grey killed a bongo bull,
which he gave to us to complete our group for the National
Museum. He had done much hunting of dangerous game
and was so fearless and possessed such prowess that he un-
derestimated their dangerous qualities; elephants he held
in some slight respect, but we heard him say that he did not
regard buffalo as more dangerous than tame cattle, and he
cared but little for lion, and nothing for rhinoceros.

194 AFRICAN GAME ANIMALS

Pease, who was out with him, is an exceptionally good
hunter, a fine rider and shot, very hardy and cool. He has
killed very many lions, a number of them under circum-
stances of much hazard; and, being a close observer, he has
been struck by the wide variety of conduct among them,
and the occasional great danger from them; and he com-
bines to an unusual degree extreme boldness with very
good judgment. After we left him in East Africa, he wrote
describing two lion hunts in which he had taken part, Harold
Hill and his brother Clifford being along. In one a lioness
literally behaved like a cur; she got into a thicket and would
not go out until he fired buckshot into it, and even then,
although slightly wounded, she ran like a rabbit and was
killed like one. In the other hunt, a big-maned lion fought
with savage ferocity, charging the mounted men again and
again, almost escaping, and finally being killed in full charge
at the distance of a few feet by the rifles of his three pur-
suers. Pease then went back to England, but returned to
Mombasa early in 1911 in company with George Grey, who
went with him expressly to try riding lions on the Kapiti
Plains. The fatal hunt occurred on the 29th of January.
In a letter Pease writes of it as follows:

During the voyage Grey often talked of lion hunting and
told me he had never seen lions ‘“‘ridden”’ and bailed up, the
method I had found so successful for many years. I explained to
him on several occasions what was required and how it was done
and over and over again impressed on him certain rules, which if
observed reduced danger to a minimum. The rule I pressed
most was when galloping a lion never to get within 200 yards of
one and that it must never be forgotten that no horse can get
away from a lion with only too yards start: but that with a fast
handy sure footed pony on sound ground a lion could not catch

THE LION | 195

you if you had 200 yards start of him. I told him he must have
a good steady fast pony, and he cabled to Nairobi while en route
to have one sent to my place. I told him that unless the ground
was very sound and open I kept off lions (when ‘“‘riding”’ them) at
least 300 yards and that I never went anywhere near them on
rough cracked and broken ground where a pony could not gal-
lop, nor into high grass. I also told him that when we bailed up
a lion we always waited, at a safe distance, till the others came up
and our gunbearers with heavier and more powerful weapons: and
that whilst riding the lions we carried handy rifles, I myself always
a .256 with which I might shoot at long range but never at close
quarters. He often talked of the degree of danger in hunting
dangerous game; he did not think there was much danger with
buffalo saying that he had shot scores of them and he had
found no more danger than there would be in shooting cows: he
did not think rhino dangerous but thought lions very dangerous;
but to his mind elephant hunting under the conditions he had
been used to the most dangerous of all. I often said nothing is
so dangerous as a lion and no shot will save you with any cer-
tainty in a lion charge but a ball in the brain, a shot that can only
be fluked, for you have to hit a thing no bigger than a partridge
with your bullet and which you cannot see, and have no time to
draw a bead on, that is coming at you at the pace of the fastest
greyhound. I estimate a lion covers 100 yards in his charge in
3 seconds, perhaps less. Were it possible to draw a bead on his
head between or above the eyes he would have passed under your
bullet’s course before you had pressed the trigger. I also dis-
cussed with him the merits of modern cordite rifles and urged
that powerful penetration and shock with steel enveloped and
nickel-coated bullets gave no advantage over old fashioned big
bores and solid lead bullets either soft lead or hardened lead,
when it is the case that you must smash down with a huge blow
a lion within 30 or 15 yards of you. Hewas inclined to agree with
me but said that for himself he had more confidence in hitting a
lion right when charging with a light rifle that was familiar and
handy than with a heavy .450 cordite and he added these words
“IT can never shoot so well with a heavy rifle.”

I told him I never went near a lion without having in my hand

196 AFRICAN GAME ANIMALS

an antiquated 1o bore (invisible rifling), black powder long bar-
relled hammer gun carrying a heavy charge of black powder and a
heavy solid lead bullet, that no lion had ever got at me with this
in my hand but I did not believe that any man could alone be cer-
tain of stopping a lion with any weapon, that I had on one occa-
sion put a Io bore solid lead bullet in a charging lion’s neck low .
down at g paces and for anything I knew had missed him and a
second similar bullet an inch or two above the first broke his neck.
On the same occasion Mr. Harold Hill at my side hit the lion at
9 paces in the nose (just at the base of nose, with .404 cordite)
and he too might just as well not have fired that shot at all for
anything it did to check the lion. The lion that charged George
Grey had his head up the last 20 yards of his charge; I doubt if
Grey saw anything but mouth, nose, hair, and a tawny mass
catapulted at him.

On the 29th of January 1911 a rendezvous had been fixed
at Lanjoro with Messrs. C. A. and H. D. Hill for 8.30 a.m.
the place being situated between their place Katalembo and
my farm and residence; and handy to the ground we thought of
drawing.

Lions had destroyed many of my ostriches at Kitanga, four
had just been seen on a rocky hill, Kilima Theki, on my ground,
and the Hills had news of 2 lions the day before not far from
Philip Percival’s (Potha farm). Captain Slatter and the Hills
were most anxious to get rid of these brutes and my party were
very keen to assist and have the chance of killing them.

Our party from Kitanga consisted of—

1. Captain Slatter carrying a double .450 cordite rifle, and his
gunbearer. He undertook to stand by Grey at the post he was to
occupy on Theki hill which it was decided to drive first. Captain
Slatter was mounted on a little slow, weak Somali pony.

2. George Grey carrying his .280 Ross, and his gunbearer his
shotgun. He was mounted on a strong, rather slow, but good
pony hired from Newland and Tarlton.

3. Mr. Howard Pease carried a .256 Mannlicher and his gun-
bearer a Holland and Holland .390 (? or .375?) cordite rifle and was
mounted on a very good pony hired from the Boma Trading Co.

THE LION 197
Mr. Clifford Hill undertook to stand by him during the first

drive.

4. Mr. Edward Pease carried a .450 Lancaster cordite rifle
and his gunbearer a double barrelled Purdey .500 express rifle
and I took charge of him during the first drive. He rode a bay
sudan arab.

5. Myself. I carried a .256 Mannlicher and my gunbearer a
double barrelled 10 bore. I rode a good pony purchased at Nai-
robi. The Katalembo party consisted of

6. Mr. Clifford Hill who carried a .404 Jeffrey cordite rifle.

7. Mr. Harold Hill who carried a .450 (?) cordite rifle, and
shotgun with bearer. He took charge of the beaters and the man-
agement of the first drive. They had a mule with them.

On reaching Theki hill Grey and Slatter and gunbearers took
one post: Howard Pease and Clifford Hill, and gunbearers another;
I took my son out onto the open plain so that I might round up
any lion that broke cover and left the hill.

H. Hill drove the hill round and just when the drive was over
a lioness was started in some bush between the rocky sides of the
Kopje and the plain. Grey got a long shot (180 yards) at her
standing and had rather alarmed Slatter by abandoning a safe
post and running down after the lioness into the bush. Grey
missed his shot and for awhile the lioness was lost in the bush
but from the plain below I saw her lobbing along across open
patches in the scrub and then lie down in a small bush surrounded
by long grass. I left my gunbearer with my son Edward Pease
telling my son to keep his eye glued to the bush and mark her if
she went away and if she came to him to leave her alone unless
she attacked. Then I galloped up through the scrub to where I
could see Grey and Slatter standing, and told them we had her
marked down. I intended them to wait there or a little nearer
the bush she was in till the whole party came up but to Slatter’s
and my astonishment in spite of my calling on Grey to wait he
started to run as fast as he could towards the bush I had pointed
out. He was about 30 yards from it when I managed to get up
to him on my pony. I begged him to come back and wait, but
he hurried right up to it, and I, in great fear that we should both
be downed by her, ran up to him hoping she would not charge

198 AFRICAN GAME ANIMALS

before at least a gunbearer or Slatter joined us (they were hurry-
ing after Grey). To my relief my son and party below in the
plain shouted and signalled that she had left the bush and gone
up the rocks. If she had been there it was an even chance that
she would have charged Grey and me and when I saw he had
only his little Ross I was alarmed: but regarding him as an ex-
perienced, confident, and fearless man I felt he would not like
me to find fault, and I only said, ‘you know you ran right to
the very spot where she was and I expected her to charge out.”
He was breathless with running and just smiled.

We decided to leave her alone for another day and proceeded
across to a donga that runs down from Wami (on the Theki side
of Wami) and joins the main dry donga that runs east of Wami
into the Potha River. On reaching this donga (which is the one
where Col. T. Roosevelt killed his first lion) my gunbearer found
the spoor of two lions going down. We drew this donga down,
I myself riding a little wide of it on the Wami side in order to hold
up any lion that broke away on that side. Others were riding
out on the Theki side. After drawing the donga down about a
mile, Clifford Hill walked across to me and said “they are going
to draw that donga on to the junction; we will cut across here
(pointing south) and draw the main donga and cheetah bush
down to them.” ‘Taking some boys we did this, Clifford Hill and
the boys walking in the donga and I riding on the east side to hold
up anything that might break that way: anything that broke the
other side would be in full view of the main party. We drew
the cheetah bush blank and our boys joined the main party at
the junction. I was then about 4% a mile from the main party
on the east side and just about to come down and join them and
get on to lunch at the water at Potha, when Clifford Hill ran up
to me on foot and was joined by two boys and said it was just
about here the 2 lions were seen yesterday. I saw my son’s bay
arab being led and called for him and told Hill to mount the horse
which he did. We rode up the hill and the boy showed us the
little thorn bush with unmistakable evidence of the truth of his
story. Clifford Hill then said ‘“‘we had better go on a bit and
then we can draw the little spruit running down to where they
(1. e. the party) are, it is a likely place for them”’ (the lions). We

THE LION 199

were now about a mile from the party at the donga and above
them on the hill side and getting near the top of the spruit we
were going to draw down, when we both reined up and Hill
looking towards the ridge (towards Bondoni) said ‘‘what are
those?’’ I saw two objects in the distance near the sky line and
said “they look rather like ostriches.” Hill said “are you
sure?” and I got out my glasses: he said “I am sure they are
lions”’ just as I with my glasses came to the same conclusion.
We at once sent the boys off to inform the main party, and off we
started for nearer inspection, discussing where we could turn
them to. We both agreed there was a chance of their stopping
in the cheetah bush if we could get their heads turned that way.
We were galloping fast and the lions were galloping too; but the
rear lion, slower and fatter than the other, was getting behind:
so I suggested not pressing them too much, and Hill also thought
that if we kept well to the left behind them they would be more
likely to stop in the cheetah bush donga. ‘‘Anyway we must
not let them get up Wami,” he said. After going thus about a
mile, and the heavy slow lion who objected to being hurried
losing ground all the time, Hill said ‘“‘look here, I will take the
leading lion, you stick to the big one”’ and off he went to get on
terms with the fast-lion now 14 mile ahead of the other.

My lion went very slowly down to the donga, and the other
lion and Hill were going up Wami side when my lion reached it.
I lost sight of him for a minute here and as I was rather near and
did not like the look of him I pulled up and waited to see him
come out hardly expecting he would go any further. I was a
minute or two watching the spot where he had gone into the
donga and then saw that he had come out further down and was
rapidly catching up to the leading lion. Hill was now 4% a mile
ahead of me, so I just put pace on and caught up with him.
Both the lions were now turning off Wami and my lion had now
passed Hill’s lion and I was thinking of nothing else but where
they would stand and guessed it would be in the little donga
(towards Theki) when Hill called my attention to Grey and How-
ard Pease, who I saw to my surprise galloping hard directly behind
the lions, perhaps 350 yards behind, but gaining on them. Hill
shouted to me “‘They will drive them right up Wami; we must

200 AFRICAN GAME ANIMALS

get ahead; for goodness sake stop them.” We were still galloping,
and till that moment had never thought of anyone else interfering
so that I felt certain that we had both lions collared and safe, and
should hold them till the whole party came up. I could see no
one else in sight besides the two who had appeared so suddenly
upon the scene. I yelled waved and blew my whistle as hard
as I could; but Grey, who must both have seen my frantic efforts
and heard the noise, paid not the slightest attention, and to my
surprise I saw him gaining rapidly on the rear lion, Howard Pease
coming across to me in response to my screams and signals.
Still, I never dreamt that Grey was going literally to try and ride
them down and was more annoyed than anything else that the
lions were being pressed up the hill just as we had begun to turn
them down it. Then Hill called out, ‘Grey is getting too near.”
What happened next all took place in about 4 seconds and as
quick as you can think and see. I put the facts down as they
flashed on me—

Moment 1. Grey galloping hard 100 yards directly at the
tail of the rear lion, the other lion some 60 to 80 yards ahead of
the rear lion.

Moment z. The rear lion whipped round and stopped (as
did the other) fora moment. Grey 80 to 90 yards directly below
him on the hill side.

Moment 3. The lion charged at once.

Moment 4. Grey leapt from his pony in a flash and stood in
shooting attitude all in the same moment, the lion then say
40 yards from Grey. (Hill is positive that Grey dismounted be-
fore the charge began).

Moment 5. Lion at 25 yards. 1st shot from Grey. No effect.

Moment 6. Lion at 5 yards. 2nd shot immediately after Ist.
No effect.

That is what I saw in say 4 seconds of time.

This is what happened on our side—

Hill cried out to me at moment I, some despairing exclama-
tion, such as, ‘‘My God they will have him for a certainty,” and
leaping from his horse said “‘shoot him, shoot him.”

At the moment 1 I was bewildered with horror, and hearing
Hill’s appeal ‘‘to shoot” pulled up dead and had got to the

hd

THE” LION 201

ground; when I saw the lion prepare to charge and charge all at
once. I had just got my rifle to my shoulder and had cried to
Hill “it is too far, it is over 300 yards” when Hill said ‘shoot!
shoot! 200 yards” and he fired. His bullet falling short by about
10 feet, he as a last desperate chance took two hurried snap
shots. Realising the position, conscious that Howard Pease
was either on his feet or pony close by, I leapt to my horse, and
galloped over roughish ground in the hollow dividing me from
Grey with H. Pease, my eyes fixed on the horrible scene in front
of me. The 300 yards seemed like half a mile and the lion lifted
his head when we got within 50 yards and ceased mauling Grey.
At about 20 yards or 25 yards we left our ponies and hardly got
to the ground and ran in when Hill who had not stopped to re-
mount but ran with great speed was there too. The rest of
the story 1s not what I seek to dwell on—Hill gave the lion the
knock out shot with his .404 before it jammed. I fired 3 shots.
The position was rendered worse by the other lion preparing to
- charge and grunting less than 100 yards above.

The questions that need deciding are these:

1. Did Grey hit the lion at all, or once or twice—

To me it looked as if he missed both times but I am sure the
lion had a very bloody mouth when he raised his head when I
galloped up, and I hold the view that Grey missed the first shot
and hit the lion in the mouth the second. I think Grey would
have been dead but for that injury. I understand that Clifford
Hill holds the opinion that Grey missed both shots and that the
lion’s bloody mouth or at least his broken jaw was done by one
of my shots. J am sure this was not done by me or any one but
Grey.

2. Did Grey dismount before the lion charged? or when he had
started. I should have said (but am not clear about it) Grey
only dismounted when the lion’s charge had begun. C. Hill is
positive that Grey dismounted as soon as the rear lion stopped.

3. Could he have got away on his pony when the lion charged?

I say emphatically no he could not even have turned his pony
in the time. I do not know what else he could have done that
would have been better, if he was certain of himself and confident
of his power to stop a lion charge, as he said he was. Otherwise

202 AFRICAN GAME ANIMALS

if he had taken the charge on the pony or got off the pony so that
the pony was between him and the lion, the lion might not have
got at him so quickly and the pony might have got some of it,
and Grey would have had a chance of killing the lion when he and
the pony were brought to earth. Hill considers that if Grey had
turned and galloped off immediately the lions stopped he would
probably have got away.

4. Could we have got across to Grey when first we saw he
was in danger.

Most certainly not. We did all as quick and fast as it was
possible to do a thing from the moment we realized Grey was get-
ting near 100 yards distant.

5. Did the lion charge before Hill’s shot.

I cannot say whether the lion had begun his charge before the
shot or whether the shot started the charge. I can say, both lions
looked like charging, the rear one was going to charge and that
I saw no possible escape for Grey unless he had the fortune to
stop the lion. I never expected to see him stop the lion. C. Hill
is quite clear that he (Hill) fired twice, once when the lion stopped
and prepared to charge and once when he was in full charge.

6. Could any others of the party have come to the rescue or
have stuck to Grey?

No certainly not, even had they known what he was going to
attempt.

Slatter had a little old weak pony that could not gallop, let
alone with Slatter on his back, and he pumped the pony in fol-
lowing and did not even get within sight of the scene.

H. Hill was on foot 2 miles away.

E. Pease was on a mule about 2 miles away.

Towards the clearing up of this incident I can give Grey’s ac-
count in answer to my remarks and questions as well as what he
volunteered. Here are some of his sentences: many of them he
repeated.

1. Remember I say it that nobody is to blame for what has
happened, that it is entirely my own fault.

2. It was very foolish of me.

3. I was not riding directly behind the lions but a Jitile to the
right of them. (H. Pease corroborates this)

THE LION 203

4. I don’t think I was as near as 100 yards when the lions
stopped. I saw both the lions. (C. Hill thought Grey could not
have seen them stop)

5. I did not believe that lions would charge unless shot at.

6. As soon as Hill shot I leapt off my pony.

7. I thought that after galloping the lions they would stop
somewhere and that I should get a quiet shot at them.

8. (To C. Hill going to the station) ‘‘I thought I was a match
for any lion but find that is not so.”

I suppose from the time the charge took place to the moment
when the lion was dead would be about 3 minutes, not more.
We had 300 yards to gallop (Hill ran it almost as fast as we gal-
loped, mounted and dismounted). The lion ceased to maul
and savage Grey when he saw us galloping up, and lifted his
great shaggy head and stared at me with his mouth wide open
and all streaming with blood. At about 20 yards Howard and I
had dismounted and walked in, the lion looking at us, the other
lion crouching grunting and lashing his tail just beyond the one
we had to kill. Hill’s and my first shots were simultaneous, Hill
fired another and I another, and Howard two.

To the first shots the lion responded by savaging Grey and
then he bounced up and came at us and went back and lay down
on Grey again and Hill’s next shot knocked him out and then
Hill’s rifle jammed. I put the muzzle of my rifle to the right
side of the lion’s head and fired and he just flopped it on the
ground. We thought Grey was dead. Hill called to me “‘the
other lion is going to charge my rifle is jammed you have got to kill
him.” I said “‘for God’s sake don’t look at him I cant stop him
with my rifle” (.256)—I told Howard not to look at him. Grey
said to Hill, “you want a rifle, take mine”’: then in a minute or
two the lion moved off slowly.

A couple of days later poor, gallant George Grey died in the
hospital at Nairobi. A gamer and hardier man, and a stauncher
and more loyal friend never trod the African wilderness.

As we have said above we do not entirely agree with
Pease about the rifles proper to use for lions. He explicitly

204 AFRICAN GAME ANIMALS

states that even the ten bore, reinforced by a powerful
four hundred cordite, both hitting a charging lion at short
range in the head, but missing the brain, failed to stop him
for even a moment. Surely this shows that even the most
powerful rifles may do no more than lighter ones, under
the very—and only—circumstances when the big rifles are
supposed to be superior. A Springfield with the pointed
bullet, or a Winchester .405 with a soft-nosed bullet,
could not have done any worse than the two big rifles
in the above instance, and we believe they would have
done better; at any rate, even at close range, they are
almost, if not quite, as good as the very heavy rifles, and at
all ordinary ranges are far superior. Personally we regard
the mere fact of a rifle being a magazine gun as conclusively
establishing its superiority over a double barrel for lion
hunting, if both are good modern weapons. While we were
in East Africa a Mr. Williams, a friend of Mr. McMillan,
was badly bitten by a lion, because with his two barrels he
slightly wounded it at a distance, and was seized before he
could reload, whereas if he had been using a magazine rifle
he would probably have stopped his assailant. The experi-
ences of the two cowboys, Loveless and Means, who with
Buffalo Jones roped the lioness, south of Kijabe, a few
months after we were there, shows that a lion cannot catch
a really good horse, already at full speed, even if it has only
a few yards start. Doubtless a lion can catch an ordinary
African hunting pony, which is standing or has to wheel when
the lion starts, if it is under a hundred yards off. But the
cow-punchers with Jones were riding big American cow
horses of the best type, fast, agile, and thoroughly trained.
The lioness was at bay under a bush, and the punchers kept

THE: LION 205

riding by her at a run, throwing the rope; the bush always
prevented the noose going over the lioness, and she would
charge at full speed (as the photos show); she was only the
length of the rope behind when she started, and gained for
the first fifteen or twenty yards, but never quite succeeded
in catching the horse. Only the best and coolest riders,
on the best horses, could have performed the feat, how-
ever. The lioness finally became cowed, crouched in a
donga, and was roped, thanks to the nerve of Jones, and
the extreme skill of the cowpuncher who did the actual
roping.

Light, agile men who keep some distance from the lion
when he is at bay can leap on their horses and ride off after
shooting if the lion charges. Says Mr. Roosevelt: “ Per-
sonally, I was too old and stiff to try this, and on the occa-
sions when I was riding and dismounted to shoot at the
lion, I left the horse, advanced as close as I thought the lion
would stand without charging, and then trusted to straight
powder, and my repeating rifle. I stopped one lion while
in full charge, another as it began its charge, but while it
was only trotting, and another just as, I believe, it was about
to charge; and I killed a wounded lioness in tallish grass,
which ought to have charged but did not, and at eight
yards merely wounded a lioness, which I ought to have
killed outright, but which galloped away instead of charging,
so that a minute or two later I got.an even closer shot, and
bagged her. Thus I personally met with no adventures
and neither did Kermit, although, when in company with
Leslie Tarlton he one afternoon galloped a party of eleven
lions and killed five, one of them, a lioness, charged with
fierce determination. But since I left Africa Tarlton per-

206 AFRICAN GAME ANIMALS

formed an interesting feat with lions which shows what a
really good hunter can do when luck is not hostile. He
writes me as follows under date of June 2, 1911:

Provided you can decipher this missive, I can entertain
myself by hoping that a letter from a “far countree” breathing
blood and ‘‘deeds of derringdo”’ may take your thoughts away
for a time from the sterner work of life, and recall some pleasant
hours spent together, what time we carried a lioness through the
darkness one night, and fondly scanned the blackness for the
bellicose rhino! Ina recent letter to Kermit I mentioned a rather
successful hunt at Simba, and last Saturday I took a run down
to a point half way between Athi and Kapiti Plains Stations, and
arriving there at 3 P. M. hunted unsuccessfully for lions until dark.
Next morning I was at it early, hoping to return in time for office
Monday, but my hopes were considerably damped by finally see-
ing four lions galloping away some two miles ahead in that open
country about five miles nearer Nairobi than Wami Hill where
you got your first lions. I was on foot, and without much chance
of a stalk in that country. I sat down and watched them until
they stopped under one solitary thorn tree close to the only piece
of bush within sight—about enough to really cover the four if
they wanted to hide. Here they lay down, but after a long and
painful stalk, the nearest I could get was 500 yards before they
saw me and slid into cover—z2 lions, 1 lioness and 1 large cub.
Again I made a very careful stalk on the bush, and finally getting
up to, beat it carefully—to find the birds flown! So I sat down
and had no trouble in picking up my quarry trotting steadily
away in almost perfectly flat country a mile and a half off. So
the game being up apparently, I sat down under my bush and
gazed through glasses at the quarry. Presently they started to
walk, and the youngster lagged behind—sat down—and then
went on. So we all hid under the bushes, and after half a mile
of walking and stopping, the four finally came to a halt about
2 miles away, the youngster lay down, and presently went to
sleep. Whereupon I set a boy on guard and followed suit. It
was about 8 a. M. and a cool cloudy day and I had lots of time

COWBOYS WITH BUFFALO JONES ROPING A LIONESS

__ By courtesy of Everybody's Magazine
Copyright, 1910, by the Buffalo Jones African Company

THE LIONESS CHARGING THE COWBOYS AT FULL SPEED

By courtesy of Everybody's Magazine
Copyright, 1910, by the Buffalo Jones African Company

COWBOYS ROPING A LIONESS

THE LION 207

—so although without any idea of a shot, feeling quite comfort-
able and lazy I thought I would try and experiment. At 10.30
the boy waked me and said they were all lying asleep—and
through my glasses I saw that three out of the four were really
fast asleep—only one—the lioness—lying head on paws looking our
way. Sol told the boy to go to sleep and took up the watch myself.
Presently the lioness got up—stretched—walked 20 yards or so
and again lay down—this time broadside on—and then to my
joy her head gradually rolled over until she too was flat on the
bare plain like a great dog. Now you can imagine that in that
almost flat plain to be able to see the lions properly, when lying
on their sides, two miles, not much in the way of grass obstructed
the view, but a finished piece of cunning on their part eventually
aided me. They had chosen for their resting place the top of a
very slight rise, which commanded a view of the country all
round, rather like this:—

i Siig eee Ses edie a ees SE Ree eee
eae Aa pi OR ASS AO Onsite oe Sates ae hp en ===
x SWALE Ta INCHES LONG. wien Sees ee ae ee ee
eG) wah fh Cone a ali TS EeANoS WSeoey SRE °
& ee oe F ~~ eet te Ae eX Benes ~~
Fad Aor eB ie ie ene et eu anti ae Ses
~~ as : Ln MSS ma

Sy yet Suerte

mae

“ne ~e-:-:

but by going back and making a detour like arrows > A I was
able to take advantage of about half a mile of longer grass, and get
to point (1) before the old lioness got suspicious. Here she got
up suddenly and I dropped flat and lay still for half an hour.
At the end of that time as she was still on the gui vive (I’ve marked
her © to distinguish her from the other sleepy ones) I took an-
other nap, this time for an hour. At the end of that time, she
was fast asleep again and then came the tug of war. Another
100 yards of my long grass (up till then about 2 feet long) and
the ground was devoid of cover except for very scanty grass
well-fed-down and the whole absolutely commanded by the little
rise where the lions lay. By filling my hat with grass, as did my

208 AFRICAN GAME ANIMALS

boy—and crawling on my stomach, I began slowly to draw nearer,
and the flatness of the country helped me in that the nearer I got
the less advantage did the hill give the lions. Occasionally the
lioness would look up but of course, we at once jammed our
noses in the ground and kept stock still, and a friendly kongoni
discovering them asleep distracted her attention and the sun com-
ing out she got sleepy again and finally rolled over once and for
all. Doubtless to the crocodile, the python and the puff adder,
to say nothing of the subordinate government official or the
baksheesh seeking gunbearers, a recumbent position may be both
the natural and comfortable one for locomotion, but I grow old,
rheumaticky and stiff, and what with the blazing sun, swarms of
ticks, thorns, etc., etc., I should have been thankful if early in
the game the lions had cleared out—as it was, begun in a spirit
of pure curiosity, as the distance slowly but surely lessened be-
tween us, I felt as if the whole of my future depended upon besting
them and I think if necessary I would have stayed out all night.
However, the longest lane ends at last and finally I got them so
beat that I rose to my knees within 80 yards of them, and having
thoroughly got my breath—crawling winds me terribly—I care-
fully wiped the dust off my foresight, saw the other rifle-sight was
clean, drew a steady bead on the biggest of the lions—and not
fancying a shot at them lying flat on the ground, I “‘tschissted”’
in what I flattered myself was an exact imitation of the kongoni
which had been puffing at them for so long. I did it very softly—
but those sleeping lions were on their feet and saw me about one
second—just one second—too long for the biggest lion who got
it on the point of the shoulder as he raised himself. I gave him
another which decided him to leave and then swung on to the
other lion, who went head over heels to the shot, but was up and
at me like a streak—whereupon his mate joined in. This was
bad luck as between them they gave me so much trouble that
by the time I had killed them the lioness and cub were out of
range. They were two fine big fellows, but as customary with
Athi lions, almost maneless. I wish Kermit could have been there
—it was the most difficult interesting stalk I’ve ever made. I
took the time and found it was just 3:30 P. M. so I think I earned
my trophies.

THE LION 209

In this letter Tarlton does not mention that the two
wounded lions charged him from different quarters, coming
on with savage fury; and they were stopped because Tarlton,
in addition to having great coolness and nerve, is an ex-
ceptionally good shot with the rifle at dangerous game.

We do not regard marksmanship as the most important
quality in the chase of dangerous game; but it is very impor-
tant, and, of course, no man has a right to follow dangerous
game at all unless he is a good shot, while if he is a really
first-class game shot his task is very much simplified.
Pease, Delamere, Tarlton, are all of them excellent shots.
Stewart Edward White in his trip to East Africa in 1911,
during the course of which he killed a dozen lions with his
little Springfield rifle, was with our old friend and hunting
companion, Cuninghame; and Cuninghame wrote us that
of all the hunters, of every description, whom he had ever
seen in Africa, White was the best shot. Among other noted
lion hunters, Selous, Akeley, Stigand, and Rainey are all good
shots. Of the above, Delamere and Stigand have been
mauled by lions; Akeley was mauled by a leopard and nearly
killed by an elephant. Mrs. Akeley and Mrs. McMillan—
whose husband is among the successful lion hunters—have
both of them killed lions to their own rifles.

Occasionally lions are killed with the rifle, as in the in-
stance above by Tarlton, and in other instances in the
careers of almost all veterans like the men named where the
conditions are such as to imply high prowess on the part of
the hunter and the running of grave danger. Normally,
however, of course, lion hunting with a modern rifle does
not mean danger of the kind incurred by coming to close
quarters. Sucha feat as that of Jones, Means, and Loveless

210 AFRICAN GAME ANIMALS

in roping the lioness implies much greater daring, skill, and
risk than is normally attendant upon shooting a lion. The
same is true of killing a lion with spears, after the fashion
of the Masai and Nandi. In this kind of hunting, however,
much depends on the type of spear. It happens that the
long, very heavy, narrow spearheads of the East African
cattle-owning foot-nomads are much better suited for this
particular sport than the light spears of the equally fearless
and gallant Zulus to the south of them and Somalis to the
north of them. In Somaliland the lions subsist largely, and
often mainly, on the flocks and herds of the Somalis, and
frequently become man-eaters; but the Somalis only attack
them under exceptional circumstances, for their spears,
formidable enough against men, are too light for lions, and
the danger to the lives of the hunters in the contest is very
great. The Zulus also ring and kill lions with their spears,
in the Nandi and Masai fashion, but their spears are, for
this purpose, much inferior, and in the fight men are far more
often killed and mauled than is the case in middle East
Africa. We saw the Nandi spearmen kill a big-maned lion;
he mauled two of their number; but a couple of days pre-
viously they had killed two lions without getting a scratch.
Akeley saw ten lions thus killed and only one man was
hurt. «Mien are rarely killed im these. contests: “Unisys
because the Nandi and Masai spears are so heavy that they
drive right through the lion, and into his life, from any
angle; the first spear driven into the big-maned lion above
mentioned entered at his left shoulder and came out through
his right flank near the hip. In consequence a single spear
will not infrequently kill a lion. For lighter and more agile
foes the spear is too heavy and slow, and for this reason the

THE LION 211

Nandi and Masai find the leopard more formidable than the
lion in this kind of hunting. The Masai or Nandi spear is
made of soft iron and is given a fine saw edge by whetting
ona stone. This is a most effective cutting edge, being so
sharp that it cuts its way through the toughest hide.

Among the horse-owning tribes of northern Africa there
are a few in which it is customary to kill not only the lion
but the elephant, rhinoceros, and buffalo from horseback
with the spear or sword, some using one weapon, some the
other. Of course, the spears and swords used in such hunts
are not dull and blunt like ordinary cavalry sabres kept in
metal scabbards; they have fine, carefully guarded points
and razor edges.

Mr. Heller accompanied Paul Rainey on his African trip,
in which Mr. Rainey killed several score lions with his pack
of American bearhounds. Mr. Heller describes certain in-
cidents of this trip as follows:

“The Loita Plains form a region of grassy veldt inter-
spersed by areas of scattered acacia scrub and dotted by
several high rocky kopjes which dominate the district.
These kopjes being both rocky and scrub-covered are used
as day retreats by the lions which have been attracted to the
district by the immense herds of game which graze on the
plains. The driving of one of these hills gave us the most
exciting lion hunt we experienced during the entire year’s
safari with Paul J. Rainey.

“Six months had been spent in a neighboring district
hunting lions with dogs until the pack were practically
annihilated. After the loss of all the best hounds or trail-
ers this form of sport had to be abandoned and lion driving
with porters was resorted to. Leaving the Amala River

212 AFRICAN GAME ANIMALS

drainage we entered the Loita Plains from the west and
struck across country to a high rocky kopje which domi-
nated the whole western section of the plains. Camp was
pitched on the open plains near a few muddy pools of red
water some three miles from the kopje and preparations
were made for beating the hill the following morning.
“The hill from camp stood out like a great headland, but
at the back it sloped away gradually and connected by a
wide, grassy pass with two smaller hills. The summit was
strewn with huge reddish granite boulders interspersed with —
a few dark-green bushes. The plain at its base was every-
where grass covered and open except on the camp side to-
ward which a scattered growth of thin bushes ran out as a
long spit and connected with the bush-covered area about
the water-pools near camp. The beating was done by a
hundred porters who were sent in a body to the low pass
behind the hill, from which point they began their opera-
tions so as to prevent the lions from breaking back to the
shelter of the two smaller hills. Rainey and the hunters
and photographers stationed themselves west of the hill in a
slight depression of the veldt and awaited results. As we
lay there waiting we saw a lioness appear on one of the huge
boulders at the summit. She stood silhouetted against the
sky scanning the plains for a means of escape. The noise
which the beaters were now making on the lower slopes of
the hill was beginning to cause the lions uneasiness. Soon
other lions appeared on the rocks, sometimes as many as
four appearing together and searching the horizon for the
best means of escape. At intervals two magnificently
maned lions appeared in sight, one with a deep black mane,
his yellow head framed by a wide circle of black; the other

THE LION 213

a yellow maned lion of equal size but less conspicuous. We
could form no idea of the number of lions on the hill, but we
were all wildly excited by the prospect of the numbers which
had been seen. Our position was an exposed one on the
open plain, past which the lions would flee only if driven
by the beaters. At the eleventh hour I was sent over to the
cover of the spit of scattered bushes to turn the game into
the plain if it used this, the only bit of cover available.
My position placed me between the lions and the camp
toward which it was thought they would not attempt to
flee. Several of them came out of their cover on the hilltop
and surveyed me and my gun-bearers as we stood clustered
about the base of a gouty little tree. These great, wily
brutes were displaying a very intelligent interest in our
movements, and I have no doubt but that the leaders of
the troop had decided at all events not to escape by way
of the open route past Rainey and his formidable array
of gun-bearers and photographic assistants. When hard
pressed by the beaters they left the hill in a pack, coming
down the hill in close formation like a herd of sheep. They
came down at a fast trot directly toward me, led by one of
the lionesses. The two maned lions were the rear-guards.
Rainey and his men were in action as soon as the lions had
reached the plain, speeding toward them on their horses at
a terrific pace. The two maned lions upon observing these
hostile movements turned back into the hill and were never
again seen. The rest of the troup numbering approxi-
mately sixteen, kept on their course along the spit of scat-
tered bushes toward my position. They came past me at a
swinging trot in a long line one hundred and fifty yards
away. It was my duty to turn them toward the hunting

214 AFRICAN GAME ANIMALS

party. I took several shots at the leaders but these had
no effect; in fact, they paid not the slightest heed to them
but trotted past in an unbroken, unswerving line. The last
of the line, however, was a lioness which displayed a threat-
ening attitude toward me. When she got within two hun-
dred yards she left the troop and charged me. She came
in a crouching trot, growling and lashing the air with her
tail. I could not understand her hostility, for only the lead-
ers had been fired upon. I clutched the 405 Winchester and
waited for her to come within range of my excited marks-
manship. During this wait the gun-bearers tugged at my
coat, whispering to me to shoot before the dangerous animal
got any closer. At seventy yards I shot and hit her low
in the chest, the shock stopping her for a second. She was
up in an instant, but a second shot in the chest sent her
down again. She came on again at once, growling savagely.
A third shot through her leg stopped her again. Finally
another shot brought her down and she lay with her head
raised, growling defiance. At this juncture one of the gun-
bearers excitedly called my attention to two half-grown lions
which were approaching us from the rear. These were the
offspring of the lioness, which had charged me because my
position had accidentally placed me between her and her
cubs. Rainey’s two stag-hounds had attacked these young-
sters which were several times the weight of the dogs,
and had annoyed them so that they had lost the troop.
They strayed in my direction, the dogs following and snap-
ping at them whenever opportunity offered. After ridding
myself of these two enemies by a couple of shots, I again
turned my attention to the lioness and finally killed her
with one of my last cartridges. Before this occurred, how-

THE: LION 215

ever, she was so broken down by the numerous body shots
that she was only able to shamble along and quite incapa-
ble of doing any damage.

“The troop after passing me was overtaken by Rainey
and his mounted men and six of the lions were shot. After
firing had begun the danger from attack was greatly in-
creased, owing to the number of wounded animals concealed
in the scattered bushy country. The horsemen unwittingly
rode quite close to some of these dangerous beasts, but es-
caped injury by rapid firing or retreat. Three of the fleeing
lions ran through the camp and were fired upon by the
askaris on guard. The morning’s bag netted nine lions,
three of which were large lionesses and the others maneless
lions.

““An attempt was made in the afternoon to beat out the
two large maned lions which had retreated to the hill.
They, however, could not be found, owing without doubt
to their wisdom in fleeing far away and not stopping in the
vicinity of the hill from which they had been driven in the
morning.

“TI was greatly surprised to catch lions in steel traps set
for hyenas baited with carcasses. Four lions were caught
in this manner on the Rainey safari, all within the space
of a few months. This very much astonished me, for I had
used the same methods of trapping and the same traps for
two years in the region without experiencing a single in-
stance of the kind. The traps used are ridiculously small,
the jaws being very little greater in diameter than the paw
of a lion, and the weight only about six pounds. They are
so small that it is only possible to catch a lion by a single
toe or at most by two toes. |

216 AFRICAN GAME ANIMALS

“The first lion caught in one of the hyena-traps showed
the great holding power exerted by a single toe. Four traps
were set about the carcass of a newly shot hartebeest.
During the night the carcass was visited by two male lions,
one of which was caught in one of the traps by the middle
toe of his right hind leg. The jaws of the trap took a firm
hold of the toe behind the great toe pad. To this trap was
attached a drag, consisting of the limb of a thorny acacia
eight feet in length and much branched. The lion, after the
first struggles with the trap, dragged it a few yards to a
termite hill. About this mound he rushed many times, wear-
ing in the earth a path a foot wide. Finding this manceuvre
of no avail in freeing him of his encumbrance, he proceeded
some fifty yards further with the drag and climbed up the
slanting trunk of a small tree which was bent over at a very
oblique angle. At a point five feet above the ground the
tree forked and through these forks he bounded to the
ground. The trap, however, caught in the forks and held
his hind foot. Here he hung, his forefeet and nose touching
the ground, and the hind part of his body suspended from
the tree. His plight was now irretrievable. In the morning
I found him in this position with his companion some three
hundred yards from him, still feeding upon the hartebeest
carcass. The latter bolted upon seeing me and left his
companion to his fate. The trapped lion, upon perceiving .
me, began to growl and thrash about in a last effort to free
himself in the presence of this new danger. I prudently
shot him at a distance of one hundred yards. Upon exam-
ining him I was dumfounded to find that he was caught by
a single toe. He was a full-grown male, but maneless, and
in excellent condition with a heavy layer of fat under his

THE LION 217

skin. Later in the morning, after the return of Rainey with
the hounds, the dogs were put upon the trail of the lion
which had bolted upon our approach. But a single one of
the hounds was able to follow the trail which had now been
subject to three hours of sunlight. Carefully working his
way along this faint scent, the hound took the pack some
two miles away into a bushy ravine. Here they found his
majesty and routed him out. He ran a mile and then sav-
agely took his stand in another dense thicket. Rainey soon
discovered him and waded into the bush to within forty
yards of the bayed brute. As soon as an opportunity offered
of shooting the lion without endangering any of the dogs, he
shot him. ‘This lion was also maneless and in fat condition,
being practically the same age as the trapped specimen.

“To those who have witnessed the savage way in which a
wolf attacks with its teeth and the entire strength of its
body a trap which holds it by a foot, it will appear quite
beyond the powers of belief to give credence to the catch-
ing of adult lions in traps of similar size as here narrated.
The miracle, however, is due to the extreme caution with
which the lion preserves its teeth from breakage. It re-
fuses to risk them on hard substances like iron and will not
even attempt to break heavy bones with them. When
caught by a steel trap its caution is still dominant, and its
rage asserts itself merely in futile tugs at the drag without
any assault upon the trap or attempt at amputation of the
limb by which it is held. The hyena, on the other hand,
though a thoroughly cowardly beast, will attack the trap
savagely, doing great injury to the crowns of its teeth, and
at times as a last resort freeing itself by the amputation of
the limb by which it is held.

218 AFRICAN GAME ANIMALS

“In order to avoid trapping more lions the traps were
baited with small bits of meat which it was thought would
only be attractive to hyenas or jackals. Four of the traps
were set near the skinned carcasses of two small wart-hogs
in a sandy ravine some four hundred yards from camp.
Another trap was set close by within the mouth of a large
burrow into which a few fragments of meat had been cast.
During the night a party of three or four lions visited the
baits and sprung the traps at the carcasses. Their tracks
were perfectly preserved in the dry sand of the ravine.
Finally one of the lions in investigating the burrow which
had been baited was caught by one of the toes of a forepaw.
After thrashing about in the sand, he made off through the
dense bush growing on the sides of the ravine. In the morn-
ing on my visit to the place I found him by trailing on the
hillside some two hundred yards from where the traps
were set. He threatened me with deep growls and thrashed
about in an effort to escape. The sun was still very low
and shone directly in my eyes when I was in a position free
enough from bushes to make a shot possible. I shot at him
in this unfavorable light twice without much apparent
effect. He then reared up on his hind legs and drew the
trap up near his side. I fired again when he was in this
position and then he disappeared in the bushes. He could
not be seen lying dead or wounded, and we could not with-
out foolhardy risk venture into the thicket for fear he was
still there, and very angry. Luckily Rainey was near by,
returning from a morning’s hunt with the hounds. With
the pack we entered the bush, and they at once picked
up the trail. When we reached the spot where the lion had
taken his stand, we found the trap broken apart, one of the

THE LION 219

jaws having been severed by the last bullet I had fired.
Within half a mile the dogs bayed him in dense brush. In
order to shoot the lion, Rainey had to approach within fif-
teen yards. He was alsoa fully grown male but maneless.
Upon examining his forepaws the marks of the jaws of the
trap were distinctly visible on the inner toe, the congested
condition of which upon skinning gave further proof of his
having been caught by that member. No growling was
indulged in by this lion during the night, absolute silence
being maintained. The failure of this lion to charge when
freed from the trap by my last shot is quite unaccountable,
as he had been put to several hours of irritation.

“The next lion captured by the hyena traps was taken in
a very different region. While we were camped on the
lower Guaso Nyiro at the point where the Marsabit road
crosses the stream, one of the mules died of tsetse infection.
The country about this camp was a low, dry desert, intensely
hot, having an altitude of only two thousand feet. The
district was rather uniformly covered by a scattered growth
of acacias and wait-a-bit thorn-bushes. The body of the
mule lay half a mile away on the brush-covered plain. Near
it I placed some of the steel traps for the purpose of catch-
ing striped hyenas which were abundant in the vicinity.
Several days previously the mule had been guarded from the
vultures and spared for lion bait, but no lions ventured near.
On the night the traps were set two lions visited it, one of
which was captured. This lion gave us a long trail to fol-
low which wound about through the brush for a mile and a
half, and finally into a dense reed bed of tall bulrushes much
higher than our heads. My gun-bearers finally sighted the
tip end of the acacia-bush drag sticking out of the reed
patch. The lion had only entered the edge of the reeds, but

220 AFRICAN GAME ANIMALS

they were so dense that he was quite invisible to us. We
stood within thirty yards making some disturbance, but no
noise or sign of life came from the lion. In order to get a
view into the swamp I climbed a thorn-tree growing near- -
by, but without the desired result. From this elevated
perch I began to shoot into the reeds at about the spot I
thought the lion lay, using for the purpose a small-calibre
rifle. Not until the sixth shot did the lion come out, and
then only because he had been hit. A more persistent case
of attempt at ambush I have never witnessed. He charged
out desperately in extreme ferocity, leaving us in no doubt
as to what our fate would have been had we entered the
reed bed. He was a terrifying sight as he stood there in his
desperate rage roaring and straining to reach the fleeing
gun-bearers. I placed a 405 bullet in his shoulders which
sent him back into the reed bed again. In a few seconds
he was out again to receive another bullet which quite
took the fight out of him. He staggered back into the
reed bed and died. After he had quieted down the gun-
bearers cautiously pulled the drag into the bare veldt, where
we could examine our late foe at our leisure. This lion was
a large-bodied male, considerably older and bigger than any
others we had trapped. What astonished us most was to
find two traps attached to him. His forepaw was held in
one trap by two toes, and his hind foot by one toe in another
trap. With these encumbrances he had gone for more than a
mile over rather rough veldt. Over this course he had been
followed by a lioness, doubtless his mate. Her footprints
were boldly marked in the smooth trail left by the trap
drags. No evidence of her was found, however, in the reed
beds, although the negroes asserted ‘that she would surely
remain with her mate and fight.

THE: LION 221

*““An instance of the caution and intelligence of lions
was observed in the circumstances surrounding the trap-
ping of the fourth and last lion secured in such an un-
sportsmanlike manner. The battle-ground of this last
episode was on the broad summit of Mount Lololokui.
The mountain is a great table-topped mass standing isolated
out in the Marsabit desert, above which it looms with quite
perpendicular slopes to a height of six thousand feet. The
summit is several square miles in extent and as level as the
floor of the plain at its base. Two springs occur on the
summit, which are the only waters known for a radius of
some fifteen miles. While camped at the larger of the
springs my guide told me of the existence of the other which
he offered to show me. I readily accepted, and next morn-
ing we set out with the guide for the spring. It was situ-
ated in a bushy ravine, along the floor of which we ap-
proached the spot. When within a few yards of the spring
we were startled by several short, growling ‘whuffs.’ Peer-
ing down the few open avenues in the bushes we beheld
three lions scattered out in a semicircle some twenty yards
away. We had awakened them from their noonday nap
and they were greatly startled at our presence. —The moun-
tain being without a single human inhabitant, it was the
custom of these lions to come to the spring at dawn for
water and lie there most of the day and then go down to
the plain below at night for their prey. After glaring at
us for a few seconds, the lions retreated through the brush
and were not again seen that day. Great confusion reigned
among my followers; the guide nimbly mounted to the top
of a large rock, my gun-bearers peered about under the
bushes to locate the lions, and I brandished a shotgun. At
the spring we found the footprints of the lions and also

222 AFRICAN GAME ANIMALS

others of bush-pigs. Specimens of the latter animals I
greatly desired to add to the collection of mammals I was
making, and steel traps were accordingly set that night at
the spring in the hopes of catching some of the pigs. Upon
approaching the spot in the morning, we heard the growls
of a lion and finally made him out in the brush. He was
caught by a toe of his forepaw in one of the traps, but all
we could see of him was his head through an opening in the
bushes. I shot him at twenty yards. He proved to be full
grown, but not yet fully mature. The tracks about the
spring showed that his two companions had been to the
water with him. They, however, never again returned dur-
ing the fortnight we remained there. A single instance of
capture by trapping was sufficient to cause these cautious
animals to avoid the locality and seek other water several
miles away.”

Key To THE Races OF leo

Body color buffy or olive-buff; mane long-haired, yellow, black usually
only at tips on shoulders or along midline of nape; teeth
larger; female tawny, sides of belly and inside of legs
marked by pale, indefinite darker spots massaica

Body color of male tawny or ochraceous; mane short-haired, tawny, black
or dark only on shoulders and nape; teeth smaller
NYANZE

East Arrican Lion
Felis leo massaica
Native Names: Swahili, simba; Kikamba, mwenyambo; Kikuyu, ngatia;

Masai, ol ngatuny.
Felis leo massaicus Neumann, 1900, Zool. Jahrb., XIII, p. 550.

Rance.—The whole of British East Africa northward
through the Uganda highlands east of the Nile and the

THE EION 223

Rift Valley of Abyssinia, south through German East
Africa, and west as far as the Tanganyika drainage.

The lion is universally distributed throughout eastern
Africa from the coast inland to the high plateaux, and is
wanting only in the dense forests and absolutely waterless
desert tracts. Almost a century ago an Austrian natural-
ist, von Meyer, published in Vienna descriptions of the
Barbary, Senegal, and Persian races of the lion which were
in vogue for a time. These names have remained long in
obscurity, however, and have only recently been brought to
light by Hollister.* Naturalists are now convinced that the
lion shows definite geographical variation which is con-
stant enough to warrant the recognition of many of the de-
scribed races. In 1900 Oscar Neumann named the East
African race, massaica, basing his description upon the skin
of an adult male which he had obtained in Kibaya, south of
Kilimanjaro, in German East Africa. Some years later
Lonnberg renamed the same race sabakiensis, from speci-
mens collected on the lower slopes of Kilimanjaro.

The East African race may be known by its light-buffy
color, yellow mane, large body size, narrow skull, and the
small size of the cheek teeth. ‘The body color ranges from
an ochraceous-buff to an olive-buff, often showing a dis-
tinct gray cast. The mane usually shows black hair along
the nape and throughout its posterior border on the shoul-
ders. Occasional specimens are found in which the whole
mane is black with the exception of the fore part on the
throat and sides of the head. Of the large series of speci-
mens from British East Africa in the National Museum
there is not one which can be classed as black-maned. A
specimen killed by Carl E. Akeley, on the Mau summit,
at 9,000 feet, near the station of Molo, has the greatest
amount of black in the mane of any specimen examined.
The dark extremes are seldom met, however, the usual
coloration being a light buff, lined dorsally by black-tipped
hairs, with a mane showing black-tipped hair only on nape
and shoulders. One of the color characters often assigned
to this race is the spotted sides in the adult. An examina-
tion of a large series of old or fully adult male skins, how-

* Proc. Biol. Soc., Wash., 1910, p. 123.

224 AFRICAN GAME ANIMALS

ever, showed the sides unspotted, but the adult females
and the young show darker spots on the sides of the belly
and inside of legs of an indefinite sort which are only evi-
dent on close inspection. The spots are large blotches of
ochraceous color and show very little contrast to the buffy
or whitish ground-color. The lioness is decidedly darker
than the lion, the body color being tawny and only ex- -
ceptionally as light as the buff or olive-buff of the lion.
The lioness of the South African and Abyssinian races is
quite identical in tint of color to the lion. The ‘former,
however, is not spotted on the sides of the belly. The lioness
of the Abyssinian race shows spots on the sides of the
belly and inside of the legs similar to those of massaica,
with which they also agree closely in the tawny body color-
ation. The nearest geographical ally of the East African
is the desert race from Somaliland, from which it is doubt-
fully distinct. The coloration is quite identical in the two
forms, but the mane is apparently shorter, the body size
less, and the cheek teeth smaller in the Somali race. The
characters assigned by Noack in the original description of
Felis leo somaliensis, of larger ears and longer tail, are not
applicable to the race, these parts having the same pro-
portionate size as in other members of the group. Noack’s
description was based on a pair living at the Berlin Zo-
ological Gardens, and the characters he assigned to the race
are merely such as appeared upon casual observation and
are not founded upon actual measurements of specimens.
Doctor Matschie has informed us that the types have been
exchanged by the Berlin Zoological Gardens’ to animal
traders, and their present abode is unknown. The unfor-
tunate condition of these types is a good illustration of the
loss and confusion to systematic work so often attendant
upon the pernicious custom of naming species from living
specimens. In the present case we have no exact char-
acters and no knowledge of the skull structure of the race
described as somaliensis, merely a few casual observations
to which are attached a general locality of doubtful value.
Several of the types of African big-game mammals are to-
day living in various zoological gardens. Special efforts
should be made by such institutions to keep trace of these
types, and upon their death deposit the specimens in the

THE: LION 225

- largest available public museum where they can be preserved
and accessible to zoologists for comparison.

Adult lions vary surprisingly little in size, as evidenced
by the slight variation in skull dimensions. The extreme
skull lengths show a difference of only one inch in a series
of fifteen old adult males in the National Museum from
British East Africa. The size variation in the lioness is
greater, the extremes being 17% inches apart in a series of
twenty old skulls from the same general locality. The
largest male skulls in the National Museum are quite iden-
tical in ‘greatest length. The very largest is one killed by
Paul J. Rainey, which has a length of 1414 inches, the next
in size being one shot by Colonel Roosevelt, measuring only
zs inch less in length. An average skull length for an
adult male is 14% inches. The smallest old male has a
skull length of 1334 inches. The skulls of adult lionesses
range in greatest length from 123 inches to 11 inches. It
is doubtful if skulls from East Africa ever equal 15 inches
in length, notwithstanding Ward records several exceeding
this dimension. ‘The largest known skull in length is one
now in the Berlin Museum, which represents a South African
specimen which lived formerly in the Transvaal Zoological
Gardens near Delagoa Bay. The length of this skull, as
measured by Heller, is 1675 inches. Ward records this
skull also as the longest, but overstates its length, giving
it as 17 inches. The greatest or zygomatic width of this
skull is 1o}% inches. A skull in the National Museum,
known as the Richardson lion, formerly living in the Cen-
tral Park Zoo of New York City, exceeds that width by 3
of an inch and is the record specimen in this dimension.
Much stress has been laid upon the skull dimensions owing
to their importance in determining the actual relative size
of specimens. Sportsmen often attach great weight to flesh
measurements, consisting usually of the total length from
nose to tip of tail and the height from the pad of the fore-
foot to the withers. Such measurements, however, are sub-
ject to much personal variation, no two sportsmen taking the
measurements quite alike. Measurements taken of bone
lengths, such as the skull, show the relative size of speci-
mens much more accurately. The largest adult male lion
shot by Colonel Roosevelt was a specimen from the Loita

226 AFRICAN GAME ANIMALS

Plains and had the following flesh measurements: head
and body, 77 inches; tail, 40 inches; hind foot, 15 inches;
ear, 61% inches; weight, 412 pounds. The average speci- —
men is somewhat less than these dimensions. An average
female specimen measured in the flesh: head and body, 62
inches; tail, 38 inches; hind foot, 1234 inches; ear, 434
inches.

Ucanpa Lion
Felis leo nyanze

Native Names: Uganda, empologoma; Bari, kamiru; Unyoro, linant.
Felis leo nyanzeé Heller, 1913, Smith. Misc. Coll., vol. 61, No. 19, p. 4.

Rance.—Uganda and the Nile Valley from the German
border northward to the Sobat and the Bahr-el-Ghazal
drainages, east as far as Mount Elgon and the western
shore of the Victoria Nyanza, and west to the Congo-Nile
watershed; limits of range unknown.

The Uganda race of the lion has been recently described
by Heller from a flat skin from Kampala presented to
Colonel Roosevelt by the European residents. From the
East African lion it may be distinguished by its darker
coloration and shorter mane. The dorsal color is tawny
and quite identical in tone to that of the Abyssinian, but
the mane is light-colored with little or no blackish cast and
short-haired. It differs further from the Abyssinian race
by its narrow skull, relatively shorter cheek teeth, and
narrow mesopterygoid fossa. The Uganda race does not
apparently share the close approximation to its West African
representative that is exhibited by much of the mammal
fauna of the region. The broad-skulled, large-toothed race
senegalensis of West Africa is strikingly different from the
narrow-skulled, small-toothed nyanz@. The coloration of
the type skin is ochraceous-tawny, vermiculated very slightly
with black over the median line of the back. The sides of
the body are ochraceous without darker spots and the belly
is somewhat lighter, being buffy-ochraceous. The mane is
short and does not extend over the shoulders except on the
median line of the withers, where it forms a narrow ridge
of hair 2 inches wide by to inches long. In color the
mane is tawny with a seal-brownish patch on the nape and

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MAP 8—DISTRIBUTION OF THE RACES OF THE LION

1 Felis leo massaica 2 Felis leo nyanze 3 Felis leo rooseveltt

The numerals 1, 2, 3, etc., on the map mark the exact spots where specimens have been collected or observed by reputable
sportsmen, and the lines limiting the distribution are drawn around these numerals so as to map the approximate area
occupied by the race to which the numeral refers.

Represents type locality or exact spot from which the type or original specimen came.

The heavy black line on this and the following maps indicates the route of Col. Roosevelt's African expedition of 1909-10.
227

228 AFRICAN GAME ANIMALS

another at the terminal tip of the narrow stripe on the
withers. The length of the hair on the nape is only 3
inches. The legs are tawny like the back in color on the
outside, buffy on the inside, with the hair on the under side
of the toes seal-brown. ‘The head is like the back in color,
with somewhat stronger black vermiculation, and the ears
have the black spot on the back narrow.

Four complete specimens of this race from Uganda have
been examined at the British Museum, consisting of a lion
and two lionesses from Mulema and one lioness from An-
kole. The skin of the lion has a small tawny mane quite
similar to the flat skin from Kampala. The lionesses are
tawny in color, similar to the lion. The skull of the lion
has a length of 14,5 inches and a breadth of 9;'s inches.
The three lionesses range in skull length from 11356 inches
to 121% inches and in breadth from 7% inches to 8 in-
ches. In the absence of flesh measurements of specimens
the dimensions of the flat skin from Kampala may be taken
as showing approximately the size of the race, the meas-
urements of which are: head and body, 75 inches; tail, 35
inches; ear, 4 inches.

CHAPTER VII
THE LEOPARD AND THE CHEETAH

THe LEOPARD
Felis pardus

THE leopard is widely distributed from China, India,
and the islands of Japan and Java westward through
Persia to Asia Minor, Arabia, and the whole of Africa. It
is over this territory a universally distributed animal,
being found in all the forest and bush country as well as
in sheltered nooks on the plains. The leopard still occurs
over practically all its original range, being able to hold
its Own against man owing to its stealthy habits. Ex-
ternally the leopard may be distinguished from the other
species of large cats by the small size and rosette-like char-
acter of its dark spots. Several well-marked races of
leopards are recognizable, but the Indian races as a whole
are not distinguishable from the African, there being ap-
parently no characters of continental scope in the group.
Skins from Java, for instance, are not distinguishable in
coloration from the Soudan leopard, or typical pardus.
Chinese specimens may, as a rule, be recognized by the
distinctness of the black median stripe formed by a series
of closely linked, elongate, solid black spots, while others
from Central Asia exhibit large rosettes or ocellations

widely separated by the ground-color, somewhat after the
229

230 AFRICAN GAME ANIMALS

style of the snow-leopard. The African races may all be
considered small-spotted, although they show racial differ-
ences in the degree of smallness of the spots. Indian leop-
ards are comparatively uniform in size compared with the
African, the largest races of which exceed any Indian, while
the diminutive nanopardus of Somaliland represents the
smallest of all known races. The African races number
some eight or ten forms, which are distinguishable by differ-
ences in body size, skull shape, arrangement of spots, and
general coloration. In Africa the leopard occurs every-
where, except in waterless desert tracts, from the seacoast
to the limits of vegetation on the highest mountains. It
is rare, however, in the depth of the forest and in the alpine
zone of the high mountains. About the forest edges and
in bush country it is most abundant. The sexes differ
much in size, the males usually showing an excess of forty
pounds over females. There are, however, no differences
in color, but female skulls may be recognized, aside from
the difference in size, by the relatively much smaller size
of the canine and carnassial teeth. The young are much
more finely spotted than the adults, owing to the spots
which later coalesce and form the rosettes being widely
separated and appearing as distinct solid spots, no rosette
formation being evident. This broken condition of the
ocellations, or rosettes, remains until the animal is of large
size, and has given rise among some hunters or sportsmen
to the belief in the existence of two species inhabiting the
same locality, a fine-spotted and a large-spotted form.
Save where it has been killed out, the leopard is found all
over Africa, as well asin southern Asia. From the standpoint
of animal coloration it is interesting to compare the distri-

FOREST LEOPARD, MALE

Man-eater trapped by Edmund Heller on body of an elephant at Meru, Mt. Kenia, B. E. A.
United States National Museum

EAST AFRICAN LEOPARD, ADULT MALE EAST AFRICAN LEOPARD, ADULT FEMALE
Lake Naivasha Shot by Kermit Roosevelt, Nzoia River

FOREST LEOPARD, ADULT MALE NILE LEOPARD, ADULT MALE
Meru, Mt. Kenia Gondokoro, Uganda

RACES OF THE LEOPARD

THE SLEOPARD AND THE CHEETAH: “231

bution of the mono-colored lion and spotted leopard of the
Old World with that of the mono-colored puma and spotted
jaguar in the New World. The facts in the two cases are
mutually contradictory if the effort is made to build the-
ories of concealing coloration on them. In America the big
spotted cat has a range restricted to the dense tropical
and subtropical forest regions; while the mono-colored cat
has a far more extensive range, being found in the north and
south temperate zones, and in the high mountains and on
the open plains, as well as among the forests in which the
jaguar dwells. In Africa the spotted cat is found almost
everywhere that the lion goes, while the latter never ven-
tures into the dense forests which are favorite haunts of the
former. The lion is not only absent from the vast equa-
torial forests of West Africa in which the leopard dwells, but
also from the isolated forest tracts of East Africa, which are
also haunted by the leopard. On the other hand, the leop-
ard is found in most of the country wherein the lion lives.
Along the Guaso Nyiro, in the Uasin Gishu, and on the
Athi Plains, for instance, and in the Sotik and the Lado, we
found the lion and leopard in precisely the same country.
In this open country of grass and thorn-scrub each was
extraordinarily difficult to make out when it was hiding or
even when merely lying motionless; and in thick cover they
were also extremely and equally difficult to see. We could
not find out that the color of the coat had any effect in
making one more conspicuous than the other. This was
true when they were lying, as they ordinarily were, in ex-
actly the same cover.

The wide distribution of the leopard in altitude and
latitude, its ability to lie concealed alike in the thorn-scrub

232 AFRICAN GAME ANIMALS

and in heavy timber, its lithe activity, stealth, and cunning,
its ferocious daring and extreme wariness and caution, make
it as universally known as it is rarely seen. It preys on the
smaller antelopes and the young of the larger antelopes and
zebras; also on birds and small mammals. It is fond of the
wild swine, but avoids the old boars. It is the especial
enemy of the baboons, which are always on the watch for it.
A female or young baboon is helpless against its assault, but —
an old male, unless seized in the first rush in such fashion
that it cannot retaliate, will put up a good fight, for its
teeth are as formidable as the leopard’s. The weight and
claws of the leopard, however, give it the advantage over
even a male baboon of the ordinary kinds (the hideous man-
drill of the western forests may stand in another category).
But if the horde is large and contains a number of full-
grown males the latter may combine for the rescue, and
in their turn assail the assailant. Baboons are fearsome
beasts under such conditions; the father of one of the
*Ndorobo who was with us when after bongo had been killed
by baboons which rallied to the aid of one of their number
that the man was trying to kill with his knobkerry. They
frequently drive off leopards and have been known to kill
them. Lord Delamere came across one such instance. A
leopard attacked some baboons among the rocks, and a tre-
mendous row ensued as the old dog baboons drove the
leopard from the animal that had been seized. ‘That eve-
ning the same leopard, very thin and hungry, came into
camp and was shot; it was already fatally wounded, the
bites it had received being such as only baboons inflict.
There have been authentic cases of leopards being killed by
old boars and by old male bushbuck. Save in exceptional

THE LEOPARD AND THE CHEETAH 2338

instances a leopard does not attack animals bigger than a
half-grown waterbuck, hartebeest or zebra. Unlike wolves
and dogs, which usually snap at the flanks and hams of
their prey, the leopard, like the lion, tiger, jaguar, and
puma, ordinarily kills by bites in the head, neck, or throat.
It will kill a serval or jackal, and is itself, although very
rarely, killed by the lion and crocodile; and it has been
known to go up a tree to avoid wild dogs. With the hyenas
it lives in a state of partial feud. Neither preys on the
other, unless that other is sick or crippled; and a single
hyena dares not meddle with a leopard. But two or three
will attack a leopard and drive it from its just killed prey.
Sir Alfred Pease once saw a leopard leap over the thorn
fence surrounding a Somali encampment, seize a sheep, and
leap back with it over the fence, only to be at once robbed
of its booty by the hyenas outside, which cackled, yelled,
and laughed like so many ghouls as they devoured the spoil
they had stolen from the stealer. It is undoubtedly because
of the hyenas that the leopard is so apt to hang its prey in
the branches of a tree. The strength shown in carrying the
prey up the tree is astonishing. Once on the Guaso Nyiro
we found half the carcass of a big Grévy zebra colt which
had thus been dragged into a thorn-tree. The attack is
always made by a sudden rush from ambush on the sur-
prised quarry; occasionally the leopard lies in wait, but more
often it prowls about and stalks its quarry when seen or
scented, the approach being a marvel of slinking and noise-
less stealth. 7

The leopard is a bold marauder against man, killing the
calves, goats, sheep, pigs, dogs, and chickens of any village
which it visits, and sometimes breaking into the building

234 AFRICAN GAME ANIMALS

in which they are secured. It will also occasionally take to
man eating. Twice we came across instances of this in
East Africa. In each case the offender was a big male
leopard, so old that its teeth were worn down; evidently it
had begun to find it difficult to kill game, and so took for
its quarry the females and young of the most helpless of
animals, man. Both of them made their assaults by day.
One took to killing the small boys that minded the herds of
goats; sometimes he took a boy, sometimes a goat. He
was killed by two old men with spears, on the last occa-
sion when he thus sought to carry off a boy. The other,
shortly before our arrival at Meru, had begun a career of
woman killing. He strangled and ate one woman, and seized
and mauled another, but was driven off. We caught him
in a trap. Both of these leopards killed to eat. But on
another occasion a leopard attacked a man ina fit of
freakish ferocity. A Masai chief, with some followers, was
sitting under a tree at noon, when a leopard, without warn-
ing, sprang on him, clawed him severely, but did not bite
him, and then escaped.

Like other cats, the leopard is sensitive to pain, and is
easily held by even a small trap. Nor does it, like a wolf or
hyena, tend to splinter its teeth by biting the iron. On
several occasions we caught leopards in steel traps such
as those used in America for coons and foxes. In two or
three cases the leopard was caught by only one toe, and
went off with the trap and drag—the drag being a fairly
large branch. As soon as it knew it was followed it charged
ferociously, although much hampered by the trap and drag,
and was shot while charging. It was in each case firmly
held by the single toe.

THE LEOPARD AND THE CHEETAH 235

On the whole, the leopard is the fiercest and pluckiest of
African game. It charges as freely as the lion, and is so
quick and agile, and offers so small a mark, that it is harder
to hit. A bullet stops it more easily, however, and in
strength and size it is so inferior to a lion that its assault is
much less likely to be fatal. Kermit was fiercely charged
by an unwounded leopard, which, after he had crippled and
mortally wounded it, caught and mauled a native. Yet it
weighed less than seventy pounds. The biggest males we
got weighed about one hundred and thirty pounds. Carl
Akeley was bitten and clawed by a leopard he had wounded;
he killed it with his naked hands, partly by choking, partly
by crushing its breast with his knees—an extraordinary
feat. Many hunters have been wounded, and a few have
been killed, by leopards. It is incomparably fiercer, pluck-
ier, and more dangerous to man than the cougar, although
the latter in some of its forms is larger and has longer teeth
and claws. Yet the leopard does not attack and master more
formidable game than the cougar. A good pack of hounds
will kill it as readily as they will kill a cougar. Paul Rainey,
in the course of his lion hunting, killed several leopards.
Four of these, all full grown, the pack finished without as-
sistance. They could always kill a leopard if they got it
in open ground; but among dense bushes or rocks it was
generally able to stand them off, as they could only get
at it in front. Two or three big fighting dogs will kill a
leopard without especial difficulty.

236 AFRICAN GAME ANIMALS

Key To THE RACES OF pardus

Body size smaller, skull length in males less than 9% inches; ground-
color on back lighter than cinnamon-brown; rosettes
often forming ocellated or completely ringed spots

Spots numerous and crowded, the interspaces of light ground-
color narrow; ground-color of hind feet ochraceous,
like the flanks; ventral surface of tail tip with a
narrow whitish stripe
Spots smaller on back and more widely separated
suahelica

Spots larger on back, the interspaces forming narrow reticu-
lations, the predominating color being the black of
the rosettes, or ocellations TUWENZOTIL

Spots fewer, large and widely separated by the light ground-
color which is the predominating color; ground-color
of hind feet white; ventral surface of tail with a
broad white stripe chut

Body size larger, skull length exceeding 9% inches in males; color-
ation dark, ground-color cinnamon-brown, spots on
back small and broken, not distinctly ocellated, and
with central portion not darker than ground-color

fortis

East AFRICAN LEOPARD
Felis pardus suahelica

Native Names: Swahili, chui; Kikamba, ngo; Kikuyu, ngart; Masai,
ol owarukert.

Felis leopardus suahelicus Neumann, 1900, Zool. Jahrb., XIII, p. 551.

Rance.—British and German East Africa, from the
coast districts westward through Uganda to the Edward
Nyanza and Lake Kivu, north to Abyssinia and western
Somaliland.

The East African race of the leopard was described in
1900 by Herr Oscar Neumann from some flat skins which
he obtained in German East Africa during his journey in
1893. Unfortunately, however, his description is very
meagre, the name referring merely to what he called the
large-spotted leopard without being based on any par-

PAE LEOPARD AND) THE CHEETAH 237

ticular specimen or locality. Reference to his specimens |
now in the Berlin Museum shows none marked as the type,
so that an exact idea of what he had in mind cannot now be
ascertained. He mentions obtaining specimens at Tanga,
Lake Manyara, Nai, and Usandawe, German East Africa,
and also Uganda. Neumann labored under the supposition
that two species occurred throughout East Africa, a large-
spotted and a small-spotted form, the former of which he
attempted to describe as swahelica. No substantial differ-
ence in the size of the spots in adults from the region can be
detected. ‘There is, however, a marked difference in spot-
ting in the immature and the adults, the former being fine-
spotted, owing to the rosettes being broken up into several
solid spots which later coalesce to form the rosettes of the
adult. Neumann’s skins were chiefly flat specimens ob-
tained from natives and were without skulls, so that their
relative ages were unknown, and the error of associating the
difference in size of spots with racial rather than age char-
acters was doubtless made. Old males often show the
larger and more completely ocellated spots.

The characters of the Hast African race are the small
size and great number of the black spots, which are rosette-
like in formation, being made up of three or four coalesced
spots into a circular form. ‘The ring, however, is seldom
complete or ocellated in character, but the light central
portion or eye is usually distinctly darker in shade than
the general ground-color. ‘The general dorsal ground-color
is usually light, ochraceous, or olive-buff in shade, paling
below to pure white on under-parts and the inside of the
legs. The body is of medium size, with a relatively broad
head or skull, and small cheek teeth.

The East African leopard occurs almost everywhere
throughout its range except in the low, waterless desert
tracts between the Sabaki and Tana Rivers and in the Galla
or Abyssinian desert, north of the latter river. It occurs
everywhere in the desert in the close proximity of springs,
being absent only where water is absolutely wanting.
Specimens have been examined in the National Museum
from the Kapiti and Athi Plains, Naivasha, Loita Plains,
Kisi Province, Uasin Gishu Plateau, Nakuru, and Voi,
British East Africa. The leopard ranges to a high alti-

238 AFRICAN GAME ANIMALS

tude on Kenia and the Aberdare Range, ascending to the
alpine meadows which lie at 10,000 to 13,000 feet. The
tracks were not infrequently seen at such altitudes, which
would indicate the residence of the animal and not merely
its occasional straying into the alpine region. Such resident
mountain-dwelling individuals had best perhaps be referred
to the next race rather than to this plains-and-bush animal.

The flesh measurements of the largest male specimen,
one obtained at Lake Naivasha by Heller, were: head and
body, 50 inches; tail, 35 inches; hind foot, 1034 inches;
ear, 338 inches. The skull of this specimen measured 9%
inches in greatest length by 575 inches in greatest, or zygo-
matic, breadth. A man-eating male leopard caught at Meru
Station on the northern slopes of Mount Kenia agreed very
closely in size with this specimen, being but little less in
dimensions. ‘he large carnassial cheek teeth, which per-
form most of the work of cutting up the food before swal-
lowing, were decayed and their crowns broken down to the
level of the gums in this specimen, which may have had
some influence on his abnormal habits. Adult males, as a
rule, vary little in size, but the females show much greater
size variation. The female specimen shot by Kermit
Roosevelt at Juja Farm, which showed such ferocity and
attacked him without hesitation when wounded, has the
smallest skull of any specimen and was really diminutive
in body size, the skull being 11% inches less in length than
the largest female specimen. The flesh measurements of
this female were: head and body, 39! inches; tail, 2834
inches; hind foot, 834 inches; ear, 258 inches. A large
female from the Uasin Gishu Plateau is much larger, and
measured in the flesh: head and body, 48 inches; tail, 33
inches; hind foot, 9 inches; and ear, 3 inches.

Forest LEOPARD
Felis pardus ruwenzorit

Native Name: Luganda, engo.
Felis pardus ruwenzorit Camerano, 1906, Boll. Mus. Zool., Torino, XVI,
INO5-34350p:

Rance.—Ruwenzori and Mount Kenia forest region,
ascending to high altitudes; limits of range not known.

THE “LEOPARD AND THE CHEETAH 239

The Ruwenzori leopard is based on a specimen obtained
by the expedition to Ruwenzori, led by the Duke of the
Abruzzi, in 1906. The specimen was secured at a high
altitude, eleven thousand five hundred feet, and doubtless
represents a mountain race confined to the higher parts of
Kenia and the Ruwenzori Range. The photograph of the
mounted specimen published in the scientific results of the
Abruzzi expedition shows a large spotted type of coloration,
the dorsal rosettes being separated by narrow reticulations of
light color and having the central or eye portion of the spot
small, the black ring being much wider than in the East
African race, causing the black color to predominate. The
tail is chiefly black with a few light reticulations on the
basal part. The man-eating specimen already referred to
from the Kenia forest at Meru, and now mounted in the
National Museum, resembles ruwenzorit more closely than
any other specimen in the collection. It shows strong
evidence of the same large spots and reticulated ground-
color, but the eyes of the rosettes have the appearance of
being larger than in ruwenzor1i, giving a less blackish tone
to the general color. The two specimens have the same
bodily size, judging by the skulls which are quite identical
in dimensions. The similarity in coloration is due doubt-
less to the similarity in environment, both being forest-
dwelling mountain inhabitants.

Nite LEoparp
Felis pardus chut

Native Names: Bari, koka; Acholi, kwat; Aluru, kwats.
Felis pardus chui Heller, 1913, Smith. Misc. Coll., vol. 61, No. 19, p. 6.

RancE.—Lowlands of the Nile from the Albert Nyanza
northward to the Sobat and Bahr-el-Ghazal River systems;
limits of range unknown.

The Nile leopard is distinguishable from the East
African by the fewer number of spots, the wider inter-
spaces between them, the greater extension of the white
ground-color of the under-parts to the upper surface of the
hind feet, and the ventral surface of the tip of the tail, the
larger body size, and larger skull. There is a tendency in

240 AFRICAN GAME ANIMALS

the spots to become completely ringed or ocellated. Quite
a number of such occur scattered over the dorsal surface
among the more numerous rosetted spots. The West
African race of the leopard, leopardus, differs by having
actually as well as relatively larger cheek teeth, more
numerous spots, and smaller body size. The leopard of
upper Egypt, which should stand as the typical race, instead
of the Indian, according to Oldfield Thomas, is a smaller-
spotted race than chut, with a bright ochraceous-tawny dor-
sal ground-color. The type specimen was caught at Gon-
dokoro by Heller, who also secured another at Rhino Camp,
on the west bank of the Nile. The dorsal body color is
ochraceous, paling on the sides to buffy and on the belly
and the under-parts of the legs to pure white. The spots are
black-rosetted, the centre being usually darker-ochraceous
than the ground-color and the margin in some forming a
complete ring without any breaks, being distinctly ocellated
in character. The spots on the under side of the body and
on the legs are generally solid black without lighter central
portion. The spots on the head and the sides of the face
are broken up into numerous small blotches. ‘The tail is
marked basally with rosetted spots which become length-
ened into solid streaks on the middle portion and terminally
have a ringed appearance with much white from the under
side showing-on the sides and nearly separating the black
rings of the extreme tip.

The measurements of the type in the flesh were: head
and body, 49 inches; tail, 33 inches; hind foot, 10 inches;
ear, 34% inches; skull, greatest length, 93 inches; breadth,
5% inches. The Rhino Camp specimen exceeds the type
somewhat in size, which would indicate a really large
average size for the race. Rowland Ward records an ex-
ceptionally large leopard skull taken in the White Nile
region by the game warden of the Soudan, A. L. Butler,
measuring 10% inches, which doubtless represents this
race. Judging by the dimensions of this skull, the Nile
leopard is an animal quite equal in size to fortis, the high-
land leopard of British East Africa.

Pah LEOPARD AND THE CHEETAH 241

HIGHLAND LEOPARD
Felis pardus fortis

Felis pardus fortis Heller, 1913, Smith. Misc. Coll., vol. 61, No. 19, p. 5.

Rance.—Mau highlands of British East Africa; limits
of range unknown.

The highland leopard is a large race, which attains the
maximum size, the skull exceeding in length that of any
other African or Asiatic race. ‘The skull is further distin-
guishable by its narrowness, the small size of the tympanic
bull2, and the absence of the first upper premolar tooth.
The body size is large, with long pelage, dark coloration, and
numerous small, rosetted spots, the central color of which
is not differentiated in shade from the general ground-color.
Fortis resembles in color suahelica most, but is easily distin-
guishable by the darker ground-color, which is uniform in
shade with the central portion of the rosetted spots. Male
skulls of swahelica differ by their smaller size, the largest
being seven-eighths of an inch less in length than the type
of fortis, decidedly larger bulla, and by the presence of a
well-developed first upper premolar. The bullz in fortis do
not rise to the level of the mastoid process, the skull when
placed on a level resting upon the bulle; but in suahelica
they extend well beyond the tips of the mastoid.

The ground-color of the type on the median line of the
back is cinnamon-brown, paling on the sides to ochraceous-
tawny, and on the under-parts and inside of limbs to whitish
or pale buff. The rosetted spots on the back are small and
broken into two or three sections, the central portion being
uniform in color with the cinnamon-brown ground-color.
On the sides of the body the central portion is darker than
the ground-color, as usual among leopards. The under-
parts, legs, head, and basal part of the tail are marked by
solid black spots, the terminal portion of the tail being
solid blackish with a narrow median whitish band on the
under side. Ears tawny, like the ground-color of the head,
and marked by a broad band of black across the middle of
the back.

The type is without flesh measurements, but the skull

242 AFRICAN GAME ANIMALS

indicates a very large race, almost equalling a lioness in
size, and measures, greatest length, 104 inches; greatest
breadth, 9; inches. No other specimen closely approx-
imating the type in size and skull characters has been de-
tected in the collections of the United States National,
British, or Berlin Museums. We may possibly be dealing
here with an abnormal or freak specimen, but the limits of
the individual variation in leopards, as shown by the large
series examined, show that this specimen is well beyond
such limits in its peculiarities.

THe CHEETAH
Acinonyx

Acinonyx Brookes, 1828, Cat. Anat. and Zool. Mus., Joshua Brookes,
London, p. 333 type 4. venaticus of India.

The cheetah is strikingly like a greyhound in body form
as well as in its non-retractile claws. It is the only living
member of the cat family that has succeeded in breaking
away from the typical cat type. Although the long, slender
racing body and canine claws give it conspicuous external
characters, the skull shows no less marked peculiarity com-
pared to that of the true cats. ‘The skull is extremely high
and dome-shaped, with a much shorter jaw than any of the
cats, and is furnished with short canines and very narrow
carnassial teeth which have lost the large inner cusps so
conspicuous in cats and consequently the crushing power
of the teeth is less. The skull of the lynx approaches more
closely that of the cheetah than any other of the cats, but
it is more distinctly elongate and decidedly lower or less
dome-shaped, with typically feline teeth. The houndlike
form of the cheetah has been brought about by similarity
of habit and is not in any way due to closer blood-relation-
ship with the Canide, for the cheetah is much more widely
removed by its skull structure from dogs than such typical
cats as the lion and the leopard. The cheetah may be
taken as an excellent illustration of the effect of habit on
body form. In general style of coloration the cheetah re-
sembles the serval cat, with which it shares much of its
territory. Both are marked uniformly by solid black spots

ae LEOPARD AND THE CHEETAH 243

which show no tendency to form rosettes, ocellated spots,
cross-bands, or stripes. The female is slightly smaller than
the male, the sexual discrepancy in size being much less
than in the true cats. No color differences are apparent
between the sexes. Newly born young have long, woolly
hair and show little evidence of spotting or color marks
except the dark tail rings and tear stripes on the face.
When half grown, at an age when they are shedding their
milk molars, they are still somewhat woolly but are spotted
by small black spots quite uniformly over the back. The
spots on the feet, throat, and base of the tail show a distinct
tendency at this age to form cross-bars as in the African
wildcat, F. ocreata. In a fossil condition the cheetah is
known from the Pliocene of India. At the present time a
single living species, 7uwbatus, is known which covers an ex-
tensive geographical area ranging from the Cape of Good
Hope north through eastern Africa and the Nile Valley to
Arabia, Persia, and India. It is lacking from the Congo
forest area and the wooded portion of the west coast of
Africa and from all forest tracts within its general range.
The cheetah, like the other large cats, shows considerable
geographical variation in the coloration of its coat and may
be divided into several recognizable races on color charac-
ters. Authors have, however, generally failed to find dis-
tinctions between the Indian or the African. Much of the
failure to recognize such long-known or described races as
gubatus and venaticus is due to the absence of specimens
for comparison. Wild-killed cheetahs of the Indian race,
venaticus, are very rare in collections, most of the specimens
representing this race being captive ones used for hunting,
which have lost their slight color characters through the
changed environment.

The Indian cheetahs may, as a rule, be distinguished
from the African by the absence of the short mane on the
neck and the ruff on the under side of the tail, the smaller
body size and less numerous spots, the small, interspaced
spots between the larger being absent. The African animal
splits up into at least four fairly well-defined geographical
races. In the Cape region occurs a small-bodied form, ju-
batus, rather like the Indian, with the spots well separated,
their color not predominating over the ground-color. Far-

244 AFRICAN GAME ANIMALS

ther north, in the equatorial highlands, this race is replaced
by a large-bodied form, velox, with more black spots and
darker coloration. The arid regions of the White Nile and
Lake Chad are occupied by a small-spotted, light-colored
race, semmeringi1, which shows a decided pinkish tone to
the coloration. The lower altitudes of East Africa and
Somaliland support a similarly pinkish-buff animal, but in
this race, rainey1, the spots are much larger and cover an
area fully as great as the ground-color.

The cheetah is fairly common in East Africa. It is
diurnal and is found on the bare plains; therefore, it is much
more often seen than the leopard where both are equally
abundant. It has been suggested that the lion has lost his
spots because he grew to frequent open country where he
no longer needed them, as do the spotted cats which dwell
in the jungle. Unfortunately for this theory, the cheetah,
a much more specialized cat than the lion, dwells even more
in the open, and nevertheless is spotted, while the mono-
colored cougar, a beast of the woodland, is as difficult to see
there as any spotted cat. The cheetah lies up in clumps of
bush or tall grass, and roams over the plains after its prey.
In the sunlight, at a little distance, the spots cease to show,
and its color seems like that of a lioness.

The cheetah is far less ferocious and formidable than the
leopard. It ordinarily preys on the smaller antelopes, such
as gazelles, steinboks, and oribis, up to animals the size
of a half-grown hartebeest; yet there are well-authenticated
instances of its killing koodoo and waterbuck. It kills its
game in fair chase, for it is the fastest animal on earth for a
quarter of a mile, or perhaps half a mile. It stalks up to
within a hundred yards or so, and then gallops into the
fleeing quarry. But it is winded by any long run. If it is

RAINEY CHEETAH, ADULT MALE

From Kapiti Plains, B. E. A.
United States National Museum

RAINEY CHEETAH

From Nairobi
Presented by W. N. McMillan to the National Zoological Park, Washington, D. C.

RAINEY CHEETAHS

Pre LEOPARD AND ATE CHEETAH 245

within half a mile of cover which it desires to reach, and a
horseman seeks to cut it off from one side, it will go two feet
to his one, and will probably reach the cover in safety. But
in a stern chase a hard rider on a good horse will run it down
in a mile or two. Kermit ran down seven in this manner.
Seemingly the cheetahs, on these occasions, never put out
their full speed, as when seeking to catch fleeing quarry.
They were content to gallop easily ahead of the galloping
horse, without spurting, until their wind was gone and it
was no longer possible to spurt. After a mile or two of
fast galloping, according to the start the cheetah had ob-
tained, the horse would be close up. Then the cheetah
would suddenly crouch flat on the ground. Apparently
when it did this it was completely done up, and it made no
further effort either to escape or charge, and was shot as it
lay. It is very rare for one to charge; but one did charge
Stewart Edward White, and with the utmost determina-
tion, too.

Cheetahs are easily tamed, and make good-humored
pets. At Juja farm there were three, which Mrs. McMil-
lan used to lead about witha leash. They did not lead well,
but were friendly and unconcerned. When sitting down
they were so erect that they looked like monkeys rather
than dogs or cats. They uttered a very peculiar, bird-like
chirp, twisting their lips; when we first heard the chirp we
could not believe that it came from the cheetahs, and looked
everywhere for the bird.

246 AFRICAN GAME ANIMALS

Key To THE Races oF jubatus

Ground-color of body buff without pinkish suffusion; spots on hind
feet well marked; dark spots on dorsal region numer-
ous, predominating in area over the ground-color;
pelage longer velox

Ground-color light, pale pinkish-buff; spots on hind feet small or
indistinct; dark spots on back fewer, the ground-
color predominating; pelage shorter

Dorsal dark spots larger, 34 inch in diameter; ground-color lighter
than ochraceous; hind feet spotted
raineyt
Dorsal dark spots smaller, 1% inch in diameter; ground-color
ochraceous; hind feet unspotted semmeringit

HIGHLAND CHEETAH
Acinonyx jubatus velox

Native Name: Masai, engingylasho.
Acinonyx jubatus velox Heller, 1913, Smith. Misc. Coll., vol. 61, No. 19, p. 7.

RancE.—From the Rift Valley and Laikipia Plateau
westward over the highland country as far as the Nile low-
lands; north to the latitude of Nimule and south through
German Fast Africa.

The highland cheetah is characterized by tts large, close-
set black spots, which predominate over the ground-color in
area, by the boldly spotted hind legs, long pelage, and large
body size. The race has been recently described from a
specimen shot by Kermit Roosevelt on the Loita Plains
of British East Africa. The typical race, jubatus, which
inhabits South Africa, differs by having darker ground-color,
smaller spots, and smaller body size. The two lowland
races, semmeringi1, of the Soudan, and raziney1, of the coast
of East Africa, are both distinguishable from velox by their
pinkish-buff coloration and shorter pelage.

The dorsal ground-color of adults is ochraceous, deepest
on midline of back and palest on sides and belly, where it
shades into cream-buff. The black spots on the back are
circular in outline, measuring three-quarters of an inch in

THE LEOPARD AND THE CHEETAH ‘247

diameter, and are interspersed with numerous smaller black
spots, causing the black color to predominate over the
ground-color. The spots on the sides of the body are larger
but less numerous and cover a smaller area than the
ground-color. The legs and under-parts are marked by
oblong black spots. The feet are marked by irregular black
spots, the forefeet more numerously than the hind, and the
bases of the toes also show spots on their dorsal surfaces;
but the hair on the under side is dusky-brown. The tail is
marked by large black spots above and black rings below,
with the terminal portion ringed all around with broad
black and narrow pale buffy rings, the tip whitish. The
crown of the head and the nape are marked by small black
spots as far as the interorbital region. The whole snout
from the eyes is uniform ochraceous and banded on the
sides by a heavy black tear band from the eyes to the
mouth. ‘The sides of the head from the eye to the base of
the ear are marked irregularly by small black spots, with-
out, however, any suggestion of a line from the eye to the
ear. The chin and upper throat are whitish or cream-buff
in color. The backs of the ears are chiefly black, only the
tips and the inner sides being buffy. The hair of the nape is
long, forming a mane from the head to behind the shoulders.
The hair on the median line of the breast and belly is long
and mane-like in character. The ventral surface of the tail
is marked along its whole length by a broad mane or ruff
of longer hair, which at the tips becomes somewhat bushy
in character.

The flesh measurements of the typical male which has
been used as the type specimen were: head and body, 51
inches; tail, 29 inches; hind foot, 11% inches; ear, 3%
inches. The flesh measurements of adult female specimens
fall very slightly below these dimensions. The skulls of
adult males range in length from 7 to 77 inches, those of
females from 634 to 7 inches. The skull dimensions ex-
ceed considerably those from any other part of Africa, which
would indicate a relatively larger body size than in other
races,

A series of specimens have been examined at the National
Museum from the Loita Plains, the Laikipia Plateau north
of Mount Kenia, and the Uasin Gishu Plateau. The range

248 AFRICAN GAME ANIMALS

of this race no doubt extends northward over the highlands
of Elgon to the latitude of Nimule and westward and south-
ward over the higher parts of Uganda and German East
Africa.

Rainey AFRICAN CHEETAH

Acinonyx jubatus raineyt

Native Name: Somali, wer.
Acinonyx jubatus raineyi Heller, 1913, Smith. Misc. Coll., vol. 61, No. 19,

P- 9:

RancE.—British and German East Africa from the
coast lowlands inland as far as the base of Mount Kenia
and the Kikuyu Escarpment, north to Somaliland, and
south to central German East Africa.

The cheetah inhabiting the coast-drainage area has been
recently described from a specimen shot by Paul J. Rainey
near Ulu Station, Kapiti Plains district, British East Africa.
This race is a pale-colored, short-haired form having a light
pinkish-buff dorsal ground-color. It may be distinguished
from the Soudan race, semmeringzi, by the large size of the
dorsal spots and the spotted hind feet. ‘The ground-color
is pale pinkish-buff, darkest on midline, where it is ochra-
ceous-buff, and paling on the under-parts to cream color.
Body marked uniformly by round black spots three-
quarters of an inch in diameter, interspaced by occasional
small spots or dots. The spots on the legs and belly are
elongate in shape and cover the limbs to the toes. The fore-
feet are much more conspicuously spotted than the hind,
which have the spots small or indistinct near the toes. The
toes are much spotted above on a pinkish-buff ground
color, but below they are clothed by dusky-drab hair.
Tail spotted like the back with the terminal portion marked
by five black rings and the extreme tip white. The top of
the head and the nape are marked by numerous small
black spots, but the snout is uniform ochraceous-buff with-
out spots, and is sharply defined on the sides by a black
stripe extending from the eye to the mouth just behind the
whiskers and thence passing backward to the angle of the
mouth. The sides of the head behind the eye and below
the ear are marked by a few black spots, but no evident line

THE LEOPARD AND (THE "CHEETAH 249

between the eye and base of the ear is thus formed. The
ears are pinkish-buff marked by a wide black band covering
the lower half of the back. The chin and the upper throat
are white. The pelage is short on the dorsal surface, being
seven-eighths of an inch in length on the rump. The nape
is furnished with a short mane three inches wide extending
from the head to behind the shoulders. The midline of
the breast and the belly and the whole length of the ven-
tral surface of the tail is further furnished by a ruff or short
mane of longer fluffy hair.

In body size and proportions this race is quite identical
with the highland race. An adult male from Ulu Station
had the following flesh measurements: head and body, 49
inches; tail, 3014 inches; hind foot, 12 inches; ear, 3 inches.
Skull length in this specimen is 71% inches. Specimens
have been examined, from the Kapiti and Athi Plains, at
the National Museum. Others are recorded by sportsmen
from the flanks of Kilimanjaro, near Taveta, and also on
the Upper Tana River southeast of Mount Kenia, north
of which point they are known to occur in parts of Somali-
land. Cheetahs have been reported from the Rift Valley
region of central German East Africa, which are doubtless
referable to this race.

SouDAN CHEETAH

Acinonyx jubatus semmeringit

Cynailurus semmeringit Fitzinger, 1855, Sitz.-Ber. Math. nat. cl. d. K.
akad. Wiss., XVI, hft. 2, p. 245.

Rance.—Lowlands of the Nile Valley, from the Albert
Nyanza northward to Kordofan and westward to Lake
Chad and northern Nigeria.

The German naturalist Fitzinger, in 1855, described the
Soudan cheetah from a specimen secured on the Bajuda
Plains, Kordofan, naming it for the noted physiologist
Samuel Thomas von Soemmering, of Frankfort-on-the-
Main. The race may be recognized by the small size of the
black spots, which do not exceed a half inch in diameter,
and are widely separated by the ochraceous ground-color and
the absence of spots on the hind feet. A specimen from the

250 AFRICAN GAME ANIMALS

White Nile, purchased at El Dueim, has been examined in
the National Museum and another from Lake Chad in the
British Museum. It is a rare animal in the Nile district
and is seldom secured by sportsmen. Schweinfurth fails
to mention it in his list of mammals met with during his
exploration of the Bahr el Ghazal, and Von Heuglin refers
to it very briefly without mentioning any localities. Heller
saw a pair near Gondokoro, but no other member of the
Roosevelt expedition met with this race.

CHAPTER VIII

THE SPOTTED HYENA, STRIPED HYENA, AND
HUNTING DOG

INTRODUCTION To Famity Hyenide

HYENAS

Tue hyenas are at once recognizable by the peculiarity
of their body shape, which is decidedly high-withered, fall-
ing away rapidly toward the hips which are supported
by very short, weak hind legs. The head is somewhat en-
larged but short-jawed and armed with heavy, conical,
crushing teeth. The teeth of the hyenas are highly spe-
cialized for crushing bones, and for this purpose some of
the cheek-teeth have developed into stout, rounded, conical
cusps of great strength. The canine teeth are much smaller
than in either the dogs or the cats, being more fitted for
crushing than for tearing flesh or for grasping prey. In
order to make this formidable dental apparatus of great
crushing value the jaws are armed with enormous muscles,
the bulk of which find attachment on the high median dor-
sal crest of the back part of the skull. The development of
this bony sagittal crest is greater than in any of the other
carnivores and furnishes one of the marked distinguish-
ing features of the skull, which otherwise closely resembles
that of the cats. The feet are four-toed and furnished by
stout, non-retractile canine claws. Large anal glands or
sacks are present, and the testes are internal, but beyond

this similarity there are marked differences in the sexual
251

252 AFRICAN GAME ANIMALS

organs in the two genera comprising the family. The
hyenas, to the casual observer, appear to be fairly close
relatives of the dog family, and few people suspect the really
close relationship they bear to the cat family. The struc-
ture of the skull and the arrangement of the teeth are
strikingly feline, the feet alone being distinctly canine.
Some fossil genera have been discovered which tend to
link the hyenas with the Viverride, of which the well-
known mongoose is a representative, but the evidence is
yet far from satisfactory in this regard. Two strikingly
different groups or genera comprise the family: the striped
hyena, genus Hyena; and the spotted hyena, genus
Crocuta. Hyena, from which the family name has been
derived, is the least specialized of the two. It is character-
ized externally by the long dorsal mane of hair extending
from the nape to the tip of the tail, the large pointed ears,
dark vertical stripes or bands across the body and legs,
and black throat. The sexual organs are normal in struc-
ture, but the skull shows decided dental differences. The
upper molar is well developed and fits into a long heel on
the last lower tooth or carnassial. The skull is somewhat
smaller, narrower, longer-snouted, and the body size less
than in the spotted hyena, or Crocuta. The latter differs
by its shorter hair growth, absence of a well-developed dorsal
mane, by short ears and the spotted type of coloration. The
female sexual organs are quite peculiar in structure and
closely resemble those of the male in external appearance.
The dentition is more reduced, the upper molar being mi-
nute, as in the cats, and the lower carnassial has only a rudi-
mentary heel. The skull is broad, with very broad mesop-
terygoid fossa, and the body size is considerably greater.

EASTERN SPOTTED HYENA, FEMALE

From Forest Station, West Kenia
United States National Museum

HIGHLAND STRIPED HYENA, FEMALE

From Loita Plains
United States National Museum

SPOTTED AND STRIPED HYENAS

2 ae
tna 2

ie Ea YvENA AND THE HUNTING DOG’ 253

StripeD HyEeNA
Hyena

The genus Hyena has at the present time two living
species, the striped hyena of North Africa and Asia and
the brown hyena or strand-wolf of southwest Africa. In
former times, however, the genus was much more widely
distributed, as evidenced by the several fossil species which
have been found in the Pliocene and Pleistocene beds of
southern Europe and Asia. The striped hyena is readily
recognizable by the long dorsal mane, which extends from
the head along the nape and over the back to the tail, and
by the blackish transverse stripes on the body and legs. In
coloration, body shape, and general appearance it is mi-
nutely imitated by the aardwolf, a somewhat smaller car-
nivorous mammal having rudimentary teeth and an insec-
tivorous diet. In the east equatorial regions of Africa the
two animals share the same territory. The close resem-
blance in color pattern appears to be purely an accidental
weatiair and not a case of mimicry. The dentition of the
striped hyena is less reduced than that of the spotted
genus, the upper molar being well developed and func-
tional, and the lower carnassial tooth has a long heel behind
which opposes the molar when the jaws are closed and is
functional during mastication. The female sexual organs
are quite normal in structure, and the relative body size of
the two sexes is that usual among carnivores, the female
being somewhat less in size than the male. No sexual
color differences are apparent. In German and British
East Africa the striped hyena reaches its southern limit
of distribution and is there represented by two races, a
highland form and a low desert form.

This hyena was not uncommon in certain parts of East
Africa, but we never saw it abroad in the daytime, and
only caught it in traps. It seemed more nocturnal, shyer,
and less bold and ferocious than its big, spotted brother.
There are spotted, striped, and brown hyenas. Their hab-
its are all substantially alike, and their ranges overlap. In

254 AFRICAN GAME ANIMALS

view of their habits it seems quite impossible that the col-
oration of their coats can have even the smallest effect
upon their well-being; and it is absolutely impossible that
the totally different types of coloration of the three species
can all have the same effect from the standpoint of conceal-
ment. It seems probable that these totally different col-
oration patterns have been developed without reference
to the concealing quality of any of them; and it is cer-
tain that no one of them is now of more value than the
others in the animal’s life, from the standpoint of con-
cealment.

Key To THE Races OF hyena

Dorsal ground-color pale-buff without pinkish suffusion; stripes
black and broader; mane tipped with black; feet uni-
form dark-brown schillingst

Dorsal ground-color pale pinkish-buff; stripes dark seal-brown and
narrower; mane tipped with dark-brown; feet buff or
ochraceous mottled with brownish-drab bergert

HIGHLAND STRIPED HYENA
Hyena hyena schillingst

Native Name: Swahili, kungugua.
Hyena schillingsi Matschie, 1900, Sitz.-Ber. Ges. Nat. Freunde, Berl., p. 55.

Rance.—Kilimanjaro region and Rift Valley of German
East Africa, northwest to the Victoria Nyanza, Loita Plains,
and Kedong Valley in British East Africa.

The striped hyena is a rare animal in East Africa, and
is seldom met with by the sportsman. Willoughby and
Hunter saw specimens during their trip to Kilimanjaro in
1887, and were the first to report its occurrence, but it was
not until a decade later that specimens were actually se-
cured by Schillings, for whom the race was named by Mat-
schie in 1900. More recently specimens have been secured

THE HYENA AND THE HUNTING DOG 255

by various expeditions on the Loita Plains, the Kedong
Valley south of Lake Naivasha, and at Voi.

The highland race of the striped hyena shows a grayer
coloration and more distinctly striped coat than the desert
race. The ground-color is pale-buff without any pinkish
suffusion, the stripes are quite blackish, the feet are uni-
form dusky-drab, and the dorsal mane is tipped by black.
The body size is somewhat larger than the other equatorial
races. An adult female of this race secured on the Loita
Plains by the Roosevelt Expedition and now in the National
Museum had the following flesh measurements: head and
body, 43 inches; tail, 1134 inches; hind foot, 814 inches;
ear, 6 inches. Greatest length of skull, 954 inches.

DeEseRT STRIPED HyENA
Hyena hyena bergert

Native Names: Somali, whera or didar.
Hyena hienomelas bergeri Matschie, 1910, Sitz.-Ber. Ges. Nat. Freunde,
Berl., p. 361.

Rance.—Desert region of British East Africa from the
southern slopes of Mount Kenia and the Mau Escarpment
northward throughout the Lake Rudolf basin, southern
Abyssinia, and Somaliland.

A specimen of the striped hyena from the Elgeyo
Escarpment in the Uasin Gishu Plateau region has been
made the type of the desert race by Matschie. The race has
been named for its collector, Doctor Berger, who has given
us “In Afrikas Wildkammern,” an excellently illustrated
account of his hunting expedition through East Africa and
Uganda. The desert race may be distinguished from the
highland race by its pale-pinkish ground-color, less con-
spicuous stripes, brownish tips to the dorsal mane, light-
colored buff feet, with little or no dusky-brown mixture,
and smaller body size. Considerable individual variation
in color is shown by a series of skins from the Northern
Guaso Nyiro River in which the feet vary from pale-buff to
wood-brown and the hair tips of the dorsal crest from dusky-
brown to seal-brown or blackish. The dorsal stripes also
vary in intensity, but they are never as heavy or black as

256 AFRICAN GAME ANIMALS

in the highland race. The tail varies much in length,
ranging in adults from 1214 to 14% inches. The skulls
show considerable variation in size and shape, quite equal-
ling in this respect the spotted hyena. In a series of
seven female skulls the range in length is 8% to 834 inches,
and in breadth 53 to 57% inches. The male skulls show
less variation, being usually 9% inches long by 6 inches
wide. The flesh measurements of an old male from the
Northern Guaso Nyiro River were: head and body, 42
inches; tail, 1334 inches; hind foot, 814 inches; ear, 534
inches. The females average somewhat less than these
dimensions.

Many years ago Latreille described a race, hienomelas,
from the Atbara River or northern slopes of the Abyssinian
highlands, which may upon the study of further speci-
mens be found to be indistinguishable from the present |
race, in which case the latter name would give way for the
older one, proposed by Latreille. Recently Lonnberg has
given a new name to the striped hyenas of the Northern
Guaso Nyiro River, but the large series in the National
Museum from that locality are quite indistinguishable in
color or other characters from the description of bergeri, the
type of which was secured at the extreme southwestern
limit of the race, near the base of the Elgeyo Escarpment.

SPpoTTED HyENas

Crocuta

Crocuta Kamp, 1828, Oken’s Isis, XXI, Heft XI, p. 1145; type Hyena
crocuta.

The spotted hyenas have by most writers been com-
bined with the striped in a common genus, Hyena. A
careful comparison of the two groups, however, leads to the
discovery of many differences of a fundamental character
and shows a really wide gap in structure between the two
hyenas. There are decided differences in dentition, in
skull shape, in the genital organs, in body shape, and hair
growth, any of which are of generic importance. ‘The
spotted hyena has the dentition much more reduced, the
molar being a mere rudiment or missing altogether, while

Tee TYENA AND. THE HUNTING DOG. 257

the lower carnassial tooth lacks a well-developed heel. The
skull is wider, shorter-snouted, and has a narrower mesop-
terygoid fossa. The sexual organs of the female are very
peculiar and resemble in external appearance those of the
male—a condition quite unique among carnivorous mam-
mals. It is owing to this peculiarity that the animals are
believed by most of the negroes and many sportsmen to
be hermaphrodite. Externally, the spotted hyenas differ
from the striped by the small size of the ears, the absence
of a dorsal mane of long, stiff hair extending from the nape
to the tip of the tail, and the spotted character of the coat.
In the spotted hyena the female exceeds the male in size, a
sexual peculiarity not found in the striped hyena or in any
other existing carnivore. The young at birth are uniform
seal-brown, without any indication of spots or markings of
any sort. After a few months have passed, their coloration
becomes paler in spots and assumes gradually the heavily
spotted coat characteristic of the immature. Two young are
produced ata birth. The genus Crocuta is much more highly
specialized than Hyena, and is quite catlike in dental char-
acters and skull shape. A single living species, crocuta, is
known, which is confined in distribution to Ethiopia or Af-
rica south of the Sahara Desert. Several geographical races
have been described; but, owing to the great individual
variation to which some of them are subject, their status is
at present quite uncertain. During the Pleistocene age a
species very closely related to the existing one occurred in
southern Europe as far west as the British Isles and also
on the Mediterranean coast of Africa and eastward as far
as India. :

The spotted hyena was common through East Africa
and Uganda. Its strange cry was heard on most nights,
wherever we were, and we often came across it in the day-
time; for, although most active after nightfall, it frequently
prowled abroad in the mornings and afternoons, in lonely
lands. Generally, however, in the daytime it retired to its
lair, which might be a cave or a hole under a stone, or

258 AFRICAN GAME ANIMALS

merely a patch of reeds or bush. Normally it was solitary,
but quantities assembled around any big dead beast. Ker-
mit rode down several, although sometimes, if the beast had
a long start, only after a good chase. Hyenas are noisy;
besides the usual questing cry, they at times cackle and
screech and more rarely laugh.

They are queer creatures. Ordinarily they are scaven-
gers, feeding on carrion. Yet they occasionally kill mules,
cattle, donkeys, and young rhinoceros. Their jaws are so
powerful, their strength is so great, and they are so tough,
that only a number of big, fierce dogs can kill one. When
hungry they pounce on and carry off dogs. Yet when
hunted they are very cowardly. A big Airedale terrier, be-
longing to a ranchman we visited, would unhesitatingly
attack a hyena single-handed, and harass the beast so that
the hunter could come up and shoot it. Toa man there is
absolutely no danger in hunting them; the utmost they will
do is to snap at a stick when wounded and at bay. They
have not a thousandth part of the fury and reckless prowess
of the leopard. Yet they are far more apt than the leopard
to prey on human beings, even full-grown men; and in cer-
tain districts where they are bold and numerous it is dan-
gerous for any man to sleep alone in the open. Their at-
tacks are always made at night, with extreme caution, and
when the victim is sleeping. Usually they seize the face,
and we saw more than one native who had been frightfully
scarred in this manner, although the beast had been driven
off. An English officer, Major Coryndon, was once seized
by the hand by a hyena while he was sleeping in the open;
he struggled so that the beast left him and ran off into the
darkness; whereupon he lay down again with his loaded rifle

THE HYENA AND THE HUNTING DOG _ 259

ready, and when his hungry assailant returned he shot it
dead. In Uganda, when we were passing through, the
hyenas had become confirmed man-eaters because of de-
vouring the innumerable natives who died of sleeping sick-
ness. ‘They haunted the ghastly sick-camps into which the
officials gathered the sufferers. After a few experiments
they found that the dying men and women could offer little
resistance, and after nightfall they broke into their huts
and carried them off; and they grew steadily in numbers
and ferocity. It became necessary not only to build stiff
fences round the camps but to station strong guards of
soldiers in them, and every night there were repeated shots
fired in killing or driving away the marauders.

In fact, the hyena is a singular mixture of abject coward-
ice and the utmost ferocity. Usually feeding on carrion,
and often hesitating to attack even the weakest animal if
it is unhurt and on its guard, the ravenous beast will, on oc-
casions, even when single but especially when in troops, as-
sail very formidable creatures. A troop has been known to
kill a half-grown rhinoceros which had returned to the body
of its mother, slain by hunters. Not only sheep and goats
but donkeys, mules, cattle, and dogs are at times destroyed.
In killing men, women, and especially children, a man-
eating hyena will penetrate big villages; one took a native
from a hut in Nairobi itself. When in troops they have been
known to seize animals that have been wounded by hunters,
and to attempt to stand off the hunters. We have never
known them in such a case actually to attack the hunters.
But under certain circumstances they do attack lions,
which seems quite as extraordinary. Ordinarily the hyenas
merely attend the lion at respectful distance, eager to get

260 AFRICAN GAME ANIMALS

whatever he leaves, and they occasionally pay with their
lives if they grow too impatient. But Donaldson Smith has
related an instance in which a pack of hyenas entered into
a regular fight with a couple of lions, near a bait, and got
the best of it. Once we were sleeping near the body of an
elephant. There were many hyenas feeding at the car-
cass. Toward morning a lion came near, and menaced the
carrion feeders, uttering a long-drawn, moaning sigh. The
hyenas, however, refused to be frightened by the self-
invited guest, and they responded to his challenge with the
most extraordinary chorus of noises to which we ever lis-
tened. They refused to leave the carcass, or to admit the
lion to his share of the banquet; and the lion made off with-
out venturing to try conclusions with them. More extraor-
dinary still is the fact that the hyenas sometimes kill and
devour very old and crippled lions. Once Lord Delamere
shot and very badly wounded a lion, which in the course
of the fray mauled him and both his gun-bearers. Camp
was pitched where the wounded men lay, while the lion,
too badly hurt to leave, crawled into some bushes but a
short distance off. After darkness the hyenas assembled,
fell on the wounded lion, and after a very noisy fight killed
and ate him.

Hyenas are easily tamed, and in many cases are docile,
intelligent, and affectionate. At the Cairo zoological park
we saw one which was as devoted as a dog to its keeper. It
loved to be stroked, and would lie belly upward while he
rubbed it. In ancient Egypt they were tamed and used
with greyhounds for the chase; and, extraordinary to relate,
they were also fattened for the table, being stuffed with
food like so many Strasburg geese.

THE HYENA AND THE HUNTING DOG 261

KeEyY To THE RACES OF crocuta

Pelage long, decidedly more than one inch in length on the rump;
spots large on the back
Ground-color ranging from buff to tawny, seldom showing a
gray cast germinans

Ground-color pale-gray thomasi

Pelage short, less than one inch in length on the rump; spots smaller on
the back; ground-color pale-buff fist

EASTERN SPOTTED HYENA
Crocuta crocuta germinans

Native Names: Swahili, fisi; Masai, ol ngojine.
Hyena germinans Matschie, 1900, Sitz.-Ber. Nat. Freunde, Berl., p. 26.

RanceE.—Highland and coast drainage area of British
and German East Africa from Mount Elgon southward
throughout the whole of German East Africa to Nyasaland
and the Zambesi drainage.

The spotted hyena inhabiting German East Africa was
named by Doctor Matschie germinans, owing to the great
individual variation which they exhibit in coloration. This
color variation has been observed and mentioned by almost
every sportsman who has recorded his observations of
spotted hyenas from eastern equatorial Africa. The
typical specimen upon which Matschie based his name
came from Lake Rukwa, a small body of water lying be-
tween Lakes Tanganyika and Nyasa on the western fron-
tier of German East Africa. In 1908 Lonnberg twice re-
described the present race, giving distinct new names to the
two extreme types of coloration observed in specimens
from the southern slopes of Mount Kilimanjaro. One of
these names, kibonotensis, he applied to a reddish animal
having a short tail, the other, panganiensis, to a grayish
one with a longer tail. These two color and tail extremes,
together with every intermediate stage, are represented in
specimens from the Loita Plains and the Uasin Gishu
Plateau in the National Museum collection. ‘They will be

262 AFRICAN GAME ANIMALS

found to occur in any large series from any locality through-
out the distribution of this hyena and are to be regarded
simply as evidence of the variability of the race.

It is difficult to define a race so variable as germinans
in color and proportions. The ground-color of the upper
parts ranges from a light-buff through ochraceous to tawny,
and occasionally a light-grayish cast also occurs, but the
usual ground-color may be said to be ochraceous-buff.
The dark spots are fairly numerous and large on the back
and occupy an area quite equal to the ground-color in
extent. A few large spots, two or three inches in diameter
and several times larger than the others, are usually pres-
ent on either side of the midline near the middle of the back.
The feet and lower part of the legs also show much varia-
tion, ranging in color from clove-brown to pale-buff. The
young and immature animals are dark-colored, with large
spots and dark feet, and are distinctly uniform in appear-
ance, giving no suggestion of the great variation which
occurs in the adult. Other races of the spotted hyena do
not appear to share this extensive color variation of ger-
minans, but have well-established color differences. The
eastern spotted hyena may usually be known from either
fisi or thomasi by the larger size of its dorsal spots, long
coat, and tawny or buffy coloration. From the Cape race
of South Africa it is distinguishable by its smaller body size
and narrower skull.

An adult male from the Loita Plains had the following
flesh measurements: head and body, 50 inches; tail, 11
inches; hind foot, 9% inches; ear, 414 inches. The great-
est length of the skull in this specimen was 1014 inches.
The female averages greater than these dimensions some-
what, the largest specimen examined being a female
from the Loita Plains which has a skull length of 107%
inches. Specimens have been examined in the National
Museum from the Loita and Kapiti Plains, Laikipia and
Uasin Gishu Plateaux, and the Kenia forest on the south
and west slopes of the mountain. The spotted hyena is
universally distributed from the coast as high as the moor-
land at ten thousand feet altitude, and is absent only from
the inner depths of forests and from the waterless desert
tracts.

THE HYENA AND THE HUNTING DOG 263

MarsaBiT SPOTTED: HyENA
Crocuta crocuta fist

Crocuta crocuta fist Heller, 1913, Smith. Misc. Coll., vol. 61, No. 22, p. 5.

RancE.—Desert lowland region from the Northern
Guaso Nyiro River drainage north through the Lake Ru-
dolf region to Abyssinia and eastward to the coast.

Specimens of the spotted hyena inhabiting the low,
arid desert region of the Northern Guaso Nyiro and Mount
Marsabit region are decidedly shorter-haired, lighter-col-
ored, and smaller-spotted than the variable germinans
of the highlands to the southward. This hyena has
recently been described as fist by Heller from specimens
from the Merille water-holes situated in the desert midway
between the Guaso Nyiro River and Mount Marsabit. It
is further distinguishable from the Somaliland race, ruji-
picta, also a desert inhabitant, by the reddish color of the
Bee and the more buffy or ochraceous ground-color of the
atter.

The ground-color is pale-buff, spotted irregularly over
the back by rather small black spots varying in diameter
from a quarter to three-fourths of an inch and covering an
area only about one-fourth that of the ground-color. The
spots are heaviest and most distinct on the midline of the
back and faintest on the neck, sides of the body, under-
parts, and shoulders, but the head and the feet are un-
spotted. The tail is basally like the back in color, with a
few dark spots, the terminal one-half being covered by a
bushy black tuft of long, coarse hair. The feet are uni-
form olive-brown to the toes, in contrast to the spotted
upper portions of the legs. The head is grizzled-buffy and
blackish on the crown and sides, the snout, muzzle, lips,
and chin being uniform dusky-black in contrast. The
whiskers and the long hairs above the eyes are black. The
throat and the sides of head below the ears are uniform light-
buff without darker spots. The back of the ears show a
grizzled appearance, due to the scantiness of the short
buffy hair which allows the blackish skin to show through,
but the ears on the inner side are more heavily clothed by
longer whitish hair. The hair is short, the length on the

264 AFRICAN GAME ANIMALS

rump being only one-half of an inch. The nape is furnished
with a short mane of uniform light-buff color in which the
longest hairs have a length of two inches.

The size and relative proportion of parts in /isi are quite
identical to those found in germinans and need not be re-
peated here. A large series of specimens are in the National
Museum, collected by the Rainey expedition in the desert
region between Mount Marsabit and the wagon ford on
the Northern Guaso Nyiro. ‘They are chiefly from the

water-holes of Merille, Longaya, Koya, and Karo, and from
the Guaso Nyiro River near its junction with the Lakiundu.
These specimens show remarkable uniformity in ground-
color and size of spots compared to the variable germinans,
from which they are readily distinguishable by color and
shorter pelage.

ANKOLE SpotrED HYENA

Crocuta crocuta thomast

Native Name: Luganda, empisi.
Crocuta thomast Cabrera, 1911, Proc. Zool. Soc., p. 98.

A specimen of hyena in the British Museum from
Ankole, Uganda, collected by Sir Harry Johnston, has
recently been made the type of a new race by Cabrera and
named for the curator of mammals of that institution,
Oldfield Thomas. ‘The character assigned to the race by
the describer, who had a series of but three specimens, is the
pale-grayish ‘coloration. It is doubtful, however, if the
race is a valid one, being founded on so ‘small a series and
living in close proximity to the highly variable germinans.
More specimens are needed from Ankole to establish the
limits of individual variation in this questionable race.

INTRODUCTION TO FAMILY Canide
Docs anv WOLVES

The only African member of the dog family which at-
tains sufficient size to be classed as big game is the hunting
dog. It is quite the smallest member of the big-game

Peony ENA “AND: THE. HUNTING DOG 265

carnivorous mammals, but owing to its organized ferocity
consequent upon its gregarious habits the species assumes
an importance far beyond what its bodily size would war-
rant. The true dogs of the genus Canis are represented in
East Africa only by the jackals, which seldom exceed the
dimensions of a red fox and live a solitary or at most a
paired existence of rapine on smaller animals.

Huntinc Doc
Lycaon

Lycaon Brookes, 1827, in Griffith’s Cuvier’s Anim. Kingd., V, p. 151; type
Lycaon pictus.

The hunting dog was considered by the first naturalists
to whom it became known, namely, Burchell and Tem-
minck, as a species of Hyena. ‘They were led to this con-
clusion, no doubt, by the spotted coloration, thin coat,
four-toed feet, the broad skull, and short, wide jaws. The
hunting dog, however, is closely related to the typical dogs
of the genus Canis, from which it is separated chiefly by
the four-toed character of the forefeet and slight differences
in dentition. There are no differences in number of teeth
between the two genera, but the lower carnassial tooth
differs from that of the true dogs by having only one cusp
on the heel, a peculiarity which it shares with Cuon, the
dhole of India to which it is most closely related. The
skull of Lycaon is easily recognizable from that of a wolf or
a jackal by its much greater breadth across the palate and
the shortness of the jaws. Externally, the hunting dog ex-
hibits marked peculiarity by its blotched coloration, which
lacks any definite pattern on the body, the three colors of
black, yellow, and white being combined in almost every
conceivable proportion and pattern in different individuals.
The hair covering is usually short and often so scanty that
the black skin forms the predominant color. The head
has, however, a definite color pattern, the snout being black
as far back as the eyes, the rest of the head ochraceous in
color, with a broad black median stripe over the crown and

266 AFRICAN GAME ANIMALS

down the nape. The ears are extremely broad and con-
spicuous, resembling in their great size some of the foxes,
and are black on the back with yellow inner sides. The
tail is of medium length and rather scantily haired, the
terminal tuft of white hair being much smaller than the
brush of a wolf. At the present time the hunting dog is
confined to Ethiopian Africa, where it ranges from the Cape
north throughout the eastern part of the continent to the
limits of the Abyssinian highlands and Lake Chad. For-
merly it ranged beyond the limits of Africa through Europe
to the British Isles, where fossil evidence of it has been ob-
tained in the Pleistocene beds of Wales. A single living
species, pictus, is recognized by naturalists, of which several
geographical races have been named on differences detected
in the proportions of the three colors in the coat, by rela-
tive differences in body size, and thickness of pelage. The
characters given for the various races, however, are based
on the examination of so few specimens that they can in
no case be said to be established, but the study of more
material in the future will without doubt lead to the dis-
covery of definite characters for many of the races now
named. Names have been applied to British East African,
Somaliland, and Lake Chad races, but it seems best at the
present time to recognize only the East African race, which

alone has recognizable characters over pictus of South
Africa.

This pied creature is found throughout East Africa,
although it is rare in most places. Hunting hounds usu-
ally go in big packs. They master all the smaller and the
young of all the larger antelopes, and there is reason to
believe that at times, although rarely, they kill even the
biggest antelope, and half-grown buffalo also. Yet we
saw zebras feeding near them without heeding them, and
also rushing at them and driving them off when they came
too close. They are extremely destructive to game, and at
times to goats and sheep; and they will menace man, al-

EAST AFRICAN HUNTING DOG, MALE
In the New York Zoological Park

ee

EAST AFRICAN HUNTING DOG, MALE
Caught at Merrille near Mt. Marsabit
Presented by Paul J. Rainey to the New York Zoological Park

: HUNTING DOGS

DRE THYENA‘AND THE HUNTING DOG - 267

though we have no authentic instance of their actually
attacking him. But Mr. Rainey, in the Northern Guaso
Nyiro desert, saw a party of wild dogs chasing a lion; they
did not bite him, but he was manifestly uneasy and con-
cerned and trotted sheepishly along, endeavoring to get out
of their way.

East Arrican Huntinc Doc
Lycaon pictus lupinus

Native Names: Swahili, biz1, mbwa-mwitu; Masai, oswyat; Kavirondo,
sudthi; Dinka, kwaty.
Lycaon pictus lupinus Thomas, 1902, Ann. &F Mag. Nat. Hist., TX, p. 439.

RanceE.—East Africa and Uganda from the coast and
desert tracts inland to the high plateaux, lacking only from
the forest and alpine country.

Hunting dogs were met with by some of the first ex-
plorers to enter the upper Nile region. Heuglin reported
them in 1869 from the White Nile, and a few years later
Schweinfurth mentioned them from the upper waters of
the Bahr el Ghazal and the Nile-Congo watershed. Speke
and Grant, however, were less fortunate in this regard
and failed to learn of their occurrence in the region they
traversed. On the East Coast, however, it was not until
comparatively recently, in 1887, that Willoughby first re-
ported the hunting dog from the Kilimanjaro country. It
is now well known that they range from the coast and low
desert country inland to the highlands wherever there is
sufficient game for food. They are nowhere numerous,
but wander about in packs over much territory and are
seldom met with by sportsmen. The East African race
has been described by Oldfield Thomas from specimens shot
by C. S. Betton near the Ravine Station on the headwaters
of the Nyuki River, at the point where it is crossed by the
equator.

A typical specimen of the East African race has the
color on the back irregularly blotched black and ochraceous-
yellow in about equal proportions, with scarcely no white,
and with the back of the ears black. The hunting dog

268 AFRICAN GAME ANIMALS

from South Africa differs by being chiefly ochraceous in
color, the black being but a small per cent of the dorsal
coloration, and the back of the ears differ by being chiefly
ochraceous instead of black. Much individual variation
in coloration, however, exists and no two specimens are
marked on the back quite alike or in the same proportion
with the three colors, black, ochraceous, and white. This
variability in coloration is well shown by a series of six
adults, two females and four males, shot from the same pack
and collected by the Rainey expedition on the Loita Plains.
In two of these specimens the black areas predominate over
the yellow on the body, while in the other four the yellow
is somewhat in excess of the black. One of the specimens
has a white spot on the shoulder, but in the others no white
appears in the coloration. All of the specimens show some
white blotches on the under-parts, particularly the breast,
but the predominant color below is black. ‘The legs are
irregularly striped and marbled on the outer side by white,
the white area being equal to that of the black and yellow
combined. The tail is another extremely variable feature
in color. The white tip varies from a few white hairs to
two-thirds the tail length in extent. The median portion
is usually black, this part varying from a narrow ring one-
quarter of an inch wide to two-thirds the whole tail. ‘The
basal one-third is ochraceous in all the specimens. The
head and nape are the only portions which show a definite
and constant color pattern. The nape and crown of the
head are uniform ochraceous, marked by a wide black
stripe along the median line. The snout from the eyes
forward and the sides, chin, and forethroat are uniform
black, as are also the backs of the ears. The inner side and
the extreme base of the latter are ochraceous, like the
crown and cheeks. Aged specimens often show very little
hair, being quite hairless and slaty-black like Mexican hair-
less dogs. The coat is longest and heaviest in the young
adults. There are no sexual differences in color. Puppies
and immature specimens are quite blackish, with little or no
yellow and whitish except on the terminal tail tuft.

The flesh measurements of an old male from the Loita
Plains were: head and body, 38 inches; tail, 15 inches;
hind foot, 914 inches; ear, 514 inches. The female is some-

Tae HYENA AND THE HUNTING DOG 269

what smaller than the male, and has considerably smaller
teeth. The longest skulls in a series of eleven are: male,
81% inches; female, 77% inches. Specimens have been
examined in the National Museum from the Kapiti and
Loita Plains and Kamiti Farm, British East Africa. Others
have been seen in the Northern Guaso Nyiro and Marsabit
districts, Maji-ya-Chumvi Station near Mombasa and in the
Gondokoro district of northern Uganda.

CHAPTER IX

PIGS
Suide

Arrica is remarkable for the peculiarity of its pigs. It
is a land in which pigs have run riot and evolved such
grotesquely ugly forms as the wart-hog. Three very dis-
tinct genera occur wide-spread throughout the tropical part
of the continent. Two of these, the wart-hog and the
forest hog, have doubtless originated on the continent, while
the third, the bush pig, is of European or Asiatic derivation.
The true pigs as a family are distinguished by the simple
or primitive condition of the feet, which have all four toes
well developed, but the lateral pair show signs of atrophy,
while the middle pair are decidedly enlarged and support
the weight of the body. The canine teeth are enlarged into
cutting tusks, and the tip of the snout is broadened into a
flat disk for rooting. ‘This expanded snout is supported by
a peculiar bone, the prenasal, which lies embedded between
the tips of the nasal and the premaxillary bones. The
stomach is simple, without the complicated chambers of
the ruminants. The pigs are moderate or small-sized un-
gulates, without horns or antlers. They are confined to the
Old World, ranging from the temperate region of Asia south-
ward throughout the tropics and westward through Europe

to the British Isles and south over the continent of Africa.
270

PIGS 271

KrEy TO THE GENERA OF Suid@e

No warts on cheek below the eye; upper tusks short, bent downward,
their points cut obliquely off by wear on the short
lower tusks; body well haired, colors reddish or black,
with a white dorsal mane; skull of boar with a bony
process above upper canine; first lower premolar
tooth wanting Potamocherus

A prominent wart below the eye; upper tusks long, curved upward and
inward, only their lower edge coming in contact with
the cutting edge of the lower incisor tusks

Wart below eye shelf-like and horizontal, not conical, in shape;
body well haired, but dorsal hair not forming a longer
mane; upper canines smaller and less curved inward;
skull with a high lambdoidal crest and a deep hol-
low on crown or parietal region; last molar trans-
versely ridged, not wearing down to a flat surface,
short; head not enlarged Hylocherus

Wart below eye conical, and another on sides of head near mouth; a
prominent ridge, covered by white whiskers, on lower
part of cheek; upper tusks enormously developed and
curved inward and upward in a semicircle; skull
without hollow in parietal region; last molar long,
occupying more than one-half the tooth row, crown
with flat surface; head much enlarged; hair covering
scanty on sides and under-parts Phacocherus

Busu Pic

Potamocherus

Potamocherus Gray, 1852, Proc. Zool. Soc., p. 131; type P. kotropotamus
of South Africa.

The bush pigs are closely related to the genus Sus, of
which our common domestic pigs are familiar representa-
tives. They represent an African branch which has di-
verged slightly from the original Sus stock. The boars
have developed on the skull above the upper canine teeth a
bony growth, or process, which projects upward an inch or

272 AFRICAN GAME ANIMALS

two to the level of the nasal bones, the tip of the process
giving rise on the sides of the snout to a low, hairy wart.
Bush pigs differ further by the loss of the first lower pre-
molar, which in the common pig is a fairly well-developed
tooth lying isolated midway between the second premolar
and the enlarged lower canine tooth. ‘The female differs
from Sus by the lesser number of mammez, which are but
three pairs. In coloration bush pigs may usually be known
by their bright-red color, and on this account they are often
called red river-hogs, a term, however, which is applicable
only to the West African species, porcus. In eastern Africa
the old boars usually lose their red color, which is replaced
by black in adult life, at which time they assume quite a
dark coat, relieved only by the whitish mane and head.
The newly born young are marked by longitudinal stripes
of pale-yellow or buff on a dark-brown ground-color. Bush
pigs are inhabitants of forest or dense bush country and
are found everywhere in Africa where such cover occurs,
from the Cape to the northern frontiers of Abyssinia and
the Soudan, and from sea-level to the upper limits of forest
growth, or to an altitude of ten thousand feet in the equa-
torial highlands. Several fossil species have been found in
the Miocene of India which have a close resemblance to
bush pigs in tooth structure, but the evidence is too frag-
mentary to warrant their identity with Potamocherus,
which is quite unrepresented in the later Tertiary beds of
Asia or Europe. A race known to naturalists as larvatus
exists at the present time on the island of Madagascar,
but as it is scarcely distinguishable in either color or skull
characters from the mainland races, it is doubtless a recent
importation through the agency of man and has no signifi-
cance as showing a recent land connection between Mada-
gascar and the continent of Africa. The mammal fauna of
Madagascar is so peculiar and limited to so few types that
the island must have remained isolated throughout most
of the Tertiary time to have prevented the introduction of
any of the modern African forms of antelopes and car-
nivores. ‘The only other existing African ungulate which
had representatives in Madagascar is the pygmy hippo-
potamus, which was represented by two species during the
Pleistocene age on the island. Two species of bush pig are

PIGS 273

recognizable in Africa, the bright sorrel-red species of West
Africa, called porcus, and the variegated species of eastern
Africa and Madagascar, named by Desmoulins koiropota-
mus, a name usually corrected to cheropotamus. Vari-
ous geographical races have been described of the latter
species, but none of them can be said to be more than pro-
visional, owing to the small number of specimens upon which
they have been founded and the great variability in color-
ation and skull shape to which bush pigs are subject. The
accumulation of further material will no doubt show the
limits of individual and sexual variation in structure and
coloration and lead to the establishment of some of the
races now considered synonymous with the older de-
scribed ones. The territory covered by the present work
is inhabited by two recognizable forms: an East African,
demonis, and an Abyssinian and Nile race, hassama.

Key To THE Races oF koiropotamus

Adult boar coloration a mixture of black and white on body and legs;
hair long and abundant; bony process projecting
above canine tooth long and knobbed, reaching the
level of the nasal bones; body size larger, skull length
more than 13 inches demons

Adult boar coloration uniform black on the sides of the body and the
under-parts, and not showing mixture with white;
hair short, on the sides scanty, allowing the black skin
to appear; bony process above canine tooth short,
only reaching half-way to the level of the nasal bones;
body size smaller, skull length less than 13 inches

hassama

East AFRICAN BusH Pic
Potamocherus koiropotamus demonts

Native Names: Swahili, ngruwe; Luganda, embiz1.
Potamocherus cheropotamus demonis Major, 1897, Proc. Zool. Soc., p. 367.

Rance.—German East Africa northward through Brit-
ish East Africa to the Tana River and Mount Kenia, and
through Uganda as far as the highlands east of Nimule.

Doctor Forsyth Major, in 1897, in his monographic re-

274 AFRICAN GAME ANIMALS

vision of the bush pigs, described the East African race as
demonis, basing his description upon a single adult female
specimen collected by the Reverend W. Morris on the slopes
of Mount Kilimanjaro. The differences assigned by Major
were the narrowness of the skull behind the orbits and the
black coloration of the body, which showed very little red-
dish or tawny hair. A female specimen collected by Doctor
W. L. Abbott on Kilimanjaro is, however, quite a bright
red on the body and is not distinguishable in color from
specimens from the Kikuyu forests of Mount Kenia and
Nairobi. An old boar from Nairobi shows a mixture of
white and black hair on the body, with a whitish dorsal
mane and a white head, practically no reddish or tawny
color showing anywhere. The ears are black and the legs
blackish with a sprinkling of white hairs down the front of
the legs. The female from Kilimanjaro is rufous or fer-
ruginous on the sides of the body, with a white dorsal mane,
black legs, and white head. Specimens from Uganda in the
British Museum are quite like these in general coloration
as well as shape of skull. The Nairobi boar measured in
the flesh: head and body, 51 inches; tail, 14 inches; hind
foot, 10% inches; ear, 5% inches. Greatest length of
skull, 1334 inches. The length of an adult female skull
is 13 inches.

Bush pigs, although quite abundant, are seldom se-
cured by sportsmen, owing to their nocturnal habits and
the dense nature of the cover in which they dwell. They
are more abundant about the edges of native shambas than
in uninhabited regions and have no difficulty in outwitting
the simple devices planned by the natives for their de-
struction. Much damage is done to the native crops by
the night forays of the bush pigs, as they travel in herds of
from ten to twenty individuals or more. Specimens are
exceedingly rare in museums, and in no institution are any
of the races represented by a series of individuals from
the same spot which would make possible the determina-
tion of the limits of individual variation. Single speci-
mens are in the National Museum representing Nairobi,
Kilimanjaro, the Mau forest, and the Taita Hills. A few
other specimens have been examined in the British Mu-
seum from Kilimanjaro, the Tana River, and Uganda.

EAST AFRICAN FOREST HOG

Escarpment Station, Kikuyu Forest, B. E. A.
Mounted group in the American Museum of Natural History, New York.

EAST AFRICAN WART-HOG, MALE EAST AFRICAN WART-HOG, FEMALE
Shot by Theodore Roosevelt, Kamiti Farm Presented by W. N. McMillan to the National
United States National Museum Zoological Park, Washington, D. C.

THE FOREST HOG AND WART-HOG OF EAST AFRICA

i

Ae

ip Pinay ot
- a a \
; ws.

PIGS 275

ABYSSINIAN BusuH Pic

Potamocherus koiropotamus hassama

Native Names: Abyssinian, assame; Argo, askarmar.
Nyctocherus hassama Heuglin, 1863, Nov. Act. Leop. XXX, pt. 2, p. 7.

RancE.—Abyssinian highlands southward through the
Lake Rudolf basin to the Northern Guaso Nyiro River in
British East Africa, and in the Nile Valley southward
through the Bahr-el-Ghazal drainage area.

Ruppell was the first naturalist to mention the Abys-
sinian bush pig, which some years later was described and
named by Heuglin, who applied to it the native Abyssinian
name of hassama. He gave the habitat of his new pig as
the highlands generally, but later in his “Reise in das Ge-
biet des Weissen Nil’’ gave the particular locality of the
species as the Hawash Valley. Major, in his monograph,
has figured the skull of Heuglin’s specimen and called at-
tention to the real characters of the race, the chief of which
is the shortness of the bony process projecting above the upper
canine. Inthe Abyssinian race this process reaches only half-
way to the level of the nasal bones, having only one-half the
development found in the other races. This lesser develop-
ment of the structure, which separates the genus from Sus,
seems to point to the intermediate or less specialized char-
acter of the most northern race and the one geographically
nearest Sus. ‘There is also a difference in the coloration in
the northern race, which in the adult boar shows white
only on the head and the dorsal mane, the sides of the
body, the under-parts, and the legs being quite black.
The female and immature animals are quite rufous-red
on the sides, and strikingly different in color from the old
boars.

Only a single specimen of this pig has been available
for examination. ‘This one is an old boar collected by the
Rainey expedition on Lololokui, a table-topped mountain
situated north of the Northern Guaso Nyiro River. The
skull of this specimen shows the short character of the bony
process above the canine well, and is in general shape quite
identical to the figures of Heuglin’s specimen. The flesh

276 AFRICAN GAME ANIMALS

measurements of this specimen were: head and body, 47
inches; tail, 16 inches; hind foot, 914 inches; ear, 7 inches.
Greatest skull length, 1278 inches. These measurements,
compared to those of an adult boar of the East African race,
show less body size, greater length of tail and ears, and
shorter feet. The specimen here described was one of a herd
of fifteen met with on the broad summit of Mount Lololokui,
at an altitude of six thousand feet. ‘This herd came nightly
to the springs to drink and were occasionally seen in the
daytime, the mountain being quite without human inhabi-
tants. The stomach of this specimen contained the remains
of the white, bulb-like roots of the Sansevierta plants which
grew abundantly in patches on the slopes of the mountain
and resembled closely in habit the smaller yuccas or Spanish
daggers of Arizona or California.

Forest Hoc

Hylocherus

Hylocherus Thomas, 1904, Proc. Zool. Soc., p. 193; type H. minertzhagent.

As recently as 1904 a remarkably distinct giant hog
was discovered in the forests of Mount Kenia and the
Nandi Escarpment by Lieutenant Minertzhagen, of the East
African Rifles. The specimens collected by this officer
consisted of two male skulls from Nandi and the imperfect
skin of a female from Mount Kenia. This material showed
the giant forest hog to differ widely in skull structure or
shape from other known pigs and to require generic sepa-
ration from them. Oldfield Thomas, the original describer
of this new genus and species of pig, has pointed out its
close structural resemblance to the wart-hog as well as the
peculiar elevated crest of the parietal part of the skull, the
concavity on the crown of the head and some peculiarities
in the teeth not possessed by any other genus of pigs.
The forest pig is more closely related to the wart-hog than to
any other genus and is not to be considered intermediate
between the latter and the bush pig, or Sus, as has been
suggested by some naturalists. It is in reality quite a
specialized animal, as shown by the peculiar shape of the

PIGS 277

parietal portion of the skull, but its relationships are best
expressed if considered a forest wart-hog. The characters
which ally it to the wart-hog are the similarity in shape of
the enlarged upper canines, which turn upward and inward
and have only their lower basal parts in contact with the
sharp cutting edges of the lower canines, the presence of
only two upper incisor teeth, and well-developed pits or
choane in the basisphenoid bone. The external similarity
consists of an enlarged wart below the eye, a patch of hori-
zontal whiskers on the cheek, and the presence of but two
pairs of mamme in the female. The cheek teeth have the
ridged structure and shape found in the bush pig, but the
general shape of the skull, especially the enlarged zygomatic
arches, are distinctly like those of the wart-hog. In general
external appearance the forest hog suggests a large edition
of the bush pig, having the same relative proportion of head
to bulk of body, short, stout legs, long tufted tail, pointed
pencillate ears, and a uniform covering of coarse hair
throughout. In bulk of body it exceeds somewhat any
other pig, but in standing height at the withers it is slightly
inferior to the larger Asiatic boars, such as the long-headed
Bornean species.

Although the forest hog eluded discovery until 1904,
the veil which had shrouded it was lifted in a remarkable
manner throughout Africa during the succeeding few years,
within which time three additional new species, or rather
races, were described from the Cameroons and the head-
waters of the Congo River. Our knowledge of the forest
hog may be said to have sprung into existence almost si-
multaneously from remote parts of Africa. ‘Two years after
the discovery of the first specimen we are confronted with
descriptions of two additional species or races of this rare
animal from localities many hundreds of miles from the
original source. Three years later still another was de-
scribed from the Congo basin. Specimens at the present
time are not at all rare in the principal natural history
museums of Europe and America. It is, indeed, remarkable
to find an animal so exceedingly rare and of such recent
discovery so completely illuminated within so few years
of its discovery, while many long-known species, such as the
Derby eland, West African bongo, Abbott duiker, and a

278 AFRICAN GAME ANIMALS

host of other long-known species, remain quite unrepre-
sented in museum collections.

Hylocherus is now known to be distributed from the for-
ested highlands of British East Africa westward through-
out the Congo watershed, to within close proximity of the
Atlantic coast of Africa. The forest hog has a great alti-
tudinal range, being distributed throughout the forest area
quite to its upper limits. In addition to the type species,
minerizhagent, three others have been described, all within
the Congo basin, but none of these really show specific dis-
tinctions, but are based on individual differences in the
skull structure. They represent a single Congo race, with-
out doubt, which is separable from the East African race
only by slight relative differences in dentition, as pointed
out by Thomas for the Cameroons race, rimator.

East Arrican Forest Hoc

Hylocherus minertzhagent

Native Names: Nandi, twmtu; Tiriki (Kakumega forest), mbirri; Masai,
elguya; Kikuyu, numera.
Hylocherus minertzhageni Thomas, 1904, Proc. Zool. Soc., p. 196.

Rance.—Forested regions of Mount Kenia, Aberdare
Range, Kikuyu and Mau Escarpments westward as far as
Mount Elgon and eastern Uganda.

The type species of the recently discovered forest hog
has been named by Thomas minertzhageni, for its dis-
coverer. Minertzhagen, however, did not actually meet
the species which bears his name in a wild state, but ob-
tained his specimens, as he has recorded, from natives and
from some American missionaries resident on the borders
of the Kakumega forest. He refers to one of these mis-
sionaries as having actually shot a forest pig, and this is
apparently the first recorded instance of such a feat by a
white man. ‘The man referred to in these remarks is with-
out doubt Doctor E. Blackburn, the medical officer of the
Friends Africa Industrial Mission, who has long been a resi-
dent of the mission station of Kaimosi, and through whom
were obtained the original skulls of the forest hog sent by
Minertzhagen to the British Museum. Very few white

PIGS 279

men have thus far succeeded in bagging the forest hog, most
of the specimens recorded having been obtained from the
Wandorobo bushmen, who catch them occasionally in their
game pits. Recently the American Museum of Natural
History of New York has received a fine adult pair from an
American hunter, Alfred J. Klein, who shot the specimens
himself near Escarpment Station, in the dense forest cloth-
ing the summit of the Kikuyu Escarpment. The National
Museum collection contains two complete specimens of
adult boars, one a gift to Colonel Roosevelt from Lord Dela-
mere, and the other obtained by purchase from a farmer
resident at Lake Naivasha.

The coloration of these specimens is a uniform black
with the exception of some inconspicuous patches of pale-
buffy or whitish hair on the sides of the snout, near the angle
of the mouth, on the cheeks below the wart, on the inner
side of the ears, and on the belly. The hair covering is
quite uniform, without any tendency toward a dorsal mane,
but rather scanty, the skin showing through the coat.
Upon the cheeks below the wart are a row of horizontally
directed whiskers, as in the wart-hog, but they are much less
conspicuous, not being differentiated by lighter color, but
having a few whitish hairs mixed with the black ones. A
mounted specimen of the Cameroons race, vimator, in the
National Museum is quite identical in coloration with the
specimens from British East Africa, the whitish areas on
the sides of the head agreeing in area and inconspicuousness.
No sexual differences are apparent among forest hogs, but
the wart below the eye is smaller in the females. The color
of the newly born young is not known, but it is doubtless
a uniform deep-brown, as in the young of the closely allied
wart-hog. No flesh measurements of any specimens are re-
corded in the literature, the known specimens consisting
chiefly of skulls or skins obtained from natives. The skin of
the specimen received from Lord Delamere, which is the larger
of the two in the National Museum, measures in a tanned
condition: head and body, 59 inches; tail, 13 inches; hind
foot, Io inches; ear, 414 inches. The skull of this specimen,
which is fully adult and shows wear on the last molar tooth
has a length of 1656 inches, which is slightly less than the
largest wart-hog skull from East Africa. A specimen from

280 AFRICAN GAME ANIMALS

the upper altitudes of the Kenia forest, recorded by Lonn-
berg, is considerably greater than any known wart-hog skull,
being 1814 inches in length. Adult female skulls are smaller
than those of the males, averaging about 1534 inches in great-
est length. The tusks or upper canine teeth of the forest hog
do not show such great development as takes place in the
wart-hog. The larger of the two boars in the National
Museum has an exposed length of tusk of 534 inches. Ward
records two specimens from Mount Kenia in which the ex-
posed portions of the tusks measure 814 and 714 inches,
respectively. The former of these may be taken as the
record tusk length.

Mount Kenia is the farthest eastern point at which for-
est hogs have been collected, there being yet no evidence
of their occurrence in the extensive forests of Mount Kili-
manjaro. On Kenia they are known to occur from the
lower edges of the forest, at four thousand feet, to its upper
limits, at ten thousand feet, where the vegetable growth is
chiefly giant bamboo. At this extreme upper limit Akeley
has noted extensive areas rooted up by forest hogs, and in
places the uprooted vegetation, consisting of herbs and grass,
massed into small mounds like haycocks. In places where
the forest hog is abundant definite paths are made through
the undershrubs of the forest, and where the growth is
dense veritable tunnels are formed. They are much more
strictly confined to the forest than the bush pigs, and never
leave it to make forages into native fields, or shambas.
Like the wart-hog, they are much less gregarious in habits
than the bush pig, often living a solitary life. Specimens
have been secured by sportsmen and naturalists on the
northeast, southeast, and southwest slopes of Mount Kenia;
on the Aberdare Range in the vicinity of Nyeri Hill; on the
summit of the Kikuyu Escarpment; in the Mau forest near
Enjoro and on its summit at Molo; in the Nandi reserve,
at the base of the Mau Escarpment in the Kakumega forest,
near the shores of the Victoria Nyanza; and on the slopes
of Mount Elgon.

PIGS 281

Wart-Hocs

Phacocherus

Phacocherus G. Cuvier, 1817, Reg. Anim., I, p. 236; type P. ethiopicus of
the Cape region.

The wart-hogs are highly specialized members of the
pig family and differ strikingly from such typical members
as the wild boar and the bush pig. The two pairs of large
warts on the face and the enlarged upwardly curved canine
teeth at once distinguish them from all other genera. The
warts are dermal structures and receive no bony or muscular
support, but are made up of a solid accumulation of gristle
or cartilaginous material. Upon the cheek there occurs a
long, ridge-like fold extending toward the angle of the
mouth upon which grow the conspicuous horizontally di-
rected white whiskers. ‘This structure is sometimes con-
sidered a wart and has given rise to the statement that
three pairs of warts are present. The upper canines turn
upward in a great semicircle and only the lower side of their
bases comes in contact with the lower canines to form a
euttme surface. The latter teeth are long and slender,
triangular in shape with sharp edges, and are very formi-
dable weapons, in marked contrast to the blunt upper ca-
nines, which are quite useless for defence on account of their
turned-in points and circular shape. The cheek-teeth also
show decided peculiarity, their surfaces being flat without
transverse ridges and composed of a number of rounded
tubercles of enamel and dentine irregularly arranged, which,
when worn, give the surface of the tooth the appearance of
a cluster of islands of enamel held together by cement.
The last molar attains enormous development in age, when
it occupies three-fourths of the tooth row and in extreme
age becomes the only functional tooth. The head of the
wart-hog is enormously enlarged, the skull of a fully adult
male being quite equal in size to that of a giant forest
pig, an animal having a bulk of body quite twice that of
the wart-hog. The body is round and barrel-shaped, sup-
ported on short legs and quite nude in general appearance.
The hair covering consists of a long, thin mane of coarse hair
from the nape to the middle of the back, where it is broken

282 AFRICAN GAME ANIMALS

by a bare interspace but continued again on the rump.
The hair covering of the sides, under-parts, and legs is very
short and sparse, the individual hairs being separated by
interspaces of half an inch or more. ‘The tail is slender and
quite naked with a tuft of black hair at the extreme tip.
The female lacks the warts of the male with the exception
of a small rudiment below the eye and the ridge of whiskers
on the cheek, has much smaller upper incisors and smaller
body size, but shows no differences in coloration or hair
covering. The mammez are only four. The young differ
from those of other wild hogs by being uniform in color
without lighter stripes or spots. The wart-hogs are con-
fined at the present time in their distribution to Africa, to
which region they appear always to have been limited; for
no trace of them has been discovered in the fossil-bearing
beds of Europe and Asia. Two Pleistocene species are
known from the extreme points of the African continent,
one from the Cape region and the other from Algeria.
Three distinct species may be distinguished by differences
in dentition and shape of skull. Two of these species lack
both upper and lower incisor teeth and occur widely sep-
arated from one another; one, @thiopicus, at the Cape of
Good Hope, and the other, delamerei, in the desert region
of East Africa and Somaliland. The other species, to which
the name africanus has been given, occupies the intermedi-
ate territory from Natal northward to Abyssinia and west-
ward to Senegal. This species has two well-developed
upper incisor teeth and either four or six lower ones. Sev-
eral geographical races of this species may be recognized
which are based on differences in skull shape. It is this
wide-spread species alone which is familiar to sportsmen,
the Cape, ethiopicus, being now doubtless extinct and the
northern one, delamere1, being quite rare in the region of its
occurrence.

Wart-hogs were common on the bare plains of East
Africa, and they occasionally went among the thin groves
of thorn-trees. We found them in the Lado. They were
never found in thick forest. When persecuted by man they

PIGS 283

became nocturnal; but we often saw them abroad at mid-
day, although their favorite times for roaming and feeding
were in the morning and afternoon. They frequently
rooted up the ground, going down on their knees; but even
more frequently they cropped grass like a zebra or harte-
beest. Their food is purely vegetable. They live in bur-
rows, which they invariably enter backward, so as to pre-
sent to any enemy a front of teeth which would daunt the
most ferocious. No matter how headlong the speed of a
hunted pig it will always turn and slip backward into-its
hole, with jack-in-the-box agility. Once we saw a young
boar thus disappear, and immediately afterward reappear
as if shot out of a catapult; evidently the burrow was
already inhabited by another pig, which had savagely as-
sailed the hind quarters of the intruder. When close pressed
they will charge valiantly; but they are certainly nothing
like as savage as peccaries. We found a sow with very
young pigs in November on the Uasin Gishu; but appar-
ently they breed at all seasons. In galloping after a boar we
noticed that in trying to look back at his pursuer he lifted
his snout, not turning his head to one side. . When in mo-
tion wart-hogs always carry the tail straight up, the tassel
perhaps falling forward. They are easily tamed, and make
amusing pets owing to their continuous grotesque starts
and grunts. They often become exceedingly attached to
their owners. At the McMuillans’ place the cheetahs and
wart-hogs were as safe companions as the gazelles; whereas
nobody could touch the leopards; and elsewhere we found
that the zebras and hartebeests were apt to be dangerous
when they became tame.

284 AFRICAN GAME ANIMALS

Key To THE SPECIES AND Races or Phacocherus

Incisor teeth present in both jaws and functional; warts conical
africanus

Postorbital portion of the skull short and narrow, one-tenth the
length of the skull; interorbital region concave
africanus eliant

Postorbital portion of the skull long and broad, one-eighth the
length of the skull; interorbital region flat
africanus bufo

Incisor teeth completely absent in the upper jaw and present in the
lower only as vestiges below the gums; warts knobbed
at the apex delameret

East Arrican Wart-Hoc

Phacocherus africanus ealiant

Native Names: Swahili, ngili; Kavirondo, mbithi ; Abyssinian, kurkerrow.
Phacocherus eliani Cretzschmar, 1826, Rippell’s “Atlas,” p. 61.

Rance.—From the highlands of Abyssinia southward
through Uganda and British East Africa to German East
Africa.

The East African wart-hog was first described from
specimens collected in Abyssinia in 1826 by the renowned
traveller Riippell. Every traveller and explorer who has
entered equatorial Africa from the days of Speke and Grant
down to the present has met with the wart-hog. The East
African race differs in no way, externally, from the Nile or
the Senegambian races, but may be distinguished by the
greater narrowness and shortness of that portion of the
skull projecting behind the orbits. The length of this pro-
jecting part is usually about one-tenth the whole length of
the skull. The wart-hog of East Africa is uniform blackish
in appearance, due to the dark color of the skin and the
scantiness of the hair covering. Along the crest of the back
from the nape to the middle of the back, extends a thin mane
of long, black hair, which hangs down on the sides in loose
strands. The middle part of the back for a distance of some

PIGS 285

six or eight inches is bare, but the rump is clothed scantily
by black hair similar to that of the neck mane. ‘The sides of
the body, under-parts, and legs are clothed by short, whitish
hair set about one-half inch apart and having scarcely any
perceptible color effect on the dark skin. The front of the
legs from the knee downward support a growth of longer,
denser, blackish hair, but the knees themselves are quite
bare and calloused from kneeling. ‘The tail is quite naked,
with the exception of the tip, which is furnished by a tuft
of stiff, black hair. The head has a conspicuous growth of
white whiskers, running from below the eye to the mouth,
which stand out stiffly at right angles to the cheeks, owing
to their insertion on a narrow, shelf-like wart. The top of
the head, snout, and lips are clothed by a scattered growth
of short, black hair, while the lower part of the cheeks and
throat show a somewhat scantier and shorter growth of
whitish hair. The short, rounded ears are thinly covered
by black hair on the back and by long white hair on the
inner side. The newly born young are a uniform umber-
brown, but soon lose this first pelage. When half grown
they are quite like the adults in color and have the white
cheek whiskers well developed.

The flesh measurements of a large male from the Loita
Plains were: head and body, 56 inches; tail, 17 inches;
hind foot, 11 inches; ear, 54 inches. An average female
measures some 12 inches less in length and 1 inch less in
length of foot than these dimensions. Males range in
greatest length of skull from 1514 to 17 inches and females
from 13 to 14 inches. The tusks average in males, in ex-
posed length measured on the curve, 9 inches. The long-
est tusk in the National Museum in a series of twenty-five
adults is a specimen from the Kedong Valley, near Kijabe,
measuring 1314 inches and exceeding Ward’s record for
East Africa by 1 inch. Female tusks average 6 inches on
the curve. The longest in the National Museum is one
from the Loita Plains having a length of 914 inches. A
series of fifty specimens have been examined in the Na-
tional Museum from the Loita, Athi, and Kapiti Plains,
Kedong Valley, Uasin Gishu and Laikipia Plateaux, and
Taveta, British East Africa. The altitudinal range of the
wart-hog seldom or never exceeds eight thousand feet. It

286 AFRICAN GAME ANIMALS

is most plentiful on the highlands between three thousand
and seven thousand feet, on the edges of the plains country,
or in scattered bush. It is rare in the low desert nyikae of
the coast slope.

Nite Wart-Hoc

Phacocherus africanus bufo

Native Names: Dinka, dyehr; Bongo, bohdoos; Niam-niam, tibba.
Phacocherus africanus bufo Heller, 1913, Smith. Misc. Coll., Vol. 61,
No. 22, p. 2.

RanceE.—Nile Valley from the Albert Nyanza north-
ward to the Bahr-el-Ghazal and Sobat Rivers; limits of
range unknown.

The Nile wart-hog has been recently separated as a new
race on the evidence of a skull collected by the Smith-
sonian African expedition at Rhino Camp in the Lado
Enclave. The type specimen was collected on the shores
of a small pond near Chief Sururu’s village in the vicinity
of Rhino Camp. The wart-hog had been killed by a lion the
night previous to our arrival and the head was the only portion
which remained uneaten. Wart-hogs were rare in the Lado
Enclave, less than a score being seen by the members of
the Smithsonian African expedition during a month’s so-
journ in the upper Nile district. In addition to those ob-
served at Rhino Camp a few were seen in the vicinity of a
small lake near Gondokoro.

The Nile wart-hog differs from eliani of Abyssinia and
East Africa by the greater breadth and length of the post-
orbital or parietal portion of the skull and the flatter inter-
orbital region. Other minor differences from e@liani are
the absence of elevation of the parietal or lambdoidal crests,
which are on the same level as the interorbital region, the
narrowness of the choanz posteriorly, and the smaller size
of the tympanic bullae. The type is unfortunately an im-
mature skull in which the last molar is just erupting and
with all the maxillary sutures still evident. Skulls of this
age from the highlands of British East Africa show a much
narrower, shorter, and more elevated parietal region. In
the flatness of the interorbital region and in the general
shape of the postorbital part of the skull the type resem-

PIGS 287

bles the skull of Phacocherus delamerei closely, but differs
from that species, as do all the africanus group, by the pres-
ence of well-developed upper and lower incisors and roofed-
over choane. ‘The skull has a length of 1434 inches, which
is large for an immature female specimen and would indi-
cate a larger body size for this race than that of the East
African. The tusks are small, owing to the immaturity of
the individual, the exposed portion of the tusks being only
344 inches. The longest Soudan specimen recorded by
Ward is one having tusks 1114 inches for the same meas-
urement.

Desert Wart-Hoc

Phacocherus delameret

Native Name: Somali, dofar.
Phacocherus delamerei Lonnberg, 1908, Proc. Zool. Soc., p. 940.

RancE.—Low desert or nyika region of the Northern
Guaso Nyiro River northward to Somaliland and westward
to Lake Rudolf.

The desert wart-hog has long remained unknown or
undescribed, although specimens have for many years been
shot by sportsmen to whom the region which they inhabit
is well known. In 1908 Lonnberg discovered two skulls
at the British Museum, collected by Lord Delamere some-
where in the desert region between Somaliland and the
Northern Guaso Nyiro of British East Africa, which differed
strikingly by the absence of incisor teeth and the shortness
or greater breadth of the skull from those of the ordinary or
africanus type of wart-hog. In these characters they agreed
with gthiopicus, a species confined to the extreme southern
point of Africa and now doubtless extinct. We have thus
among wart-hogs a peculiar condition of distribution, not
duplicated by other game mammals, consisting of two species,
closely allied, living at opposite extremes of the range of the
genus with the intermediate territory occupied by a dis-
tantly related species showing considerably less specializa-
tion. ‘The desert wart-hog must be considered one of the
most highly specialized wart-hogs, owing to the reduction
or loss of its incisors, great breadth of skull, and the enlarged
and knobbed character of its warts. The absence of the

288 AFRICAN GAME ANIMALS

upper incisors is not a recent affair but one which has been
long established, as shown by the absence of even rudimen-
tary incisors in young a few months of age. The lower
incisor teeth show less complete disappearance and still per-
sist as rudiments sunk beneath the rim of the bone in pits
well below the gums. These teeth are quite as rudimentary
and functionless in the young as in the adults. Other skull
differences are the absence of a bony roof over the posterior
part of the choanz or pits in the floor of the skull.

Two specimens of the desert wart-hog are in the National
Museum, collected by the Rainey expedition in the region
lying between Mount Marsabit and the Northern Guaso
Nyiro. One of these is an adult male but not an old ani-
mal, the last molar not being fully in place and the upper
tusks only half grown. ‘The flesh measurements and skull
of this specimen are considerably less in size than those of
the East African wart-hog, as are also those of an adult
female recorded by Lonnberg, which would indicate much
smaller body size for this species. The flesh measurements
of the male were: head and body, 42 inches; tail, 1714 inches;
hind foot, 9% inches; ear, 434 inches. Greatest length of
skull: male, 1336 inches; female, 1234 inches. Exposed
portion of upper tusk in the male measured on the curve,
534 inches. Ward records specimens from Somaliland hav-
ing a length of exposed tusks ranging from Io to 13 inches,
but such specimens may be referable to africanus, which
occurs in the highlands of Somaliland and Abyssinia in close
proximity to delamerei. The desert wart-hog is found only
in the lower drainage area of the Northern Guaso Nyiro
below an altitude of three thousand feet, the river drainage
above this altitude and the slopes of Mount Kenia and the
Aberdare Range being occupied by the East African wart-
hog. It extends southward, no doubt, as far as the north
bank of the Tana River. Sportsmen’s records regarding
this species, however, are wanting, owing to its confusion
by them with the East African race, which it resembles
closely in general external appearance.

CHAPTER X
THE HIPPOPOTAMUS

HIPPOPOTAMUSES
FamiLy Hippopotamide

Tue members of the family Hippopotamide are per-
haps best described as hairless, thick-skinned mammals of
large size related to the pigs. They are, however, only
distantly related to pigs, from which they differ in many
fundamental ways, chief of which are the primitive con-
dition or evenly four-toed character of the feet, the lateral
toes of which reach the ground and assist in supporting the
body weight. The head differs decidedly from that of the
pigs by the absence of a terminal disk to the snout and by
the enormous enlargement of the lower jaw. The canines
are developed into large tusks, the tips of which come in
contact inside of the lips. The incisor teeth grow into
long spikes, which do not meet or function as grass-
cutters, the great square lips with their horny edges
performing the work of grazing. The molar teeth are
transversely ridged and small, the last one showing no
tendency toward the lengthening and complication found
in the pigs. The body is long and barrel-shaped, and sup-
ported on very short legs placed far apart, making loco-
motion slow. The head is very peculiar, being quite square
in front, flat on top, with an enormous mouth and lower

jaw of immense proportions. Placed rather high and near
289

290 AFRICAN GAME ANIMALS

the corners of the muzzle are the elongated nostrils. The
eyes are small, pig-like, and mounted on the upper surface
of the head. Behind them, a short way, project the diminu-
tive rounded ears. The only parts of the body which show
a growth of hair are the tip of the short tail, the inside of
the ears, and the muzzle from the nostrils to the lips.
There are at the present time two living genera, both con-
fined to Africa—the gigantic river hippopotamus, H7ppo-
potamus, and the pygmy or forest hippopotamus, Cheropsts.
The latter genus differs from its gigantic relative, in addi-
tion to its smaller size, by its much longer and more slender
legs, more laterally placed orbits, more circular nostrils,
longer tail, a lesser number of incisor teeth, and much larger
brain and a correspondingly larger brain-case in the skull.
The pygmy hippopotamus spends most of its time on land
in the thick forest growth, to which it retreats when alarmed.
The, lateral position of the eyes and the circular nostrils are
differences in structure which do not fit it for an aquatic
life such as is passed by the larger genus. Hippopotamuses
of both genera were abundant in the Pliocene and Pleisto-
cene age in Europe, Asia, and North Africa. Fossil species
allied to the pygmy hippopotamus are known from the
larger islands of the Mediterranean and from Madagascar,
in which latter island they were the only ungulate and
practically the only land animal showing African affinities,
with the doubtful exception of the bush pig, whose status
as a native mammal is questionable.

Key TO THE Races oF amphibius

Nasal bones narrow, their least width less than 1% inches; upper rim
of orbit less elevated, reaching at most 1% inches

THE HIPPOPOTAMUS 291

above level of crown of skull; rostral constriction
greater, five and one-half times or more in skull length;
body size greater amphibius

Nasal bones wide, their least width greater than 1% inches; upper
rim of orbit elevated 136 inches or more above level
of crown of skull; rostral constriction less, averaging
five times in skull length; body size less kiboko

Hippopotamus
Hippopotamus

Hippopotamus Linneus, 1758, Systema Nature, I, p. 74; type H. amphibius.

The hippopotamus is one of the very oldest known mam-
mals in literature, and is found referred to in some of the
most ancient Egyptian writings. It was well known to the
ancient civilizations of the Mediterranean basin, and several
myths regarding its peculiar habit of sweating blood are
found in Greek literature. Notwithstanding the ancient
and well-known character of the hippopotamus, it remains
without an English name and is everywhere known in our
literature by the scientific generic term proposed by Lin-
nzus in 1758. To the Dutch colonists of South Africa it
was known as the zee koe or lake cow, a term later corrupted
to sea cow by the English colonists. The Germans have
bestowed upon it the name of river horse, owing to the
fancied resemblance of the crown of the head, as seen in a
floating animal, to that of a horse. A far more appropriate
term, considering its zoological affinities, is that of river
pig, applied to it by the ancient Egyptians.

The aquatic habits of the hippopotamus have favored its
development to an enormous bulk, just as in the case of
some of the seals. An adult bull attains a weight of ap-
proximately four tons and is quite equal to a cow elephant
in bulk. Record bulls no doubt reach a weight of five tons
and are only exceeded in bulk among land mammals by the
bull elephant. From the pygmy hippopotamus the true
hippopotamus differs by its great superiority in size, the
shorter legs, shorter tail, which is a mere rudiment and is
much compressed laterally, the prominent dorsally situ-

292 AFRICAN GAME ANIMALS

ated eyes, and small brain. The sexual differences in hip-
popotamuses consist of greater body size, larger canine
teeth, and heavier or more bristle-like hair in the tail tuft
in the bull. The difference in the diameter of the canine
teeth is so great that skulls can be as readily sexed by the
size of the sockets of the tusks as by the teeth themselves.
A large number of fossil species of gigantic hippopotamuses
are known from the Pliocene and more recent Pleistocene
- formations of southern Europe, India, Sumatra, Java, and
the Mediterranean coast of Africa. Some of these species
are scarcely distinguishable by their skull structure from
the living amphibius of Africa. Remains of one of these
closely allied species of amphibius have been found on the
present site of London. Some of the older species of hip-
popotamuses of the Pliocene have been separated as a dis-
tinct genus owing to the presence of the full number of in-
cisor teeth in distinction to the four which characterize the
living genus Hippopotamus. Within modern historic times
the hippopotamus occurred much more widely distributed
than at the present time. Formerly they were found the
entire length of the Nile to its delta, but to-day they are
absent from the Nile proper north of Khartoum and occur
only in its headwaters and tributary streams. At the
present time they are greatly diminished in numbers in
South Africa, being quite extinct from the Cape district
south of the Orange River. They exist to-day, south of the
Zambesi, only in a few rivers where they have been afforded
strict protection. In equatorial or central Africa they are
still abundant from Abyssinia and Senegal south to the
Zambesi watershed wherever there are rivers or lakes with
sufficient water or swamp vegetation to give them cover. A
single living species is known, amphibius, to which several
racial or subspecific names have been applied, the earlier of
these dating back nearly one hundred years. Racial dis-
tinctions have, however, in no case been well established,
owing to the small number of specimens preserved in mu-
seums and the great individual variation shown by the
skulls upon which differences alone have been founded. It
is, however, now apparent that there are not only skull
differences but slight differences in the body coloration
and size as well, which are of a geographical character, and

THE HIPPOPOTAMUS 293

further study will no doubt reveal characters by which
several races may be recognized of this wide-spread species.

The hippo often wanders far overland at night, and
doubtless thus wandered by day in the past ages before
man became the most formidable foe of all big creatures.
In consequence it is found in some land-locked waters, like
Lake Naivasha, to which it can only have penetrated by
extensive journeys across country containing no streams in
which it could swim or hide itself. Where human beings
are rare and relatively inoffensive, it is often found in
streams so small that it seems extraordinary that so bulky
a creature can dwell in them without being always in evi-
dence; but in such streams it always finds some pool or quiet
reach with a fringe of reeds or bush that offers it a chance
to hide during the daytime. Where much molested it be-
comes excessively shy, and during daylight often lies for
hours with only its nostrils above water, in the shelter of
some tuft of water-grass or the like; and under such circum-
stances it is astonishingly hard to make out. But on the
Nile, among the channels and shallows of the vast beds
of papyrus, we frequently saw small herds or family par-
ties of the obese, unwieldly creatures sunning themselves
even at noon, half out of water, or out on the shore, with
their heads resting on the mud or on one another’s bodies.
Biting flies hover over them, and instead of the ticks which
infest the elephant and rhino the hippos are hosts of small
leeches which are often found in swarms fixed to the more
tender parts. Not only have we seen herons and plover
walking about and over them, but also birds that looked
like tick-birds.

294 AFRICAN GAME ANIMALS

We frequently found single hippo dwelling in tiny pools,
usually surrounded by marsh, miles away from the nearest
lake or river, and sometimes so high among the hills that
the animals must have done some real climbing to get to
them. On one occasion while on the Uasin Gishu we were
camped by a reed-covered marsh with very little open water
in it; we had no idea there was a hippo anywhere near; but
when some of the men went to get water, after nightfall,
they were threatened and thoroughly frightened by a hippo,
and when we white men turned out to see what had hap-
pened, we could hear the hippo in the darkness snorting
and gambolling heavily about us, as we walked through the
wet meadow of short grass. At night they were always
very bold, and on their rambles came close to camp; or, if
we were in boats, they would snort and plunge as they
examined us with fearless curiosity close alongside. On
the Nile there were masses of a little surface-floating plant
which we called water-cabbage, Pistia stratoides. Evidently
when the hippos went ashore for a nocturnal walk they car-
ried these plants with them, supposedly on their backs; for
in the morning we would sometimes find them drying in
the hot sun miles inland. In spite of their clumsy build
hippos trot and gallop fast. Their feet are kept far apart by
the wide body and make paths with a ridge down the mid-
dle, so as to be recognizable at once. They swim well, but
go at their greatest speed when they can gallop along the
bottom in shallow water. They can stay under water a
long time, and when they come to the surface they may
send little jets of spray from their nostrils. We were puz-
zled by the noises they made. Occasionally at night we
heard them roar, in a way that we thought must be done

THE HIPPOPOTAMUS 295

with the open mouth; but the ordinary sound was more
like the exhaust of a steam-engine and we think it was
made through the nostrils. At night they made all kinds
of noises while in the water, and when on the bank; but we
never heard them utter a sound when far inland on their
rambles. In the daytime we once heard one uttering a
series of meditative bubbling squeaks from its secure fast-
ness behind the green wall of plumed papyrus.

The cow is devoted to the calf. When young it stands
on her back as she swims. We do not understand the un-
concern with which the hippos and crocodiles get on, for
some of the latter are certainly large enough and greedy
enough to kill a partly grown hippo. Probably the croco-
diles dread the vengeance of the truculent old bulls and
cows. They feast greedily on a hippo carcass; but so they
will on the carcass of one of their own number. The hippo
bulls fight savagely among themselves, and at times a
ruthless old bully will maul other members of the herd.
At Lake Naivasha a young bull which had been thus mal-
treated, and was badly scarred, must have gone slightly
crazy in consequence, for he came on shore and attacked the
cattle, and had to be slain.

Where unmolested hippos become very insolent and not
only ravage the gardens and fields but attack any one who
interferes with them; and in places they attack and upset
canoes, sometimes quite wantonly, sometimes because the
assailant has been wounded or is a cow witha calf. After
the canoe is upset they may wreck it with their huge jaws,
and they may or may not assail the swimmers. In one case,
in the Lado, an old native was almost bitten in two by a
savage bull after his canoe was upset; and we were informed

296 AFRICAN GAME ANIMALS

by entirely trustworthy people that in swimming cattle
across a river savage hippos had been known to assail and
kill them, wholly without provocation. After we left
Africa an English official we had met was upset in a canoe
by a hippo and then carried off by a crocodile. Usually
there is no sport in hippo shooting; it needs nothing but
good marksmanship, and, as the brain is the target, accu-
racy and penetration are the only qualities demanded in
the rifle. Ordinarily, from the circumstances of the case,
there is not the slightest danger in hippo shooting; yet
Colonel Roosevelt was once resolutely charged by a hippo
which he shot in shallow water; with jaws open it came
straight for the boat, which was between it and deep water.
A wounded hippo will sometimes attack the boat of its as-
sailant; and in rare cases an unusually truculent animal will
charge out of the water and try to reach the hunter on land.

Hippos feed on land at night, as a rule, although we
once saw two tearing up and eating water-lilies, or some
plants that were among water-lilies, in the late afternoon.
Naturally they find corn, beans, melons, and other garden
products particularly attractive, and if they are plentiful
will destroy the crops of all villages which lie along the
water-front.

Once on the Nile while two of us were watching a moni-
tor stealing crocodile’s eggs we noticed a hippo in mid-
stream. Although it was in the forenoon, when most
hippos were resting, it appeared above water at about two-
and-a-half-minute intervals, in the same place, breathed,
and sank. This continued for an hour. The current was too
rapid for him to rest; and it hardly seemed that he could
be feeding on anything. We do not know what he was doing.

EAST AFRICAN HIPPOPOTAMUS

In the Tana River near Fort Hall, B. E. A.
From a photograph by Carl E. Akeley

NILE HIPPOPOTAMUS, FEMALE, FOUR YEARS OLD

From Victoria Nyanza
In the National Zoological Park, Washington, D. C.

LIVING SPECIMENS OF HIPPOPOTAMUSES

THE HIPPOPOTAMUS 297

Nite Hippopotamus
Hippopotamus amphibius amphibius

Native Names: Luganda, emvubu; Dinka, nyang; Bongo, habba; Abys-
sinian, gumari.
Hippopotamus amphibius Linnzus, 1858, Systema Nature, p. 74.

Rance.—Nile watershed from the Victoria and Edward
Nyanzas northward to the limits of the grass-land on the
White Nile as far as Kaka, and eastward through the Abys-
sinian highlands drained by the Nile tributaries.

The hippopotamus was first known to Europeans from
the Nile, and it is to this region that we must refer the
original species described by Linneus. If we regard the
species as separable into several geographical races, then
the Nile race must stand as the typical one. Specimens
from the Nile watershed are distinguishable from those
of East Africa by the narrower nasal bones, the less ele-
vated orbits and the greater rostral constriction of the
skull, and the larger body size. At the present time there
are no skins of Nile specimens available for comparison
with East African skins, and the color and other external
characters cannot be determined. Every traveller who has
entered the upper regions of the Nile watershed has met
with this animal and has, with few exceptions, given ac-
counts of his experiences with these aquatic monsters.
The flesh forms a valuable article of food to exploring
caravans or safaris, and the quest of the hippopotamus
has been more or less of a necessity to the leaders of such
parties. Owing to the great bulk of the head, compara-
tively few specimens of skulls have reached Europe, usu-
ally the only parts retained as trophies being the large
lower canine teeth. The Nile hippopotamus, therefore, has
yet to become well known in a detailed way as regards the
variation of its structure. The Nile race of the hippo-
potamus attains a large body size, judging from the dimen-
sions of a male skull in the British Museum, which has a
length of 30 inches and exceeds any other skull known to
the writers. A female skull from the Lado Enclave col-
lected by Major Powell-Cotton shows a correspondingly
large size, measuring 2614 inches in length. In contrast

298 AFRICAN GAME ANIMALS

to these dimensions the largest British East African skulls
are: male, 283 inches, female 2514 inches. Specimens of
the skulls of the Nile hippopotamus have been examined
from the Victoria Nyanza, White Nile, Lado Enclave, Blue
Nile, and Atbara River. On the White Nile hippopotamuses
were observed as far north as Jebelein, 13 degrees north
latitude, by the members of the Roosevelt expedition in
1910. ‘This point is well beyond the sudd area, where the
conditions are quite arid and the desert margins of the
river at this point form a natural barrier to the northward
extension of the hippopotamus. Farther northward and
eastward they occur in the upper waters of all the tribu-
taries of the Nile having their sources in the Abyssinian
highlands.

East ArricaNn HIppopoTAMuUS

Hippopotamus amphibtus kiboko

Native Names: Swahili, kiboko; Kikuyu, nguo; Masai, olmakau.
Hippopotamus amphibius kiboko Heller, 1913, Smith. Misc. Coll., Vol. 61,
INO: 22; pe I

RancE.—From the Rift Valley of East Africa eastward
to the seacoast, north as far as southern Abyssinia, and
southward through German East Africa; limits of range
not known.

A series of five specimens of hippopotamus shot by
Colonel Roosevelt at Lake Naivasha are in the National
Museum. The skulls of these specimens show such con-
stant differences from Nile specimens that they have been
recently recognized as a new race by Heller. This new
East African race is characterized by the wide nasal bones,
great elevation of the orbits, and the lambdoidal crest above
the interorbital region, which has a deeply concave surface
in consequence, and the less constriction of the rostral part
of the skull. From typical amphibius of the Nile it is
further distinguishable by its smaller body size and smaller
canine teeth. It may be distinguished from capensis of
South Africa and the Zambesi River system by its more
circular orbit, the lighter body coloration, and the more
heavily haired ears and tail tuft.

THE HIPPOPOTAMUS 299

The color of the skin is grayish-olive and is deepest on
the back and lightest on the under-parts, where it is in life
a pinkish-drab on the belly, axille, and groins. The hair
covering is absent except on the tip of the tail, the inside of
the ears, and on the muzzle. ‘The tail is furnished by a
conspicuous tuft of seal-brown hair, four or five inches long,
attached like a mane to the ridge of the flattened tip. The
ears are heavily haired on the inside by olive-brown hair
three-fourths of an inch in length, and the great lips at the
muzzle, as far back as the nostrils, support a scattered
growth of stiff buffy-brown tufts of hair placed at regular
intervals apart, like the bristles of a scrub-brush.

The male skin from Naivasha differs from the three
female skins, apart from its larger body size, by the much
stiffer or bristle-like character of the tail tuft, which is also
somewhat more extensive and lighter in color, being buffy-
brown. The three females show much variation in the
amount of tuft on the tail. In the Athi River specimen
the terminal tuft is quite wanting, only a few short hairs
one-half inch long being present. The lack of a definite
tail tuft is no doubt due to the youth of the specimen,
which is quite immature and has its milk teeth still in use.
A living specimen from German East Africa, in the Washing-
ton Zoological Gardens, of about the same age is also with-
out a definite tail tuft. The other two skins from Nai-
vasha, which are fully adult, have the tail tuft composed of
fairly soft chocolate-brown hair, the longest hairs having a
length of 4% inches.

The flesh measurements of the large male from Lake
Naivasha, which has been made the type specimen of the
race, were: head and body, 12 feet 2 inches; girth behind
shoulders, 12 feet; tail, 20 inches; hind foot, 21% inches;
ear, 334 inches. The skull of this specimen measures 2834
inches. The largest adult female from the same locality
measured in the flesh considerably less: head and body, 11
feet 2 inches; girth of body, 11 feet; tail, 15 inches; hind
foot, 21 inches; ear, 3 inches. Length of skull, 2514 inches.
The record circumference for lower canine teeth for British
East Africa recorded by Ward is 7% inches, but this di-
mension is doubtless an inch or two below the real record
for the region. The type bull from Naivasha, which is of

300 AFRICAN GAME ANIMALS

but average proportions, has the lower tusks 7 inches in
diameter. Specimens of the East Africa race have been
examined from Lakes Naivasha, Elmentaita, and Baringo
of the Rift Valley, and from the Athi River of the coast
drainage. The East African hippopotamus is most abundant
in the upper course of the Tana River, from the vicinity
of Fort Hall down as far as Karo-karo. The lower stretches
of the river have been depleted of hippopotamus by the
Wapokomo tribe, who hunt them for their flesh. They are,
however, found in all the rivers having pools deep enough
to afford them shelter, and are also found along the sea-
coast in marshes. They occur everywhere in the desert,
in lakes such as Lake Rudolf, and the Lorian Swamp, where
the shore conditions are often quite arid. In British East
Africa they occur on the high plateaux, in ponds or swamps,
to a height of 8,000 feet at least. Only the highest moun-
tain ranges seem to form barriers to their migration or dis-
persal.

CHAPTER Xx
THE RETICULATED AND COMMON GIRAFFES

GIRAFFES
Girafide

Tue living giraffes are so well separated by the char-
acter of their peculiar body shape and short, skin-covered
horns from the other hoofed mammals that they have raised
the family reputation for distinctness much beyond what
it is really entitled to. The extinct or fossil members are
less grotesque in shape and are less widely separated from
the deer or stags in bodily proportions and other characters.
One of the most remarkable of these was the gigantic Siva-
therium, a genus having large, forked horns, a short neck, and
general moose-like shape, but exceeding that animal greatly
in bulk. A much less specialized animal than the giraffe
is the recently discovered okapi, a forest-haunting, giraffe-
like mammal having a rather short neck and a pair of short,
giraffe-like horns. Notwithstanding the highly specialized
character of the neck and limbs of the modern giraffe, the
family is best characterized by its very primitive horn
structure. The horns are not outgrowths of the bones of
the skull, as in the deer and the antelopes, but they develop
independently and in adult age become fused with both the
frontal and parietal bones upon which they rest. They ex-
hibit the primitive condition of being encased permanently

by a covering of hair-bearing skin, and seem never to be of
301

302 AFRICAN GAME ANIMALS

any functional value as weapons on account of their soft,
skin-covered points, which show no wear. ‘The adult giraffe
horn is homologous to that of a deer horn in the “velvet,” as
it is composed of a bony core covered by soft, hair-bearing
skin. ‘The horns in the female are either lacking or much
reduced in many of the genera. Some of the extinct genera,
however, appear to have had harderand more sharply pointed
horns of more formidable appearance than those of giraffes
and okapis. The limbs show no trace of lateral toes or false
hoofs, nor do they show hoof glands like most of the other
ruminant families. The skull, beyond its horn structure,
shows no other marked peculiarity, nor do the teeth show
any substantial difference from those of deer with the ex-
ception of the broad, two-lobed character of the lower outer
incisor or canine tooth. The family formerly had a much
wider range than at present, and ranged throughout Asia and
Europe from the Miocene on down to the Pleistocene age,
reaching its maximum development in numbers in the
Pliocene of Asia. To-day it occurs only in Africa, where
it is represented by two genera—the well-known giraffe of
the drier and more arid parts of the continent and the
seclusive okapi of the Congo forests.

GIRAFFE
Giraffa
Giraffa Brisson, 1762, Reg. Anim., p. 60; type G. camelopardalis.

The word giraffe at once recalls to mind the picture of a
large animal having an immense length of neck, tall, straight
legs, a short body, small head, and blotched coloration. This
grotesque animal stands as the extreme of his family in neck
development and height of body, and, perversely enough,

MASAI GIRAFFE, ADULT MALE
Shot by Theodore Roosevelt, Kilima Kui

UGANDA GIRAFFE, ADULT MALE
Shot by Kermit Roosevelt, Uasin Gishu Plateau

RETICULATED GIRAFFE

Northern Guaso Nviro
From a photograph by Theodore Roosevelt

THE RACES OF THE GIRAFFE

GIRAFFES 303

has been the only member capable of holding his place on the
earth. His very grotesqueness and colossal size, no doubt,
have been his salvation and have kept him aloof from close
competition with the more modern hoofed mammals with
which he shares the African continent. During Pliocene
times he occurred farther afield, as shown by fossil remains
from southern Europe and Asia, which region is generally
believed to be his birthplace. To-day a single species is
found in Africa, having a distribution throughout the whole
continent south of the Sahara with the exception of the
Congo forest area and the forested parts of the West Coast
generally. ‘The giraffe is the tallest of all mammals, large
bulls not infrequently attaining a standing height of seven-
teen feet to the top of the horns. The usual height for bulls,
however, ranges from fifteen to sixteen feet. The me-
nagerie specimens, with which alone the public are familiar,
are quite undersized and seldom exceed thirteen feet in
height. Bulls are larger somewhat than the cows and are
always decidedly heavier in build. The skull of the bull
is much more massive and is exceedingly heavy, owing to
the large size and dense nature of the horn growths at
maturity. The upper lip projects far in advance of the
lower and is somewhat prehensile, an adaptation commonly
found in many browsing mammals. The sexes are alike in
coloration, the old bulls not being consistently darker than
the cows, as is so often stated. The nursing young are like
their parents in pattern but are lighter colored, although half-
grown, immature animals may be darker than the adults.
The female has a single pair of mamme and produces but
one offspring at a birth. Several well-marked geographi-
cal races are recognizable by differences in coloration and
horn development. There is little difference in body size
or in general style of coloration, all of the giraffes being
marked by large blotches of brown or red on a lighter
ground-color which take the form of irregular reticula-
tions. As we proceed from north to south the frontal
horn decreases in size until finally, in the extreme southern
race in South Africa, it disappears entirely, leaving the
two lateral horns only. Some eight races may be recog-
nized, although considerably more than this number have
been described. At least three of these forms inhabit east

304 AFRICAN GAME ANIMALS

equatorial Africa, the diversified conditions of which coun-
try seem to be especially favorable for the development of
differences in their coloration. A fourth race, congoensts,
typically from the headwaters of the Welle River near the
Nile watershed, may also be found within our area, but at
the present time no specimens from the west side of the Nile
are available for study in natural-history museums.

Key To Races oF camelopardalis

Body marked by large blotches having even margins and separated by
narrow reticulations of lighter color
Ground-color of neck white and forming very narrow reticulations
between the large rufous blotches; sides of head and
throat to chin also reticulated; legs reticulated and
spotted below knee and hock; skull with frontal horn
and mandible at coronoid process low; body size
smaller reticulata

Ground-color of neck buff or ochraceous, not white; blotches tawny
or seal-brown, but seldom rufous; sides of head and
upper throat blotched; legs below knee and hock un-
spotted; frontal horn of skull very high; body size
larger rothschildi

Body marked by small blotches having irregular or jagged margins;
legs spotted below knees and hocks with small dark
spots on an ochraceous or tawny ground-color; frontal
horn of skull smaller tippelskircht

RETICULATED GIRAFFE

Giraffa camelopardalis reticulata

Native Names: Somali, gizi; Abyssinian, gerata.
Giraffa camelopardalis reticulata De Winton, 1899, Ann. & Mag. Nat. Hist.,
EV,.p. 252:

Rance.—Desert region from the northern slopes of
Mount Kenia and the north bank of the Tana River north-
ward to southern Abyssinia and west as far as the east
shore of Lake Rudolf.

The reticulated giraffe was described by De Winton

GIRAFFES 305

from a specimen collected by A. H. Neumann, the well-
known elephant hunter, on the slopes of the Lorogi Moun-
tains in the Northern Guaso Nyiro district of British East
Africa. The name Somali giraffe is often applied to this
race, but it is quite misleading, as the giraffe is unknown in
Somaliland proper, reaching northward no farther than the
headwaters of the Webi Shebeyli River in southeastern
Abyssinia, where it has been met with by Donaldson Smith,
Major Wood, and a few other sportsmen.

We found this very handsome giraffe only along the
Northern Guaso Nyiro. In habits we saw no difference
between it and the common giraffe; its range touched on
the northern range of two of the varieties of the latter,
but there was no apparent intergrading between them
here, whereas the two species of oryx of the same region,
the beisa of the Northern Guaso Nyiro, and the callotis of
south of the Athi, did show a tendency to intergrade.

All giraffe are conspicuous animals which make no
effort to hide or escape observation, and which are always
on the lookout for their foes. At a distance such that the
spots on the coat cannot be seen, the general tint of the
common giraffe’s coloration becomes neutral, so that in
many lights it does not stand out from the landscape, and
then it is only its size and shape that render it conspicuous.
But the handsome chestnut red of the reticulated giraffe
makes it conspicuous by its color, not merely near by, but
at a distance. We do not believe, however, that this dif-
ference in the revealing or advertising quality of the colora-
tion of the two giraffes makes any difference one way or the
other in the life-history of either. Probably no other ani-
mal is less capable of being helped or hurt by advertising
or concealing coloration than the giraffe. Although exclu-

306 AFRICAN GAME ANIMALS

sively a browser, eating only twig tops and blossoms, it lives
in dry, open country, in which the trees do not, as a rule,
grow close together; it rarely or never ventures into dense
forests of the kind in which elephant and other moisture-
loving big game often dwell. Its only beast enemy, when
adult, is the lion; even the lion attacks it infrequently, and
the lion usually hunts its prey by scent; moreover, if a
giraffe were near enough for the coloration pattern of its
coat to be distinguishable, the lion would be certain to see
it anyhow, unless conditions were such as to make all ani-
mals escape observation. In any varied landscape, where
the ground is accidented, where there are trees, and lights,
and shadows, a motionless object of any shape or color may,
under certain conditions, escape observation; and any game
may, under such conditions, escape the eye even of a wild
beast or a native hunter; but even under these conditions
the giraffe is far less likely to escape observation than is any
of the antelopes, or the buffaloes, or even the rhinoceroses.
If it is in such a position that it can be seen at all, it can be
seen far more readily than any other animal, except, per-
haps, the elephant. Its coloration is of no consequence
whatever in enabling it to escape from its foes. When
feeding, when coming to drink, when resting, it never seeks
to evade observation, and trusts entirely to its own wari-
ness, to its keen senses, especially to its sight, and its
speed, for protection.

When we say that the giraffe does not go into dense
forests we are using the words one would use in describing
tropical forests of the low, wet country. ‘The giraffe is often
found in dry thorn forests, but they are sufficiently open to
enable a horseman to follow his game at speed. Along the

MASAI GIRAFFE, CALF
Shot by Theodore Roosevelt, Loita Plains

MASAI GIRAFFE, IMMATURE FEMALE
Shot by Theodore Roosevelt, Loita Plains

MASAI GIRAFFE, ADULT MALE
Shot by Theodore Roosevelt, Kilima Kui

UGANDA GIRAFFE, MALE
Shot by Kermit Roosevelt, Uasin Gishu Plateau

THE» MASAI AND UGANDA RACES OF THE GIRAFFE

GIRAFFES 307

Northern Guaso Nyiro we found the reticulated giraffe
sometimes in bare plains with a few trees scattered over
them, sometimes in thick, dry thorn-scrub, composed either
of low acacia-trees or of tall camel-thorn or wait-a-bit
bushes. The giraffe’s hide is marvellously thick. The
animals would gallop right through patches of thorn scrub
around which we had to steer our horses under penalty of
having our clothes—and skins—torn to ribbons.

Along the Northern Guaso Nyiro most of the big animals
were drinking regularly, either in the river itself or in the
rare tributaries. The country was dry as a bone, the leaves
on the thorn-trees and the harsh grass having withered under
the intense heat of many rainless months. At the good
drinking-places the ground was cut deep by innumerable
footprints of buffalo, eland, oryx, big zebra, common zebra,
waterbuck, impalla, and gazelle. At one time or another
we saw most of these animals drinking; we are convinced
that they were all drinking regularly, probably once a day.
We never saw the gerenuks drinking or saw their tracks at
the watering-places, and we met individuals in places so far
from water that we are confident they were not drinking at
all. We never saw the footprints of the giraffe at any of the
watering-places save one, where we also once saw two of the
animals themselves; and in one place we kept in touch for
three days with a herd which we are reasonably confident
did not drink during that time. Be this as it may, it is
absolutely certain that in portions of Africa giraffes and cer-
tain antelopes—oryx, eland, hartebeest, and others—pass
long periods of time, probably months, without drinking;
and where other antelopes, such as gerenuk, never drink at
all; and this when the heat and dryness are such that it

308 AFRICAN GAME ANIMALS

would seem as if every particle of moisture in the big bodies
would be sucked out. A white man or an average native
will die of thirst within forty-eight hours in the same regions
where these big beasts thrive without any water. Doubtless
there are some bulbs, some plants, from which they get a
little moisture. But the matter is so puzzling that it would
be well worth while for a first-class field naturalist, of the
stamp of Carl Akeley for instance, to study it carefully in
the field. Such studies of the life-histories of the big game
would be more valuable than any series of specimens of all
but the very rarest species.

We needed for the museum two specimens of the reticu-
lated giraffe, a bull and a cow. A cow was killed after a
good gallop, and a bull by fair stalking. The latter was the
only giraffe we ever thus killed, as, with the exception of the
ostrich, they are the wariest African game and the hardest
to stalk; and when we examined the dead body we found
that the explanation was not the skill of the stalker but
the fact that the bull, a very old one, was evidently be-
ginning to lose his eyesight. After these two specimens
were secured, we did not have to molest giraffes again, and
devoted what time we could to studying their habits.

Any man who is much in the wilderness will have occa-
sional queer experiences with animals, which are of interest
but mean nothing; and other experiences which mean a great
deal but the meaning of which it is difficult to unravel. It
is hard either to observe or to understand the psychical and
mental workings of animals and the strange individual and
communal actions which sometimes interrupt the monoto-
nously simple routine of their ordinary lives. Asan instance
of the first kind of experience, while hunting along the North-

GIRAFFES 309

ern Guaso Nyiro, we twice came on giraffes asleep. Both
were cows. One was standing erect among clumps of wait-
a-bit thorn, its eyes closed and head nodding, precisely like a
horse which is sleeping on its feet. Our party, walking in
plain sight, got within thirty yards before it waked; and it
was so startled that it fairly gave a grotesque plunge as it
got under way. The other was also a solitary cow, which
we had stalked thinking it might be a bull. It was strolling
about, and at times thrust its head among the branches of
some tall acacia and stood motionless. When it finally
seemed to come to a permanent halt, Cuninghame said it
was asleep, and Colonel Roosevelt started toward it to see
how close he could get. Cuninghame and the gun-bearers
stayed a hundred yards behind. It was open ground, and
Colonel Roosevelt walked straight toward it. When about
forty yards off it looked drowsily at him. He stopped still,
and in a few moments it again closed its eyes and he walked
forward to within a dozen feet of it. It then waked up but
was not in the least frightened; it pouted its lips peevishly
at him,-and, rearing, hit at him with one foreleg, but struck
short. The other men ran up, and all stood talking and
laughing a few paces from it while it looked sulkily at them.
The men kept under the branches of the tree so that it could
not charge them, as they did not wish to be forced to hurt
it. Finally they pelted it with sticks and clods of earth, and
it slowly withdrew, quite unconcerned. All the giraffes in
this district were relatively tame, but no others let us get
within two hundred yards, and if they saw us they usually
ran long before we got within that distance. We do not
understand the sulky indifference to our presence shown by
this cow.

310 AFRICAN GAME ANIMALS

The other instance may—we only say “may’’—have
signified something more noteworthy, something in the way
of an understanding about a meeting-place, or at least the
desire for companionship. Generally we saw giraffes in
herds or small parties. If thirty or forty were together,
there were usually several old bulls; but a little band of
half a dozen or so might contain a bull or might consist
only of cows and calves—and, by the way, the young giraffes
always struck us as being more alert, lively, and suspicious
than the old ones. We also came on giraffes singly or in
couples; and these solitary ones might be either bulls or cows.
At one of the drinking-places frequented by the game along
the Northern Guaso Nyiro, and at only one, we saw the huge
footprints of the giraffes mingled with the hoof-marks of
bucks and zebras. Late one afternoon, while returning to
camp with the gun-bearers, one of us (Colonel Roosevelt)
thought he would glance at this water-hole. Before reaching
it he found the track of a single giraffe leading toit. Whena
quarter ofa mile away he saw the giraffe standing in the open,
perhaps a hundred yards from the water. The giraffe stood
motionless, looking toward some low hills—not back along his
own track, but in a nearly opposite direction. Colonel Roose-
velt had no thought of harming him, and wished to see him
drink—giraffes straddle their forelegs wide apart, either lat-
erally or fore and aft, in order to drink. The giraffe stood in
the same place and position for fifteen or twenty minutes.
Then suddenly another giraffe appeared round a low hillock,
a long distance off, coming toward the water. It stopped
short, apparently on seeing the first one, when not very far
away. After gazing for some time the newcomer resumed
its march. It halted on coming beside the other, and the

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GIRAFFES 311

two continued to stand near one another, or walk a few
steps in one direction or another. They did not drink or
do anything; and finally, as there was a long distance to go
before reaching camp, Colonel Roosevelt reluctantly left
them. It seemed, judging from the first giraffe’s long stand
in an attitude of expectancy, as if it had come to this spot as
a trysting-place, or at least with the expectation of meeting
one of its fellows—whether lover or friend we cannot say.
Of course, the explanation may have been merely that it
had become aware of the oncoming stranger while the lat-
ter was still a long distance off; but we doubt this, for we
were keeping a sharp lookout, and we believe that the |
second giraffe was far away, beyond ken of sense, when we
began to watch.

Most of the big plant-eaters of the plains, as distin-
guished from the flesh-eaters and also from the plant-eaters
of the forest, seem to crave companionship. Two animals
may form an attachment for one another just as human
beings do. When in the Western ranch country, in the old
days, we always found the saddle band split into little
groups, many of the horses having each his companion or
companions, from which he would not willingly separate.
Probably the same thing is true in the herds of wild game.
It is certainly true of the individuals which are found in
couples. Often a solitary individual of one species will
strike up a friendship with an individual of an utterly dif-
ferent species; or an outcast, or perhaps merely an indi-
vidual of freakish tastes, will be accepted into a herd of a
different species. At one permanent camp we found a
wildebeest bull steadily associating with a herd of the
small Tommy gazelle; and a doe of a Tommy was associat-

312 AFRICAN GAME ANIMALS

ing with, and was accepted as leader by, four of the big
Grant gazelles. We saw a zebra and an oryx which had
evidently become partners, and a hartebeest and a topi
which had done the same. We heard of a case in which
an old bull hartebeest had thus attached himself to a rhino.
Mr. Schillings, a capital observer and writer, was the first
naturalist to insist on this craving for companionship, this
genuine spiritual need of fellowship among these big ani-
mals. In his ‘‘ Flashlight and Rifle” he mentions the case of
a bull giraffe which entered into close alliance with two bull
elephants; and he was able to bring out from Africa a young
rhino, alive, largely by providing it with companions in the
shape of two goats. Doubtless the element of added safety
against foes is largely, probably chiefly, responsible for such
companionship; but we think it also meets a real psychical
need of at least certain animals.

The reticulated giraffe is marked on the neck by distinct
reticulations, formed by the large rufous squares being set
off sharply by narrow lines of white ground-color. This
color pattern is so distinctive from the usual blotched color-
ation of other giraffes that the race has been considered a
distinct species by many naturalists. Some specimens of
the Uganda giraffe, however, show as narrow reticulations,
but the ground-color is seldom so whitish in appearance.
The horns of the bull are well developed, the frontal horn
being especially large and is exceeded in height only by the
Uganda race. The body is marked by large squares of ru-
fous separated by ochraceous reticulations, and differs de-
cidedly from the small size and broken-edged spots of the
Masai giraffe. The legs from the knees and hocks down-
ward nearly as far as the fetlocks are reticulated by buffy-
whitish ground-color and tawny blotches. One of the dis-
tinctive color marks of this race is the carrying forward of
the reticulated pattern of the neck over the cheeks and the
upper throat to the chin. The mandible shows distinctive

GIRAFFES 313

characters, being low at the condyles and having short
coronoid processes. ‘The frontal horn is remarkably robust
and of great circumference, and is scarcely less in height
than in the Uganda race; but the skull itself at this point
is much less in height.

In body size the reticulated giraffe is less than the
Uganda or Masai races. The height of the bull seldom or
never exceeds sixteen feet. The skulls are also proportion-
ately smaller and range from 23 to 24 inches inlength. An
adult pair shot by Colonel Roosevelt on the Northern Guaso
Nyiro had the following flesh measurements: head and body:
male, 12 feet 6 inches, female, 11 feet 11 inches; tail, 2 feet
11 inches; hind foot from hock to hoof, 3 feet 4 inches; ear,
8% inches; height to crown of frontal horn, male, 15 feet
4 inches, female, 14 feet 8 inches. The color of the two spec-
imens is very similar, the male being but little darker than
the female. In body size they are very similar, the male
being greater only in body length and height and more
massive skull, but the length of the latter is slightly exceeded
by the female skull. The material examined has con-
sisted of the original type and some other specimens from
British East Africa at the British Museum, a mounted
specimen at the Carnegie Museum collected by Childs Frick
on the Northern Guaso Nyiro, and five specimens at the
National Museum from the latter locality. All of these speci-
mens agree in the rich rufous blotches and narrow white re-
ticulated ground-color of the neck and are in no way inter-
mediate with the Masai giraffe, which occupies the region just
south of Mount Kenia, from the south bank of the Tana River
southward. ‘The variable Uganda race, however, approaches
the reticulated more closely in color, some specimens from
the Uasin Gishu being scarcely distinguishable. Specimens
from the region immediately south of Lake Rudolf will, no
doubt, be found to be fully intermediate in character. Some
of the skulls from the Northern Guaso Nyiro show well-
developed posterior or fifth horns, a character at one time
supposed to be a peculiarity of the Uganda race. In one
of these skulls the development of the fifth horn is greater
than in any specimens of the latter race.

he reticulated giraffe has been recorded by A: H.-
Neumann as far northward as the mouth of the Omo River

314 AFRICAN GAME ANIMALS

at its entrance into Lake Rudolf, and by Donaldson Smith
from Lake Stephanie and the country east as far as the
Webi Shebeyli River. The southeastern and southern
limits of the race in the Tana district are not yet known.
Lydekker has given the racial name nigricans to a small
piece of body skin from the Kenia district bearing black
blotches separated by narrow, white reticulations, which
apparently came from an individual of this race. It is un-
fortunate that so well known a naturalist as Lydekker
should add to the burden of zoological nomenclature by at-
taching a scientific name to a mere fragment of skin of so
variable a mammal as a giraffe.

Ucanpba GIRAFFE

Giraffa camelopardalis rothschildi

Native Names: Luganda, entuga; Suk, agourt; Bari, kurr1; Sudani, zaraf;
Dinka, mehr.

Giraffa camelopardalis rothschildi Lydekker, 1903, in Hutchinson’s Animal
faites TT pia22:

Rance.—From the Uasin Gishu Plateau and Lake
Baringo northwestward over the highland and desert region
of Uganda to the latitude of Gondokoro; east probably as
far as the west shores of Lake Rudolf and west as far at
least as the east bank of the Nile; limits of range not
known.

The Uganda race has frequently been designated the
five-horned giraffe, owing to the occasional development of
two small horns at the extreme hinder part of the head
behind and above the ears. These so-called fifth horns
are never more than mere rounded knobs an inch in height,
and are only to be found in some aged bulls. The presence
of a fifth pair of horns is not a peculiarity of the Uganda
giraffe, but is shared by the reticulated as well and appears
to be carried to a somewhat greater development in the
latter race. Bony excrescences of equal prominence to the
fifth horns also occur not infrequently above the orbits
and sometimes as adventitious knobs on the sides of the
main horns. In old age the skulls of the bulls produce a
great amount of dense bony tissue which greatly adds to

GIRAFFES 315

the size of the horns and cements the whole upper surface
of the skull together, obliterating all the sutures and pro-
ducing odd knobs of bone in various places as well as over
the orbits and at the back of the skull.

The type specimen of the Uganda race is the adult bull
shot by Major Powell-Cotton on the Uasin Gishu Plateau
and is now a mounted specimen in the British Museum.
The race has been named by Lydekker for the donor of
the specimen to the museum, the Hon. Walter Rothschild,
of Tring. Sir Harry Johnston was the first sportsman to
collect specimens of this race and call the attention of
British naturalists to its apparent distinctive horn char-
acters. Some years later Major Powell-Cotton collected
complete specimens at the expenditure of great personal
labor and considerable cost in the Uasin Gishu Plateau, at
Lake Baringo, and in the Toposo country north of Mount
Elgon and east of the Nile station of Nimule. The specimen
from the latter locality was considered by Lydekker as a
different race, which he named cottonz. Its color differences,
however, are well within the range of Uasin Gishu specimens,
with which it is racially identical. The differences detected
by Lydekker were the absence of black in the neck blotches,
their larger size and more regular outline, which are the
very characteristics shown by the bull shot by Kermit
Roosevelt on the Uasin Gishu Plateau. Specimens like
these, which exhibit a narrow pattern of reticulation on the
neck, are quite intermediate between the blotched colora-
tion of the type specimen and the reticulated race.

The Uganda giraffe is distinguishable from the other
equatorial races by its much more massive skull, the frontal
horn being especially high, and the main horns being as a
rule decidedly thick and massive. The body size is cor-
respondingly somewhat greater and exceeds that of any
other race except the South African, which it equals.
Adult bulls have a height of 17 or 18 feet. The body is
marked by large, regular spots separated by narrow reticu-
lations as in the reticulated, but the neck may be either
reticulated or blotched similarly to the Masai giraffe. The
legs below the hocks and knees are uniform cream-buff,
without darker markings, and are quite similar in this re-
gard to the typical Nubian race. The bulls are not con-

316 AFRICAN GAME ANIMALS

sistently darker than the cows, but vary from seal-brown to
tawny in coloration, independent of age.

The adult bull shot by Kermit Roosevelt had a height
of 17 feet 3 inches to the top of the horns. The other
flesh measurements of this specimen were: length of head
and body, 12 feet 10 inches; tail, 2 feet 10 inches; hind
foot from the hock to the hoof, 4 feet 1 inch; ear, 8%
inches. Skull length, 2534 inches. A skull from Lake
Baringo in the National Museum has a length of 26 inches
and is the maximum in size examined. ‘Two female skulls
from the Uasin Gishu at the British Museum measure 23 and
2334 inches, respectively. » Powell-Cotton records a height of
17 feet 3 inches for his largest bull from the Uasin Gishu.

The Uasin Gishu Plateau represents the most southern
as well as the highest altitude attained by this giraffe.
It is essentially a Nile race which has made its way east-
ward to the headwaters of the ’Nzoia River on the Uasin
Gishu Plateau, and then spread over the watershed and
down into the Rift Valley at Lake Baringo, east of which
it meets the reticulated race. The race occurring on the
west bank of the Nile may be congoensis, the typical speci-
men of which came from the head of the Welle River
within close proximity of the Nile watershed of the Bahr el
Ghazal. Giraffes were seen near Rejaf by Colonel Roose-
velt, at Lado Station by Major Powell-Cotton, and they
have also been reported in the vicinity of Mahaji west of
the Albert Nyanza by sportsmen. What race occurs in
the Lado Enclave district is at present unknown, and it is
desirable that sportsmen preserve complete specimens from
the Lado or upper Bahr-el-Ghazal districts for identifica-
tion.

Masatr GIRAFFE
Giraffa camelopardalis tippelskircht

Native Names: Swahili, zwiga; Kikamba, ndwi; Kikuyu, ndoiga; Masai,
oloadokiragata.
Giraffa tippelskirchi Matschie, 1898, Sitz.-Ber. Ges.Nat. Freunde, Berl., p. 77.

Rance.—British East Africa from the south bank of
the Tana River, the southern slopes of Mount Kenia, Lake
Nakuru, the Loita Plains, and Amala River drainage

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GIRAFFES 317

southward to central German East Africa at least; limits
of range unknown.

A specimen of giraffe from Lake Eyasi in the Rift Valley
of German East Africa collected by Herr von Tippelskirch
was made the type of the Masai race by Matschie in 1898.
Previous to this sportsmen had been accustomed to separat-
ing the giraffe into two races, a northern form having a
high frontal horn and unspotted legs and a southern, which
lacked the frontal horn and had blotched or spotted legs.
The equator roughly marked the dividing line of these two
forms, the reticulated and the Uganda giraffe being classi-
fied as members of the typical or northern race, camelo-
pardalis, and the Masai with the Cape race, capensis.
The Masai giraffe, however, differs widely from the South
African by the well-developed nature of the frontal horn
and the peculiar stellate or dissected margins of the dorsal
blotches. By this latter characteristic it can at once be
distinguished from the other equatorial races. ‘The body
spots differ not only by their jagged outline but also by
their much smaller size, there being fully twice as many in
an area of equal size. The stellate character of the spots
appears to be quite the same from youth to old age. The
blotches of the neck, however, are regular in outline and
are quite indistinguishable from those of the Uganda race.
A specimen from Taveta, Kilimanjaro, in which the legs are
unspotted has been described as schillingst by Matschie,
but it has been shown by Lydekker that spotted-legged
specimens also occur at Kilimanjaro and that the colora-
tion of the legs is subject to much individual variation.
There are in the National Museum six specimens of this
race from British East Africa, five of which have spotted
legs and one, an adult female, unspotted, uniformly light
buff-colored legs, quite like those of the Uganda race. This
individual was shot on the Loita Plains from a herd of six,
together with two others, an immature female and a calf,
both of which have spotted legs. The calf was the off-
spring of the adult cow, and the immature specimen doubt-
less bore the same relation, for they were closely associated
together in a small herd. The range of coloration in the
dorsal blotches of these three is really great. The imma-
ture cow is quite the darkest giraffe examined of any race
and is a uniform seal-brown relieved by narrow broken re-

318 AFRICAN GAME ANIMALS

ticulations of buff. The adult female is marked by bistre-
brown spots, while the calf is a light cinnamon-brown color.
Three adult bulls shot by Colonel and Kermit Roosevelt
near Ulu Station on the eastern edge of the Kapiti Plains are
remarkably uniform in color and have spotted legs which
are tawny in ground-color. ‘The coloration of the series of
six specimens in the National Museum illustrates well the
great individual range and the danger of confusion which
may result from the description of races on single specimens.

The common giraffe, or that variety of it occurring over
the southern portion of British East Africa, and the variety
known as the five-horned giraffe, are identical in their hab-
its with the reticulated giraffe. , They—live im /the)eame
kinds of country, varying from desert to fairly well-watered
plains covered with a rather thin growth of acacia forest.
They browse; they drink when water is plentiful; and they
go without water for weeks if necessary. They are found in
small parties, or herds of twenty or thirty individuals, or
singly. They are usually the most wary of game; and
yet at times show foolish tameness. On one occasion, out
of a small herd of seven, two cows and a young one—the
only wary or sensible animal among them—were shot for
the museum; and the bull, the remaining calf, and the two
remaining cows hung round the neighborhood for some
hours, so little frightened that they permitted Kermit to
get good photos of them. In the land through which we
travelled we found that the giraffes slept standing; but, as
is the case with elephants, some individuals, and in some
localities all the individuals, habitually lie down to sleep.
We did not find the old bulls darker than the cows; some-
times an old bull was the lightest-colored animal in a herd,
and a small cow might be the darkest.

MAP 4—DISTRIBUTION OF THE RACES OF THE GIRAFFE

1 Giraffa camelopardalis reticulata 2 Giraffa camelopardalis rothschildi
3 Giraffa camelopardalis tippelskirchi

319

320 AFRICAN GAME ANIMALS

Giraffes are such strange, picturesque creatures, and so
harmless that they ought to be killed only when absolutely
needed for scientific purposes. To ride them down in a
headlong run is good sport, of course. Their leisurely,
awkward-looking gallop carries them along at a fast gait;
the tail twisting, the neck thrust forward at an angle, the
hind legs coming up outside the fore ones with piston-like
regularity. Buffalo Jones, with his cowboys, Means and
Loveless, roped a giraffe, as they did a rhino and a lioness.
Such feats rank even higher than the feats of the Masai and
Nandi in killing lions with their spears on foot, and the
feats of the Hamran horsemen in killing all big game with
the sword; and from the standpoint of bodily prowess, all
of these feats stand far ahead of the feats of the rifleman.

The flesh measurements of the largest bull from Ulu
were:. head -and ‘body, 13, feet 4 “Inches: tail) ea) ecm
inches; hind foot, 4 feet 5 inches; ear, 914 inches; height,
17 feet 2 inches. ‘The second largest bull had a height of
16 feet 3 inches. The skulls of these two specimens are
quite the same in size and have a length of 25% inches.
The adult female from the Loita Plains measured in the
flesh: 14 feet 7 inches in height; 11 feet 8 inches in length
of head and body; tail, 3 feet; hind foot, 3 feet 9 inches;
and ear, 9 inches in length. Besides the six specimens at
the National Museum others have been examined from the
Athi Plains and Maungu Station at the American Museum
of Natural History of New York, and a mounted specimen
from Kilimanjaro at the British Museum. In the field
herds have been seen near the railway stations of Voi,
Simba, Makindu, Kui, and Ulu, in the Rift Valley, near
Mount Suswa, and on the Loita Plains. They occur prac-
tically everywhere throughout the desert nyika of the coast
and inland through the bush country to the edge of the
grassy plains up to an altitude of 7,000 feet.

CHAPTER XII
ROAN, SABLE, AND ORYX

INTRODUCTION TO FamiLty Bovide

Tue Bovide comprise the oxen, sheep, goats, antelopes,
and their allies. There is, unfortunately, no general term
in English for the group. The popular term of antelope,
which comprises nearly all of the species here considered,
has, furthermore, no zoological significance, but is merely a
term under which have been banded together the genera
not closely related to those which have given rise to the
domesticated races of oxen, sheep, and goats. The various
so-called “ antelopes”’ differ from one another more widely
than some of them do from oxen and than others do
from goats. The roan, the hartebeest, and the eland, for
example, all differ from one another and from the water-
bucks or gazelles as much as from the buffaloes. Some an-
telopes are more divergent from the more generalized forms
than are the oxen. The family may be defined as two-toed,
ruminant, hoofed mammals bearing, in the male sex at least,
frontal appendages in the form of horns supported by bony
cores arising from the frontal bones, and having the false
hoofs rudimentary or without any bony attachment or else
absent. They differ from the deer or Cervide chiefly by
the character of the horns, which are hollow and chitinous
in structure and of dermal derivation, the bony core alone

being homologous to the horns of deer.
321

322 AFRICAN GAME ANIMALS

They range over the whole of Africa and Asia, including
Japan and the larger Malay Islands to southern Europe,
and over North America from northern Mexico to Greenland.
Geologically, the family ranged from the Lower Miocene
to the present time. The most ancient types are repre-
sented to-day by the antelopes of Africa, the bulk of which
have been found as far back as the Miocene in Eurasia. No
remote ancestral types are known. The oldest forms geo-
logically, the Tragocerine, had flattened horns resembling
those of the goat. Other Miocene species of almost equal
antiquity were the Egocerine forms allied to the sable and
the oryx, as well as several extinct genera of the Tragela-
phine or bushbuck type. In expressing the relationships
of the subfamilies of African antelope by diagram, we have
assumed that the sable and its allies represent the most
ancient group, owing chiefly to their more primitive skull
characters. As no mere linear arrangement of groups is
satisfactory, the diagram has been resorted to to express
our ideas of the relationships more clearly, and we wish to
emphasize the fact that the linear arrangement we have
actually adopted is unsatisfactory and is merely adopted
because there had to be some kind of linear arrangement,
and this was, perhaps, a trifle less unsatisfactory than any
other. The arrangement in the diagram is based on a study
of the affinities of the recent species as shown by their skull
structure and horn characters. The Bubaline are an ab-
errant group which do not fit in well with the characters
shown by the others and may represent some of the most
ancient types. The Bovine, or oxen and buffaloes, although
one of the youngest or most modern groups, has characters
which ally it most closely to the very oldest genera. The

EAST AFRICAN ROAN ANTELOPE
In the New York Zoological Park

NILE ROAN ANTELOPE, ADULT FEMALE EAST AFRICAN ROAN ANTELOPE

Gondokoro, Uganda ADULT MALE
Shot by Paul J. Rainey, Amala River

THE RACES OF ROAN ANTELOPE

ROAN, SABLE, AND ORYX 323

most highly specialized seem to be the gazelles, although
they really extend far back geologically to the Miocene age.

Dik-dik
Rhynchotraginae
Klipspringers
Oreotraginae

Gazelles, etc.
Antilopinae

Oribi, etc.
Neotraginae

_Duikers
Cephalophinae

Bushbucks, etc.
Tragelaphinae

Waterbucks, etc.
Kobinae

Buffaloes and Oxen

Hartebeests, etc. Bovinae

Bubalinae

Sable, etc.
Egocerinae

Diagram of the family Bovide showing relationships of the
subfamilies

There are no really remote ancestral types known, the oldest
fossil species being quite as distinctly antelope as the living
species; so that as yet the paleontological records afford
us little clew to the evolution of the group.

324 AFRICAN GAME ANIMALS

Key To SUBFAMILIES OF Bovide

Skull without anteorbital fossa
No anteorbital gland in front of eye
No lachrymal-nasal sinus in skull
Withers much higher than hips; horns rounded at the
base, projecting straight backward, curved back-
ward or spirally twisted, ringed at least at the
base Egocerine

Withers not exceeding the hips in height; horns flattened
or depressed at base, without rings Bovine

A well-developed lachrymal-nasal sinus; horns in male only,
recurved at tips, ringed Kobine

A small anteorbital gland in front of eye
Withers higher than hips; horns diverse in shape, but never
spirally twisted; skull without lachrymal-nasal
sinus; mMammz, two Bubaline

Withers not exceeding the hips in height; horns spirally twisted;
skull with a lachrymal-nasal sinus; mamma, four
Tragelaphine

Skull with a large anteorbital fossa in the lachrymal bone
Anteorbital gland opening on face by a line of pores; nasal bones
usually long, greatly exceeding narial chamber
in length; horns short, projecting backward in
line with the snout Cephalophine

Anteorbital gland opening on face as a rounded pore
Nasal bones long, exceeding the narial chamber in length; ante-
orbital fossa a large pit usually occupying the
whole extent of the lachrymal bone
Hair bovine in texture; snout normal, the length ex-
ceeding the diameter of the orbit; foot resting
on whole under surface of hoof Neotragine

Hair peculiar in texture, pithy, and brittle; snout
short, the length less than the diameter of the
orbit; foot resting on tip of hoof only

Oreotragine

ROAN, SABLE, AND ORYX 325

Nasal bones short, equalling or less than narial chamber in
length
Snout not elongated; tail well developed; horns long,
usually exceeding length of head; body size me-
dium; nasal bones longer, seldom less in length
than narial chamber Antilopine

Snout elongated into a proboscis; tail rudimentary; horns
short, projecting straight back; body size small;
nasal bones very short, less than one-third
length of narial chamber Rhynchotragine

SUBFAMILY LEgocering

SABLE, ROAN, AND ORYX ANTELOPES

Both sexes of the members of this subfamily bear horns,
those carried by the female sex being somewhat more slender
but often no less in length than those of the male. The
horns are either curved evenly backward, as in the sable,
straight, as in the oryx, or spirally twisted, as in the addax.
The antelopes of this group are best characterized by their
large size, high withers, long ears, and absence of lachrymal
glands on the face. Other characters are the long, tufted
tail, maned neck, and striped face. The skull lacks an ante-
orbital fossa, or depression, owing to the absence of the lachry-
mal gland, but in most of the genera a small lachrymal-nasal
sinus 1s present. ‘The teeth are very long and broad, with
well-marked accessory columns in some of the genera, being
similar to oxen in this respect. ‘The skull, however, has the
long nasals and the slender snout similar to the hartebeest.

The young are very differently colored from the adults,
being uniform fawn without any conspicuous markings.

Key TO THE GENERA

Horns curved backward in a great semicircle, at their base arising
abruptly above the orbits Egoceros

Horns straight or but slightly curved, projecting backward in line with
the face and arising behind the orbits
Oryx

326 AFRICAN GAME ANIMALS

SABLE AND RoOAN ANTELOPES
Egoceros

Egoceros Desmarest, 1822, Mammalogie, II, p. 425; type Antilope leucophea
Pallas, the bluebuck.

The well-known name of Hippotragus, which has served
so long for this genus, has been recently replaced by the
older name of Egoceros, owing to its possessing a single-
letter difference over the older genus #goceros, which was
applicable only to the domestic goats and sheep. Changes
of this sort which affect well-known names, and which have
been brought about owing to a change in the rules governing
nomenclature, tend, unfortunately, to destroy rather than
establish stability in nomenclature and are reluctantly ac-
cepted by the authors.

This genus is characterized by the long, evenly curved
horns which arise vertically above the orbits and sweep
backward in a wide semicircle and are heavily ringed for
nearly their whole length. The horns are heavier and longer
in the male. The face is somewhat striped, but the body is
uniform in color on the dorsal surface and the anteorbital
glands are absent. The body form resembles that of the
hartebeests, the withers being high, and the tail tufted and
of medium length. The ears are long and narrow; the snout
is somewhat elongate, and the false hoofs are large. Three
recent species comprise the genus: the sable, the roan, and the
extinct bluebuck. The last-named species was limited to the
extreme southeast corner of the Cape Colony, and it was no
doubt owing to its limited range that it so early became ex-
tinct. The last living specimens disappeared somewhat more
than a century ago, so that their extermination can in no way
be traced to modern firearms. ‘The sable and roan range
from the southern point of Africa northward to the Zambesi
River and thence on the East Coast to British East Africa.
The latter species continues alone through the central lake re-
gion to Abyssinia and the Nile, where it spreads westward to
Senegal along the southern edge of the Sahara. The genus is
lacking only in the Congo forest region. Five fossil species
have been recorded, the oldest from the Miocene of Europe
and four later Pliocene species, one from India, another from

ROAN, SABLE, AND ORYX 327

Europe, and two from Algeria. Horn cores have been dis-
covered in the Pliocene formation of Nebraska which evi-
dently belonged to an antelope allied to this group, but
there is no further American evidence. The genus is one
of the very oldest and most wide-spread of antelopes, and is
no doubt one of the least specialized of the living forms, as
evidenced by the absence of lachrymal glands or speciali-
zation in the facial bones.

KEY TO THE SPECIES OF Egoceros

Horns not greatly exceeding the head in length; ears very long, ex-
ceeding 10 inches; body color a brownish roan
in both sexes equinus

Horns greatly exceeding the head in length; ears shorter, not exceed-
ing 8 inches; male usually dark seal-brown or
black, female ranging from tawny to seal-brown

niger
Roan ANTELOPE
Egoceros equinus

Rance.—The whole of Ethiopia from the Sahara Desert
south as far as the Orange River, in South Africa; absent
only from the Congo forest region.

The roan antelope is distinguishable from the sable by
its conspicuous difference in color and by its shorter horns
and somewhat larger body size. The color in both sexes
is a cinnamon drab or roan varying somewhat in shade in
the several geographical races. Four races are recognizable
by slight differences in color and size: the typical race of
South Africa is the darkest; farther north, in German and
British East Africa, we meet with another, /angheldi; west-
ward the Nile Valley is occupied by bakert, while in the
extreme west, in Senegal, occurs the lightest and smallest
race, gambianus. ‘The skull, compared with that of the sa-
ble, has a much smaller nasal-lachrymal sinus, which is
sometimes lacking entirely; the nasal bones are convex on
the sides, and the occipital region is much longer.

This big antelope, second in size only to the eland, has
a very wide and curiously broken distribution, extending

from the Blue Nile and Lake Chad to far south of the

328 AFRICAN GAME ANIMALS

Zambesi. Its distribution is extensive in altitude no less
than in latitude, and is evidently far less limited either by
climate or by vegetation than is the case with most antelopes.
We found it at home on the cold, wet mountains on the
road to the Uasin Gishu, and in the hot, dry, low-lying plains
along the White Nile, the flora of the two regions differing as
widely as the weather. Yet it is plentiful in very few places,
and these places are generally isolated, and separated from
one another by large tracts from which it is totally absent.
Sometimes it is the only large antelope found in a region; far
more frequently it is found where other species of antelope
swarm. Different kinds of hartebeests, for instance, abound
in many parts of its range, and usually in a locality where
both are found the hartebeests outnumber the roan a hun-
dred to one. Yet the range of the hartebeests is far more
limited. It is difficult to conjecture why the roan, which
has characteristics that enable it to stand such differences of
temperature and of vegetational and geographical condi-
tions, and to range over such a wide extent of territory, —
should, nevertheless, hardly anywhere be able to multiply
so as to hold its own in numbers with other antelopes which
are wholly unable to adapt themselves in like fashion to
change of environment.

Both on the high, wet, cold mountain slopes and plateaux,
and in the low, dry plains, we found the habits of the roan
the same. We found them most often in herds; but we
came across solitary bulls, and we also found them in
couples, usually a bull and a cow, and once we saw three
yearlings together. They were commonly found in the
open plain, but also in thin acacia bush; and on the Uasin
Gishu Mountains we saw them freely enter and leave the

ROAN, SABLE, AND ORYX 329

patches of high forests and bamboo, which were traversed in
many directions by their beaten trails. We found them
lying just within the border of the forest, and we also saw
them lying down in the open. We found them feeding at
all hours, and on the move at all hours. They seemed or-
dinarily to rest for two or three hours, and then to graze for
a longer period. On the Uasin Gishu Mountains they were
the only animals feeding on the rank, coarse, abundant
pasturage. In one place they came to water in the day-
time, sometimes daily, sometimes every second day; in an-
other place we found where one had drunk in the night-time.
They are not very fast, but we found them wary. They
are the fiercest and most savage of antelope, with the pos-
sible exception of the sable. Hunters have been killed and
wounded by them, because of approaching them carelessly
when at bay. A wounded bull charged Kermit with the
utmost ferocity. The roan is almost the only antelope
which, when at bay, will frequently charge when its pursuer
is still some distance off. They are not, of course, to be
classed with the really dangerous game; but they are more
dangerous than a moose or black bear or cougar, for instance.

KeEyY TO THE RACES OF equinus

Front of pasterns tawny-rufous, like rest of leg; ears tawny; horns
shorter langheldi

Front of pasterns buffy-white, in contrast to the tawny leg; ears light-
colored, pinkish-cinnamon; horns longer
baker
East AFricaN Roan
Egoceros equinus langheldi

Native Names: Kinyamwesi, kolongo; Kavirondo, omuga.
Hippotragus langheldi Matschie, 1898, Sitz.-Ber. Ges. Nat. Freunde, Berl.,
p- 182.

330 AFRICAN GAME ANIMALS

RanceE.—German and British East Africa, from Central
German East Africa north through British East Africa to
Mount Kenia, and westward to Mount Elgon; very local
in distribution, the localities being widely isolated.

The East African roan was described by Doctor Matschie
in 1898, from specimens collected near labora, southeast of
the Victoria Nyanza, by Captain Langheld. In the same
year, but some months later, Herr Oscar Neumann described
the race under the name of rufopallidus from specimens
which he had collected in Irangi, German East Africa,
but the latter name is not accepted by naturalists who
follow a strict law of priority in nomenclature. The first
men to meet the roan in East Africa were the explorers
Speke and Grant, who collected a specimen at Kazeh, a
station now known as Tabora.

The East African race is recognizable with difficulty from
bakeri of the Nile Valley, owing to the great individual varia-
tion in color among adults. ‘There is, however, usually a
decided difference in the darker color of the front of the
pasterns, which are uniform in color with the legs; the ears
are also darker, being tawny, but there is no difference in
the length of the hair at the tips, which is usually given as
the chief difference. ‘The dark patch on the chest 1s larger
and darker, as a rule, and the general body color may be
somewhat darker. Two adult specimens, shot from the
same herd in the Amala River district by Paul J. Rainey,
show well how great the individual variation is. One is
dark grayish-drab, the other very light pinkish-cinnamon.
The skull exhibits a considerably shorter snout, the portion
in front of the premolars usually being less in length than
the tooth row, while in bakeri it is usually decidedly greater.

The usual coloration of the dorsal parts is vinaceous-
buff, becoming somewhat darker on the hinder part of the
body, where the black of the tail projects onto the rump as
a dark stripe. The whole dorsal surface is dotted sparingly
with cinnamon hairs. ‘The hair is rather longer on the lower
side of the neck, where it forms a throat mane as in the
waterbuck. The snout, upper and lower lips, chin, stripe
from the base of the horns past the front of the eye, and a
rounded patch behind the eye are white; while rest of face,
including the forehead and cheeks, is seal-brown or black.

ROAN, SABLE, AND QRYX 331

The white patch in front of the eye ends in a considerable tuft
of very conspicuous longer hairs. The very long, pointed
ears are cinnamon on the back and around the base, pen-
cilled with black at the tips, and have the inside white. The
dorsal mane of stiff, erect hairs, the longest attaining a length
of five inches, is cinnamon at the base, tawny above, and ter-
minally black, and extends from behind the ears to the mid-
dle of the back. The chest and the inside of the forelegs are
seal-brown, and the outside is tawny, marked by a wide stripe
of seal-brown. The belly and the inside of the hind legs
are white. ‘The tail reaches the hocks and has a tuft of long,
black hairs at its extremity. Newly born young are uniform
cinnamon without the black leg stripes, but with a narrow
band of black bordering the false hoofs. They have the tip
of the tail blackish; and the top of snout walnut-brown.
Stripe before eye, tip of muzzle, lips, chin, and forethroat
are white. The ears are cinnamon with slightly darker
tips and whitish inner surface. ‘The under-parts are white.

Four specimens have been examined from the crest of the
Mau Escarpment near Ravine Station and two others from
the German border near the Victoria Nyanza. The gen-
eral tone of coloration in these skins is very light, notwith-
standing the high, moist character of the country which
they inhabit. Skins from the Nile match them quite mi-
nutely in general color tone. In British East Africa the roan
are very local, being split up into small herds living in iso-
lated bits of territory. The crest of the Mau Escarpment
west of Ravine Station is the home of one of these herds.
Another is stationed near Muhoroni, on the Uganda Railway,
on the west face of the same escarpment, but much farther
south and lower in altitude; still another herd haunts the
region near the Ithangi Hills southeast of Fort Hall. A
herd is also known to range from Sultan Hamound north-
ward as far as Machakos by way of the Kui Hills. The
greater number, however, are found in the Amala River
drainage, on the German border, not far from the Victoria
Nyanza. In this region many herds occur.

The measurements of the different sexes in the flesh
show them to be of practically the same size. An adult
male specimen measured in the flesh: head and body, 77
inches; tail, 25 inches; hind foot, 24% inches; ear, 111%

332 AFRICAN GAME ANIMALS

inches. The horns of an average male are 25 inches long,
those of a female about 2oinches. The record horns for East
Africa, recorded by Ward, are 293 inches. The largest male
skull in a series of three measured 171% inches in length.

Nite Roan
Egoceros equinus bakert

Native Names: Sudani, abu araf; Dinka, anowm.
Hippotragus bakeri Heuglin, 1863, Nov. Act. Leop. XXX, pl. II, p. 16.

RancE.—Nile watershed from Uganda northward to the
Atbara River and westward throughout the Bahr-el-Ghazal
province, extending westward to the base of the Elgon high-
lands, west coast of the Victoria Nyanza, and northwest shore
of Lake Rudolf.

Sir Samuel Baker met with the roan which now bears
his name on the plains of the Atbara, in Kassala, and south-
ward along the Abyssinian frontier as far as the Latooka or
Acholi country in northern Uganda. He mentions in “Is-
mailia”’ the antelope as being new and compares it to the
roan of South Africa, but the differences which he gave were
merely fancied, the race really being scarcely distinguish-
able from the typical one. Heuglin, the describer of the
race, had only seen the animal on one occasion in the field
owing to the extremely local character of its distribution.
The distinctness of the Baker roan has, indeed, been much
overestimated, chiefly due to the ears having been described
as fringed or tufted, a character which they do not possess.
In the headwaters of the Bahr-el-Ghazal drainage the roan
was recorded as early as 1870 by Doctor Schweinfurth and
again some years later by Doctor Junker. ‘The most east-
ern records are based on specimens obtained by W. N.
McMillan’s expedition to the Boma country northwest of
Lake Rudolf, on the headwaters of the Sobat River.

In general shade of color this race closely resembles
langheldi as well as the typical roan of South Africa. The
whitish or buffy color of the front of the pasterns, the
smaller size and lighter color of the dark chest patch, and
the somewhat lighter ears alone serve to distinguish the race.
The fringed or tufted character assigned to the ears has
been found wanting in all the specimens examined. In

ROAN, SABLE, AND ORYX 333

fact, some of the Nile specimens have the ears less tufted
than average specimens of the other races. The horns
average larger, however, in the Nile race. The record given
by Ward is 3724 inches for bakeri, while that of langheld1 is,
according to the same authority, only 293¢ inches. An
average length, however, for the Nile race is 21 inches.
The longest horned female skull in the collection measures
231% inches. Specimens of the Nile race, shot by Colonel
Roosevelt and Kermit, have been examined from Nimule,
Rejaf, and Gondokoro. The sexes are very similar in size.
An adult male from Nimule measured in the flesh: head and
body along curve of the back, 83 inches; tail, 25 inches; hind
foot, 2414 inches; ear, 12% inches. The length of the skull
of a large male is 1734 inches.

ROOSEVELT SABLE
Egoceros niger rooseveltt

Native Name: Swahili, pala-hala.
Ozanna roosevelt: Heller, 1910, Smith. Misc. Coll., vol. 56, pt. 4, p. I.

RanceE.—From the Shimba Hills near Mombasa south-
me along the coast to the Kigani River opposite Zanzibar

sland.

The sable is the most magnificent and stately of all the
antelopes, its only possible rival being the greater koodoo.
In East Africa until recently it has been known to but very
few sportsmen. It has, however, long been known on the
evidence of native skins to occur on the coast opposite Zan-
zibar Island, but it is only recently that sportsmen have shot
specimens. Sir John Willoughby was the first sportsman
to report the species from British East Africa. He met
with a herd in 1886 west of Maji ya Chumvi, but failed to
secure any. Both Jackson and Percival have also met with
herds near Maji ya Chumvi, but all the specimens which
sportsmen have secured within the past three years have
been shot in the Shimba Hills southwest of Mombasa a
score of miles. The race was described from three speci-
mens shot in the Shimba Hills by Kermit Roosevelt in 1909.
The name irk, given by Gray in 1872 to rufous individuals
of the sable recorded by Sir John Kirk from the Zambesi,
has been by some naturalists confused with the East African

334 AFRICAN GAME ANIMALS

race. Gray’s name, however, was only based on the ob-
servations made by Sir John Kirk and was not founded on
a specimen or a definite locality and does not apply to this
race.

Kermit was the only one of our party who went into the
sable country. He took a small safari thither from Mom-
basa and collected three specimens. The sable were not
plentiful, and were shy. It was near the coast, a well-
watered, hilly country, the rushing streams bordered by
palms and the surface consisting of grassy plains varied by
groves of trees and brushwood. It was hot and wet and
the grass was long. The sable fed in the open plains until
nine or ten in the morning, and then retired to the thick
brush of the groves where they lay down until about five in
the afternoon, when they again came forth to feed. They
were grazers. One of those he killed, a full-grown cow, was
very savage and tried to charge, but was too crippled to be
dangerous. While resting during the day the animals, in
spite of their conspicuous coats, were difficult to find or
approach because they lay in cover so thick as to conceal
them, and it required careful still-hunting to get them. In
the plains they were, of course, very conspicuous, and their
wariness made them hard to stalk. They went in little
bands—they were not sufficiently numerous to gather in big
herds.

The East African race of the sable may be distinguished
from the typical race of the Zambesi basin by its much
smaller size and more rufous coloration. The female is
usually a bright rufous, seldom, if ever, ‘assuming the black
coat characteristic of the male, as she does in the south.
The male is a decidedly smaller animal than the typical
sable, with much smaller horns, the skull in the adult being

WPI “1 Aq parunoyy
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ROAN, SABLE, AND ORYX 335

some two inches shorter, while the horns average more
than twelve inches less in length. The black coat though
assumed is less intense, only the tips of the hair being seal-
brown with the basal part of the hair bright-rufous which
shows through the coat readily when disturbed. In old
males from Mashonaland the hair is uniformly dark seal-
brown to the roots.

The general color of the male is seal-brown. The face
has a median seal-brown stripe from the horn bases to the
end of the snout, bordered on either side by a white stripe
commencing above the eye, passing along a ridge in front
of it where the hairs are somewhat longer, and running down
to the upper lip; another seal-brown stripe runs from below
the eye forward to the nose on either side, but the rest of
the face, upper and lower lips, and chin are white. The
crown of the head is like the back. The ears are white inside,
tawny-rufous posteriorly and tipped with seal-brown. A
dorsal mane runs from the level of the ears to the middle of
the back. ‘The belly, inside of upper part of hind legs, and
rump are pure white, and sharply contrasted with the color of
the back. ‘The tail is seal-brown and ends in a long black
tuft.

The female has the snout medially black from its tip to
the base of the horns, the black separated from the rhinarlum
and horn bases by a narrow band of reddish-brown. A
black stripe encloses the eye and extends from the base of
the horns to the upper lip midway between the angle and
the tip of the snout. ‘The hairs covering the anteorbital area
are creamy-white, set off from the rest of the face by their
whiteness and greater length, and merging rather abruptly
into the buffy stripe which extends forward to the tip of the
snout. The cheeks and throat below the dark ocular stripe ©
are whitish. ‘The sides of the head and neck posterior to the
eye, including the base and the outside of the ear, are light
chestnut, like the general body color. ‘The inside and margin
of the ear conch are white, and the tip has short, dark-brown
tufts. The whole dorsal region, including the head, is tawny-
rufous, darkest dorsally, and slightly lighter on the sides,
where it extends well down and is sharply defined against
the white under-parts. The neck is clothed with a well-
developed mane, which extends from the ears posteriorly to

336 AFRICAN GAME ANIMALS

the middle of the back, the individual hairs black at base and
tip, with the median third tawny like the back. The chest
and lower throat are black, the black extending down the
front of the forelegs as a broad stripe to the hoofs, where it
spreads and embraces the pasterns and false hoofs; posterior
part of forelegs colored like the upper parts. The hind legs
are colored like the upper parts with the exception of the black
bands above the hoofs embracing pasterns and false hoofs.
The tail is black and well haired terminally, but the under
side basally is brownish. The under-parts are white from
between the forelegs posteriorly to the base of the tail and the
lower rump, the white extending down on the inside of the
thigh as far as the knee. The newly born young are quite
different in color from the adults, the dorsal color being uni-
form ochraceous-tawny with a conspicuous seal-brown mane
on the nape extending from the head to the withers. The tip
of the tail, a band about the hoofs, and the back of the pas-
terns are also seal-brown or black. The under-parts and the
back of the thighs are white. The head is marked bya large
white blotch before the eye and a faint indication of the dark
stripe through the eye. The tip of the muzzle is marked by
a small black blotch, but the lips, chin, and forethroat are
white. The ears are colored like the body with an indica-
tion of darker tips and are white inside.

No flesh measurements are available. The skull of the
largest male examined measures 16 inches and is equalled
by that of the female shot by Kermit Roosevelt. The horn
length in an average male is about 32 inches. The longest
horns recorded by Ward from British East Africa are 37
inches as against 5414 inches for the record for the typical
ee The horns of the only female examined measure 24
inches.

Oryx
Oryx

Oryx De Blainville, 1816, Bull. Soc. Philom., p. 75; type O. gazella, the
gemsbok.

The oryx is at once recognizable by its long, slender
horns projecting straight backward in line with the profile
of the face, and by its high withers, striped face, and grayish

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MAP 5—DISTRIBUTION OF THE RACES OF SABLE AND ROAN ANTELOPES

1 Egocerus niger roosevelti 2 Egocerus equinus langheldi 3 Egocerus equinus bakert

337

338 AFRICAN GAME ANIMALS

or whitish coloration. The tail is long and tufted, the false
hoofs are well developed, and the lachrymal glands are ab-
sent. The skull is characterized by its short snout, dis-
tinct lachrymal-nasal sinus, and narrow cheek-teeth, which
show only a rudiment of the internal accessory columns.
Both sexes are horned, the horns being slightly more slender
in the female, notwithstanding their greater length. The
sexes are equal in size. The oryx is strictly a desert or dry-
country antelope. It ranges from the deserts of Arabia and
the Sahara southward to central German East Africa, and
is found again farther south in the Kalahari Desert and in
southern Angola. Four living species are recognized: the
gemsbok of southwest Africa, the beisa of northeast Africa,
the leucoryx of the southern Sahara, and the beatrix oryx
of Arabia.

Two allied fossil genera have been described, Protoryx,
with four species from the Upper Miocene of Asia Minor,
and Palgoryx, with three Upper Miocene species from the
same region, and four more recent Pliocene ones from Italy
and Spain.

Beisa Oryx
Oryx beisa

The races of oryx assembled under beisa agree in having
the head and body markings and the shape of the horns
very much as in the gemsbok of southwest Africa. They,
however, are readily distinguishable by the lack of the
black rump patch, and the throat mane, as well as the much
narrower extent of the knee bands and the obsolete char-
acter of the dark stripes on the cannon-bones. In the re-
duction of the black marking they show some approach to
the more northern Jeucoryx and Arabian oryx which have
lost both the dorsal and lateral stripes as well as the
leg stripes. The beisa and its races inhabit the East
Coast of Africa from the Red Sea port of Suakin south
through Somaliland and Abyssinia to central German East
Africa.

7

ROAN, SABLE, AND ORYX 339

Key To THE Races oF beisa

Ears not pencillate; hair whorl situated on the rump; body purer
gray in color annectens

Ears pencillate; hair whorl situated midway between rump and withers;
body usually with a brownish cast over the drab
callotis

IpEAN BeErIsa Oryx
Oryx beisa annectens

Native Names: Samburru, sala; Rendile, ogorr.
Oryx annectens Hollister, 1910, Smith. Misc. Coll., vol. 56, No. 2, p. 7.

Rance.—From the Tana River, the northern slopes of
Kenia and Lake Baringo northward throughout the desert
and Lake Rudolf region to the Abyssinian border.

Specimens of this race secured on the Laikipia Plateau
at the headwaters of the Northern Guaso Nyiro by John
Jay White furnished Hollister with the material which led
to the description of the race in 1910. The present race of
the beisa had been collected some years previously by the
elephant hunter Arthur Neumann as well as by other
sportsmen, but these early specimens were considered by
naturalists to be identical with the Abyssinian race gal-
lorum, or else to represent true beisa of the Red Sea coast.

Oryx were plentiful along the Northern Guaso Nyiro.
One of them, a bull, tipped the scales at four hundred
pounds. They lived on the dry flats, both where there were
open stretches of grass, and where they were covered with
leafless thorn scrubs. They were found in small parties and
herds of as many as fifty individuals. They also went singly
or in couples; apparently the breeding cow stayed by her-
self until the calf was a few days old, when she rejoined the
herd. We found a cow with a just-born calf, accompanied
by a younger animal; the latter was probably the cow’s
young of the preceding year. In September all the calves

340 AFRICAN GAME ANIMALS

of the oryx along the Northern Guaso Nyiro were new-born.
Old bulls were sometimes found alone, or associating with
zebra herds; sometimes a couple of the old fellows would be
found together, each doubtless turned out of his own herd by
some vigorous master bull; and sometimes three or four
young bulls would be found together, waiting until weight of
body and length of horn might give them a chance each to
master a herd for himself. The master bull keeps the rest
of the herd in awe. Once we saw a bull, which had been
shot at, gallop off several hundred yards with his herd, and
then when the herd stopped, bawl or grunt loudly, and
chase round two or three of his family in succession. Ex-
cept on this occasion, we never heard an oryx make a
noise, save for a kind of sneeze uttered by one animal
when it saw us a hundred and fifty yards or so distant.
The oryx are grazers, but we saw one browsing. Al-
though they can go for long periods without water, they
drink every day or so when it is accessible. We found a
herd watering at a bend in a stream at about noon, and an-
other coming to the river to water rather late in the after-
noon; and at water-holes in the desert oryx came to drink
in the moonlight. In the heat of the day the herds some-
times feed, but are more apt to rest, either standing or
lying; usually we found them resting under thin thorn-trees,
but once or twice in cover that was fairly thick. Usually
the herds we saw in the early morning and late afternoon
were feeding, sometimes alone, sometimes in herds with one
or the other species of zebra, once with a couple of water-
buck, once with some of the big gazelles. When either
resting or feeding, the long tail is continuously switching;
and this, even more than the vivid black and white mark-

ROAN, SABLE, AND ORYX 341

ings, makes the oryx a conspicuous beast, unless so far dis-
tant that the only effect is that of his general coloration.
When at a long distance, so that the movement of the tail
does not catch the eye, and the effect of the black and white
markings is minimized, the oryx is less boldly noticeable
than the bonte-quagga and big zebra. At night, if sideways
to us, they were rather less noticeable than the zebra, if
either could be seen at all. Oryx are sleek, handsome
beasts. We found them usually wary, difficult to approach,
and trusting to their noses, but especially to their eyes, for
safety. Yet, although ordinarily the oryx trusts to its eye-
sight, its eyes sometimes betray it; whereas its nose never
does if it can get the wind of a foe. The sense of smell is
usually the determining sense. Near the mouth of the Isiola
River an oryx which showed extraordinary tameness was ap-
proached to within fifty yards. It remained gazing stupidly
at the observer while he remained at this distance and in the
same position, which was on the lee side of the oryx. Then
he shifted his position to the windward so that his scent was
carried to the antelope. As soon as it received this warning
of the presence of a man it bolted to a safe distance. This
sort of visual stupidity is, however, quite unusual. They
are swift and enduring, but do not rank with the harte-
beest in either respect.

Oryx are bold fighters. The bulls use their horns
chiefly for battle among themselves; but, as the cows have
them also, it is difficult to believe that they have been de-
veloped solely for sex conflict. A wounded bull which we
galloped down turned to bay and threatened a charge. It
uttered a grunt so angry and menacing that it could eas-
ily be called a growl. It defends itself effectively against

342 AFRICAN GAME ANIMALS

hounds, and doubtless would do the like against wild beasts
if given the chance. It is said on occasions to beat off the
lion. We saw nothing to indicate that such a feat, if it ever
occurs, is usual. The lions along the Northern Guaso Nyiro
were killing oryx and zebra indifferently, and there was
nothing in the record left by and around the carcasses to
show that there was more of a struggle in one case than in
the other. The lion’s attack is normally a surprise, a furious
rush at close quarters, and a grapple before the quarry has
a chance to flee or fight; and once he has hold, especially
if one paw is on the head or neck, the chance for resistance
from an antelope no bigger than an oryx is at an end.
Moreover, the antelope’s aim is to escape, not tofight. The
lion’s life—that is, his dinner—depends on his killing the
antelope; whereas the antelope’s life depends, not on his
killing, but on his escaping from the lion; so that the ante-
lope’s whole energies are bent on getting away, and unless
this is impossible he makes no fight. If a cow with a very
young calf discovered a lion in time she might oppose the
beast, and then, being on her guard, her lance-like horns
might make it very dangerous for her assailant.

The Ibean beisa oryx is somewhat intermediate be-
tween the Abyssinian beisa and the fringed-eared oryx. It
may be distinguished from the former by its much grayer
color, which lacks the vinaceous or pinkish wash, and by
the gray of the sides extending below the black lateral line,
the latter not forming the boundary between the white
under-parts and the gray sides. From the fringed-eared
oryx it may be distinguished by its grayer coloration, lack
of pencillate or tasselled ears and posterior situation of the
dorsal hair whorl on the rump. The dark head markings
are variable. Usually the dark preocular stripe terminates
freely without joining the dark face blaze or the throat stripe,

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ROAN, SABLE, AND ORYX 343

but in some specimens it unites at its tip broadly with both
the black face blaze and the mandibular stripe to form a
solid dark ring about the muzzle near the tip, as is character-
istic of the South African gemsbok.

Both sexes are alike in coloration. An adult male is
uniform écru-drab on the dorsal surface, with the median
dorsal region marked by a broad black stripe from the
withers to within six inches of the base of the tail. The
lower sides are marked by a broad lateral stripe extending
from the knee band along the lower flanks to the hind quar-
ters, the stripe being widest posteriorly. The gray of the
sides borders the band below for a few inches where the color
merges gradually into the white under-parts. The forelegs
are white, marked with a broad band above the knee and a
narrow blackish streak on the front of the cannon-bone. ‘The
false hoofs are narrowly bordered by black hair. The hind
limbs are vinaceous-drab, changing to white below the hocks,
and the false hoofs are bordered by black hair. The posterior
part of the limb below the hocks is marked by an ill-defined
blackish stripe. The under-parts are white from the chest
to the base of the tail. The tail at the base is drab like the
back and the tip has a long bush of black hair. The neck
is colored like the body and furnished on the nape with a
short mane. The median line of the throat is marked by a
broad black stripe from the head to the breast, where the
stripe forks and joins the lateral bands. ‘The head is strik-
ingly marked black and white, the crown being black, bor-
dered by a cream-buff band on the forehead from the horn
base across to the interorbital region, but narrowly separated
by a tongue from the black face blaze. The front of the face,
except the snout, is covered by a broad black blaze reaching
well down on the sides, but not extending nearer than half-
way to the tip of the snout. A broad dark streak extends
through the eye from the base of the horn to the angle of the
mouth and is separated from the face blaze by a wide streak”
of white. The snout, chin, and forethroat are wholly white.
The back of the head is drab. The ears are drab, the tips
for an inch being marked by blackish hair, and the inside of
the ear by white. A wide dark stripe extends from the base
of the ears downward and joins the dark median throat stripe.
The nursing young are very different from their parents in

344 AFRICAN GAME ANIMALS

color, being quite without the dark color markings, the dorsal
coloration being uniform cinnamon-buff, only the tip of the
tail and a blotch above the knees black. The extreme tip of
the ears is dark-brown, and the eye region has a faint indi-
cation of the dark diagonal streak. The snout is dark seal-
brown. The belly and the under-parts are buffy like the
sides. There is no indication of the dark dorsal, flank, or
throat stripes or face blaze, and the tip of the snout, which in
the adult is white, is quite dark seal-brown. The head and
the body stripes in the adults show much individual variation.
In a series of ten skins from the Northern Guaso Nyiro only
six have the dark preocular stripe free at its tip from the
face blaze or the gular stripe; in two of the others the
preocular stripes are fused with the face blaze where it ter-
minates on the snout, and in the last two they are confluent
with the dark gular and throat stripe, a condition charac-
teristic only of the gemsbok of South Africa. The dorsal
stripe also shows much variation, being quite obsolete in
one-half of the skins. The flank band is always present but
varies much in width.

The flesh measurements of a large male are: head and
body, 67 inches; tail, 181% inches; hind foot, 20 inches; ear,
8 inches. The females measured show practically the same
dimensions. The females, as a rule, exceed the males in
length of horns, but their horns are usually more slender and
distinctly thinner at the base. In a series of twenty in the
National Museum, however, the longest horns are those of a
male, and are 33 inches in length; the longest female horns
being 32 inches. The basal circumference of the male horns
is 7 inches as against 514 inches for the female. The long-
est male skull, however, is exactly equalled by the longest
female skull, the dimension being 1514 inches. The max-
imum horn length given by Ward for an East African beisa is
38 inches. Both the true beisa and the fringed-eared have
smaller horns than the present race.

The specimens examined by the writers have all come
from the Northern Guaso Nyiro drainage, ranging from
the headwaters on the Laikipia Plateau as far down as
the Lakiundu junction, and thence northward to Mount
Marsabit. Northward annectens no doubt merges into
gallorum at the north end of Lake Rudolf, but southward it

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1 Oryx beisa callotis 2 Oryx beisa annectens

345

346 AFRICAN GAME ANIMALS

does not mix directly with callotis, being separated from the
latter by the Tana River and the highland barrier of Mount
Kenia and the Aberdare Range. The northwestern limits
are McMiuillan’s records from the Boma country northwest of
Lake Rudolf, which he was the first to explore.

FRINGED-EARED Oryx
Oryx beisa callotis

Native Name: Swahili, cheroa.
Oryx callotis Thomas, 1892, Proc. Zool. Soc., p. 195, colored plate XIV.

Rance.—The desert district from the Tana River south
to Kilimanjaro and thence west to the Rift Valley, ranging
from Mount Suswa south to central German East Africa
as far as Ugogo.

The fringed-eared oryx is one of the numerous races dis-
covered by Rowland Ward among the trophies sent to his
establishment by sportsmen. He pointed out to Thomas
the peculiar character of its pencillate ears; and upon fur-
ther investigation it was described, and the specimen which
came from Kilimanjaro presented to the British Museum.
This oryx was well known to the early sportsmen who
visited Kilimanjaro previous to 1892, particularly to Jack-
son, Willoughby, and Abbott, under the name of beisa.

This species resembles in coloration and markings closely
the Ibean beisa oryx, from which it may be distinguished by
its tasselled or pencillate ears and the more brown or fulvous
suffusion of its coat. The face markings are similar to those
of the Northern Guaso Nyiro race, the preocular stripe usu-
ally showing no connection with the face blaze or the throat
stripe. Variations, however, no doubt occur in which this
stripe joins the face blaze or the throat stripe, although, in
the few specimens examined, no such variations were seen.
The body color is also variable, and in some specimens it is
quite as gray as in the more northern beisa; but the race
may always be distinguished by its fringed ears.

No flesh measurements are available, but, judging by the
size of the skull, this race is practically the same size as the
Northern Guaso Nyiro race. The skull of an adult female
measures 1514 inches in greatest length. ‘The tooth row is,

ROAN, SABLE, AND ORYX 347

however, longer than in annectens and is, no doubt, one of
the distinguishing characters of the race. The horns of an
adult female collected by Doctor Abbott are 29% inches
long, which is only a few inches less than Ward’s record of
33% inches for the race.

The fringed-eared is a much more local race than the
northern. It is found locally near the Taita Hills, near
Lake Jipe, at Useri on the east slope of Kilimanjaro, near
Makindu on the Uganda Railway, again far eastward on
the coast at Merereni. In the Rift Valley it is found just
south of Mount Suswa in the latitude of Nairobi, again at
Lakes Magadi and Natron, and thence in isolated localities
in German East Africa at Lake Manyara and in Ugogo,
where it reaches its extreme southern limit.

CHAPTER. sit
WILDEBEEST, DAMALISCUS, AND HARTEBEEST

SUBFAMILY Bubaline

Tue Bubaline include some of the most grotesque and
specialized antelopes as well as some comparatively gen-
eralized types. In affinities they are closely related on the
one side to the egocerine antelopes or sables through the
genus Damaliscus and on the other to the African buffalo
through the wildebeest, Gorgon. The hartebeests are a
highly specialized genus and represent the extreme diverg-
ing branch of the Bubalineg tree. The characters which are
common to this somewhat diverse assemblage are the elon-
gate head, especially the snout region, which has very long
nasal bones; the high withers; the long, tufted tail; the pres-
ence of horns and of two mamme in the female; the rudi-
mentary condition of the anteorbital gland, and the narrow
ears. The members are all grazers and are primarily open-
plains antelopes possessing great speed. The group is ap-
parently of more modern origin than either the egocerine
or tragelaphine antelopes. The oldest members recorded
have been found only as far back as the Pliocene of India.
More recent species of hartebeests have been discovered in
Pleistocene deposits in South Africa.

KEY TO THE GENERA

Skull not markedly elongate, the horn bases never united into a pedicle
Horns buffalo-like in shape, curving downward from the base and
extending outward at right angles to the head; nape
and throat maned; muzzle broad Gorgon
Horns not curving downward from the base but directed backward
and upward; neck without mane; muzzle normal
348

WILDEBEEST AND HARTEBEEST 349

Horns curved backward and upward; tail tuft black; lower
premolar teeth three Damaliscus

Horns curved outward and then parallel, forming a U, and greatly

exceeding the head in length; tail tuft white; lower
premolar teeth two Beatragus

Skull elongate, the horns united at the base to form a pedicle which is pro-
duced backward beyondtherestoftheskull Bubalis

DaMALIScUS
Damaliscus

Damaliscus Sclater and Thomas, 1894, Book of Antelopes; type D. pygar-
gus, the bontebok of South Africa.

The genus Damaliscus is an assemblage of four very dis-
tinct species of hartebeest-like antelopes, comprising the
bontebok, blesbok, sassaby, and topis. There is, unfortu-
nately, no general term in use among sportsmen for these
four species, each of which bears a distinct name, and the
only way of supplying this deficiency appears to be the
adoption of the generic name as a common English term
for the group. The damaliscus have the high withers of
the hartebeest, but the head is less elongated and the horns
are not united in a pedicle at the base but rise independ-
ently above the orbits as in the sable and roan antelope.
The horns are heavily ringed and curve evenly backward
as in the roan with the exception of the tips which are de-
flected slightly upward. The horns are much shorter than
those of the roan and are never curved in a semicircle, their
general direction being upward or backward in line with the
profile of the head. The close relationship of the damaliscus
antelopes to the sables and roans is obvious on the compari-
son of skulls in the structures of which thereis closeagreement
in the shape of the bones, the absenceof sinuses, and the slight
development of the anteorbital fossa. The cheek-teeth of the
damaliscus, however, are much narrower and less folded than
those of the egocerine antelopes. The genus is most highly
developed in South Africa, in which region three distinct spe-
ciesoccur. North of the Zambesi watershed we find only one
species, D. korrigum, occupying a wide range of territory from
the East to the West Coast as far northward as the southern
border of the Sahara Desert and the Abyssinian highlands.
A single fossil species is known from the Pleistocene of India.

350 AFRICAN GAME ANIMALS

EguaToRIAL DAMALIScus

Damaliscus korrigum

Included under the term equatorial damaliscus are the
three races known to sportsmen as the korrigum, tiang, and
topi. They are very closely related and scarcely distin-
guishable notwithstanding their diverse geographical ranges,
which are accompanied by considerable local isolation. For-
merly these three forms were considered as species and dis-
tinct names were applied to them under the assumption
that they were quite distinct. Owing to the scarcity of
specimens in museums for comparison the real differences
in these forms have remained unknown. It has recently
been found that they are subject to a considerable degree
of variation in their black color markings which renders the
characters upon which the old species were based void.
Damaliscus korrigum has the coloration and body size of
the sassaby of South Africa from which it is distinguishable
by decided differences in the shape of the horns. In the |
sassaby the horns spread outward from the head in a wide
curve, somewhat as in the tora hartebeest, and then curve
inward at the tips and are never parallel or gradually diver-
gent backward as in the korrigum. The equatorial damalis-
cus inhabits the territory south of the Sahara Desert and the
Abyssinian highlands from the Tana and Juba Rivers on the
East Coast westward through the Nile Valley and Lake Chad
watersheds to the West Coast and Senegal region. Three
geographical races may be recognized, the topi in the east,
the tiang in the central region, and the korrigum in the west.

Key To THE Races oF korrigum
Coloration lighter, cinnamon; pelage shorter; horns longer, averag-
ing 3 inches more in length tiang

Coloration darker, cinnamon-rufous; pelage longer and heavier; horns
shorter jimela

Tianc Damatiscus
Damaliscus korrigum tiang

Native Names: Djeng, tiang; Bongo, tanghe.
Damalis tiang Heuglin, 1863, Ant. u. Buff. N. Ost. Afr. (N. Act. Leop. XXX,
pt. II, p. 22, pl. I, col. fig. head).

WILDEBEEST AND HARTEBEEST 351

Rance.—From the Albert Nyanza northward through
the lowlands of the Nile Valley to the Blue Nile, eastward
as far as Lake Rudolf and westward to the limits of the
Bahr-el-Ghazal drainage.

Von Heuglin in 1863, in his account of the antelopes and
buffaloes of Northeast Africa, described the tiang damaliscus
as new, giving it the native Djeng name of tiang. He states
that this species is very abundant in the country of the
Djeng, who occupy the territory between the Bahr el Zeraf
and the Sobat River near its junction with the White Nile.
It is a widely distributed animal in the Nile Valley and has
been met with by such early explorers as Schweinfurth,
Baker, Emin Pasha, and a host of recent sportsmen and
travellers.

The tiang differs very slightly from the topi of East
Africa, and is distinguishable from it with difficulty. The
general tone of the coloration is lighter, being cinnamon, and
less reddish than that of the topi, but the black markings
of the Imead and legs are quite identical in extent. The
hair covering on the coat is somewhat thinner and shorter.
The horns are distinctly longer and average some three
inches greater in length. No specimens representing the
tiang are in the collection of the National Museum, but
specimens from the Nile Valley have been examined at
the British Museum. A series of specimens is greatly
needed to show the limits of variation in coloration and
skull characters of this race.

Tort Dama tiscus
Damaliscus korrigum jimela

Native Names: Swahili, top7; Unyamwezi, jimela.
Damalis jimela Matschie, 1892, Sitz.-Ber. Nat. Freunde, Berl., p. 135.

Rance.—From Mount Elgon and the northern high-
lands of Uganda southward over the Mau Escarpment and
Victoria Nyanza drainage to central German East Africa;
westward as far as the Edward Nyanza and Lake Kivu;
also near the coast from the Sakaki and Tana Rivers north-
ward as far as the Juba River.

Sir John Kirk, the British consul-general at Zanzibar,

352 . AFRICAN GAME ANIMALS

to whom we are indebted for the earliest accounts of the
game animals of East Africa, met with the topi as early as
1866 on the plains near Merereni and also at the mouth of the
Tana. Jackson, Harvey, and Hunter a score of years later
recorded the topi in the same district. At this time it was
held to be identical with the korrigum and called, accord-
ingly, the Senegal hartebeest. In 1892 Matschie described
the topi as a new species, Damalis jimela, basing his descrip-
tion on a pair of horns from the southeastern shore of the
Victoria Nyanza and a painting by Richard Bohm of speci-
mens from central German East Africa. None of the color
characters given by Matschie apply to his race, but the
horns may be taken as representing the race. More re-
cently, in 1907, Lydekker renamed the race, se/ousi, on speci-
mens from the Uasin Gishu plateau. Still more recently
another name, phalius, has been applied by Cabrera to a
white-faced specimen from the same plateau shot by Senor
Huerta. This specimen represents an abnormal or extreme
type of individual variation of the topi. Occasional speci-
mens are found in East Africa marked by a whitish or buffy
face blaze and showing a tendency toward the white face blaze
of the closely related blesbok and bontebok of South Africa.
The topi resembles closely the tiang in coloration and
proportions, but may be distinguished by its darker cinna-
mon-rufous coat, longer pelage, and smaller horns. From
other East African antelopes it differs conspicuously by the
silvery sheen of its coat which gives it a resplendent effect
very similar to that of watered silk.

The topi offers almost as conspicuous an instance of dis-
connected distribution as the square-mouthed rhinoceros.
In East Africa it is abundant along the seacoast in certain
regions; and in the western part of the country it is abun-
dant in certain regions. From the regions between it is
totally lacking, although to all outward seeming they are
physically of precisely the same character. Moreover, ani-
mals of the same or very closely allied species or sub-
species are found in regions along the White Nile and in

cone

WILDEBEEST AND HARTEBEEST 353

the Abyssinian neighborhood; and yet here again there are
great stretches of similar territory in between in which they
are not found at all. The problem is all the more puzzling
because the topi is often very plentiful where it exists at all;
if it was everywhere rare, it would be easy to understand
why it might have died out in certain places. It is another
instance of how very much we yet have to learn from the
book of nature. Many of the problems connected with the
distribution and abundance of African antelopes are, with
our present knowledge, insoluble. Compare, for instance,
the distribution of the roan antelope—treating the various
forms together—and the topi—also treating the various
forms together. The topi is usually abundant wherever
found at all; the roan is usually much less abundant where
itis found. The roan is at home in surroundings as diverse
as if they were in different zones and different continents.
We found them among the cold, wet mountains, south of
the Uasin Gishu, where a moose would have found the cli-
mate entirely congenial, and also on the hot, parched flats
of the Lado, with a vegetation as utterly different as was
the climate itself. Yet there are vast reaches of country
from which it is entirely absent; and we never came across
it in such abundance as we often found the topi. The topi
kept everywhere to country of substantially one kind; and
yet, for no apparent reason, its distribution in this kind of
country is narrowly and irregularly limited. It abounded
in company with one kind of hartebeest on the Loita Plains,
and in company with another kind on the Uasin Gishu;
but it was absent from precisely similar country—as the
Athi Plains—on which one or the other of the hartebeests
swarmed. The roan, except for its face markings, is on the

354 AFRICAN GAME ANIMALS

whole concealingly colored. The topi is one of the most
conspicuously colored of all the antelopes, being inversely
countershaded. Only the wildebeest and the old bulls of
the sable, Nile lechwi, and white-eared kob have a coloration
more clearly advertising. Yet the difference in coloration
is evidently of not the slightest consequence as regards any
effect on the respective life histories of the two animals;
wherever found at all, the highly conspicuous topi—which
is advertisingly colored under practically all conditions—is
much more plentiful than the roan, whose coloration really
does have a certain concealing value under certain circum-
stances. But neither trusts to concealment at all when on
the plains, and while the roan in certain places in the woods
may make use of cover to conceal itself the topi never does
at all. The topi is almost as conspicuous as a wildebeest; in
its haunts only a wildebeest would be observed farther off,
and it never seeks to escape observation, trusting purely
to its vigilance and its sense for safety.

In its habits, as in its outward appearance, the topi is
a somewhat less violently accentuated hartebeest—just as
in the wildebeest the characteristic habits and aspect of
this group of antelopes reach their most extreme and eccen-
tric expression. ‘The topi frequents precisely the same type
of country as the hartebeest, when the two are found in
the same locality, and it grazes, it rests, and goes to water—
in places at stated times, in others at irregular intervals—
just like its congener. Often I found herds of the two species
intermixed, and often other game, such as wildebeest, zebra,
and gazelle, were with them. Often herds would be found -
exclusively of one species. We have seen a couple of hun-
dred topi in a herd; and small parties and single bulls are

TOPI DAMALISCUS, MALE

Shot by Theodore Roosevelt, Loita Plains, B. E. A.
United States National Museum, Washington, D. C.

TOPI DAMALISCUS, MALE TOPI DAMALISCUS, ADULT MALE

Shot by Theodore Roosevelt, Loita Plains, B. E. A. Uasin Gishu Plateau
United States National Museum, Washinshon, D. C.

MOUNTED SPECIMENS OF THE TOPI DAMALISCUS

pie
kD *e ve i;

Tote ee

WILDEBEEST AND HARTEBEEST 355

also common. Like the hartebeest, the topi often stands
on ant-hills to watch and look round. In places we found
the topi shyer than the kongoni, and in places both were
tame. ‘The topi is said to be an even swifter and more en-
during runner than the kongoni. However this may be, it
is certainly both swift and enduring, and it is, morever,
very agile; when a herd is suddenly alarmed the individuals
spring clear over one another’s backs in the hurry and con-
fusion, and on one such occasion we saw one jump over an
ant-hill higher than a man’s head. The bulls fight fiercely
among themselves. On the Uasin Gishu plains we saw a
big herd of topi one of whose members had a white face
like the South African blesbok.

There are few more beautiful sights than a herd of topi
seen close by in the bright sunlight of the great African
plains. They are not ungainly, like the hartebeest, and their
coats are not only beautifully colored but have, in life a
wonderful iridescent sheen; the muscles ripple beneath the
satin skins, and every movement shows lithe and abounding
vigor.

A topi bull weighs two hundred and eighty or three hun-
dred pounds; a cow twenty or thirty pounds less. In spite
of its usual wariness, the topi is subject to fits of silliness
and curiosity that expose it to reckless slaughter by rifle-
bearing hunters. It is very tenacious of life, and will go
long distances after receiving what would seem to be crip-
pling wounds.

The body coloration is a bright cinnamon-rufous over-
laid everywhere by a silvery sheen which gives the coat a
resplendent effect. The red color: is deepest on the head,
_ throat, and sides and lightest on the rump, hind quarters,

356 AFRICAN GAME ANIMALS

and tail, where it fades to pure cinnamon. The shoulders
are marked by a broad black patch which extends down on
the forelegs as far as the knees and completely circles the
upper part of the leg. ‘The hind quarters are marked by a
much larger black patch which extends down on the limbs
as far as the hocks above which it forms a complete band
around the leg. The legs below the knees and hocks are
uniform tawny, the color being slightly deeper in front
than behind. Some specimens show a median, narrow, wal-
nut-brown stripe on the back of the pasterns. The ter-
minal half of the tail is furnished with a long tuft and crest
of black hair. The under-parts are cinnamon-rufous, like
the dorsal surface, with the exception of the groins, the
posterior part of the belly, and the axilla, which are buffy.
The dorsal surface of the head from the crown to the muz-
zle is deep-black in color as far down as the level of the eyes.
The sides of the head and throat are a deep-bay or blood-
red color, which is sharply defined against the black. The
upper lips and the chin are tawny. A black stripe or bar
is usually present below the eye, extending from the black
snout patch to behind the eye. Often this bar is absent or
represented only by a spot below or behind the eye. Situ-
ated in front of the eye is the rounded anteorbital pore
surrounded by a narrow band of black naked skin. The
backs of the ears are tawny, with the terminal third walnut-
brown and the inner side white. The sexes are identical in
coloration, but the newly born young differ widely in color.
At first they are a uniform light-fulvous or buffy-ochra-
ceous, very much the color of sable at the same age. This
lighter color, however, is soon lost and the cinnamon coat
and black leg patches assumed. The black face blaze lags
somewhat behind the leg markings and is buffy and only
indistinctly indicated until the budding horns have reached
at least four inches in length.

An adult male topi shot by Colonel Roosevelt on the
Loita Plains measured in the flesh: head and body, 67
inches; tail, 19 inches; hind foot, 22 inches; ear, 8 inches.
These dimensions represent well the average in males, but
the females are considerably smaller, the length being some
4 inches less and the hind leg 1 inch less. The skulls of the
males range in length from 15 to 1534 inches. The horns .

“ite

es

Pee etuttnaca ee

ere : \ :
2

—_

ores, cart
i Seen...”

Abaya L

/

* aw comm cae” f

ast

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ce

MAP 7—DISTRIBUTION OF THE RACES OF THE DAMALISCUS

1 Damaliscus korrigum tiang 2 Damaliscus korrigum jemala
357

358 AFRICAN GAME ANIMALS

of the males are decidedly heavier or thicker than those of
the females and are slightly longer as well. They do not,
however, vary much in size. The large series in the Na-
tional Museum range from 13 to 1734 inches. Ward re-
cords a specimen 19% inches in length from East Africa.

A large series, some thirty specimens, have been ex-
amined in the National Museum, from the Loita Plains and
Uasin Gishu Plateau. Much variation in the black stripe be-
low the eye is shown by this material, in fifty per cent of which
it is quite absent or only indicated by a spot behind the eye.
Several of the specimens show a tendency toward a white
face blaze. In one specimen from the Loita Plains there is
a strong intermixture of white hair throughout the whole
black face blaze, while in others this mixture is confined
either to the tip of the snout or the forehead between the
horn bases. The topi has been reported from southwestern
Uganda in the Ankole district by Delme-Radcliffe and in
Buddu by Gedge. This elevated portion of Uganda marks
its extreme northwestern range in the Nile watershed.

Hunter ANTELOPE
Beatragus

Beatragus Heller, 1913, Smith. Misc. Coll., vol. 60, No. 8, p. 8; type Da-

maliscus huntert.

The Hunter antelope, owing to its peculiarities of body
form and horn shape, has recently been separated from the
genus Damaliscus and named Beatragus by Heller. In the
elongate shape of the snout it‘approaches the hartebeest,
but the occipital portion of the skull is rounded as in the
damaliscus antelopes and not overhung by a horn pedicle.
The horns arise independently behind the orbits, curve
outward, and then slant upward and backward, extending
parallel in direction for the greater part of their length.
Viewed from the front, they are broadly U-shaped and quite
impalla-like in general shape and size. They are ringed on
the basal two-thirds of their length, the tips being smooth
and acutely pointed. The first lower premolar tooth is usu-
ally wanting, the cheek-teeth of the lower jaw numbering
but five on a side. Beatragus differs from the hartebeests

WILDEBEEST AND HARTEBEEST 359

and damaliscus further by the white tail tuft and the white
bar across the forehead between the eyes. The body form
is more slender than that of the hartebeest and the withers
are lower. The Hunter antelope is the sole representative
of the genus, which is restricted in range to a small area of
park-like or plains country on the north side of the Tana
River. It is doubtless a specialized offshoot of damaliscus
and not an intermediate connecting link between the harte-
beest and the latter.

HunTER ANTELOPE
Beatragus huntert

Native Name: Galla, herola.
Damalis huntert Sclater, 1889, Proc. Zool. Soc., pp. 58 and 372, pl. XLII.

Rance.—North side of the Tana River from the village
of Durani up the river as far as Massa at least, that is, from
a point seventy miles inland to about one hundred and
twenty miles up the river. North of the Tana River it ex-
tends parallel to the coast as far as the latitude of Port
Durnford.

Hunter was undoubtedly the first sportsman to meet
with the species which was named for him by Sclater.
During a sporting trip in 1887 on the lower Tana River he
shot the first specimen near the village of Durani, which
is some one hundred and fifty miles from the mouth by way
of the tortuous river channel but actually only seventy miles
in a direct line. He recognized the antelope as new, and
carefully preserved, and photographed the first specimen he
secured. Farther up the river they were met with in greater
numbers as far as Massa, at whith point the hunting party
turned back to the coast. Hunter’s specimens have for
many years remained unique in the British Museum, but
recently two others were added by Gilbert Blaine, who ob-
tained them on his way down the Tana on the north side
near Hunter’s original locality. “Two other specimens have
been examined at the Nairobi Club in British East Africa,
which were obtained by Major H. J. Kirkpatrick on the
Tana River. Owing to the isolated and restricted nature
of its habitat, less than half a score of sportsmen have met
with this antelope.

360 AFRICAN GAME ANIMALS

The dorsal color is a uniform ochraceous-tawny without
any indication of a lighter rump patch. The distinctive
color marks are the white tuft of the tail and the white
band across the forehead between the eyes. The under-
parts are whitish or light-buffy and the ears show dark
tips with white inner surfaces. ‘The sexes are alike in color.
The coloration of the young is not recorded, but it doubt-
less closely resembles that of the adult.

Hunter met with this antelope on open, grassy plains
bordering the Tana, in herds of fifteen to twenty-five in-
dividuals. He also found them occasionally in scattered
bush country. They were met with occasionally along the
river throughout a stretch of about one hundred miles bor-
dering the north or east bank but were local in distribution,
owing to the interrupted character of the plains country.

No flesh measurements are recorded. The body size is
somewhat less than that of the Coke hartebeest. The skull
of the type measured 12,% inches in basal length by 5,
inches in greatest breadth at the orbits. The horns of the
male exceed somewhat in length and size those of the female.
Ward records the horn measurements of six male and two
female specimens. The males in this series average 24
inches in length, with a spread at the tips of 13 inches. The
largest pair are 26% inches long, the widest spread at the
tips 1534 inches. The largest female pair measure 21144
inches in length by 11 inches in spread at the tips.

BRINDLED WILDEBEEST
Gorgon

Gorgon Gray, 1850, Proc. Zool. Soc.; type G. taurinus.

The brindled wildebeests show marked differences in
shape of skull and horns from the white-tailed gnu, and the
two cannot with propriety be grouped together under the
same genus, as is commonly done. The snout in the brin-
dled wildebeest is long, the nasal bones being two-thirds
the length of the skull; the orbit is small and without a
bony ridge below it; the premaxillary bones are rounded at
their tips and the lachrymal bone has a shallow hollow or
fossa in front of the orbit. The white-tailed gnu has a

WILDEBEEST AND HARTEBEEST 361

short snout, the length of the nasal bones being only one-
half that of the skull; the orbit is large and has a ridge
below it; the premaxillary bones are square at their tips
with angular corners, and there is no evidence of an ante-
orbital fossa in the lachrymal bone. The horns of Gorgon
are lateral in direction while those of Connochetes, the type
of which is the white-tailed gnu, curve forward over the eyes
and then upward and backward. The two wildebeests have
furthermore marked differences in the body shape. The
brindled is quite bison-like, with enlarged shoulders, high
withers, and small hind quarters while the gnu has enlarged
knobbed hips and comparatively small shoulders or withers.
In general shape the skull of the brindled wildebeest ap-
proaches more closely that of the African buffalo than the
gnu. It is distinguishable from the buffalo only by the dif-
ference in horn shape and narrower cheek-teeth. It is re-
markable how closely the bones of the snout of these two
members of the Bovide agree in shape and proportions.
The brindled and the white-bearded wildebeests are the sole
representatives of the genus Gorgon. ‘They range throughout
eastern Africa from the Vaal River, the northern affluent of
the Orange, to the Athi Plains of British East Africa.

WuiTE-BEARDED WILDEBEEST
Gorgon albojubatus

The white-bearded wildebeest is distinguishable from
the brindled species, as its name signifies, by the white or
light color of its throat mane. There is no other important
or obvious difference, both species having similarly dark
bodies which show a tendency to dark cross bands due to
a worn or ragged condition of the hair along transverse
streaks. Owing to the absence of specimens which show
an intermediate coloration of the throat mane between the
black of the brindled and the white or buffy of the white-
bearded, the latter must be considered a distinct species.
The white-bearded is the northern representative of the
brindled wildebeest type and ranges from central German
East Africa northward to the southern slopes of Mount
Kenia in British East Africa.

362 AFRICAN GAME ANIMALS

With the single exception of the nearly extinct white-
tailed gnu of South Africa, the common African gnu or
wildebeest is psychically—and therefore in all its life habits
—the most interesting and original of all African ruminants.
It is as totally different in conduct as in aspect from all
other antelopes. In its temper it has something both of
the ferocious and sinister and of the queer and freakish,
the eccentric and fantastic. It is by nature an extremely
savage creature. When tamed it becomes excessively dan-
gerous as it grows old, attacking with mortal fury every
human being, from sheer homicidal anger and lust for death
and mischief. When wild it will charge if cornered or close
pressed when wounded. But its moderate bulk and blunt
horns furnish it with bodily weapons so poor as to render
it entirely helpless when assailed by either lion or man, and
knowledge of this bitter fact prevents it from ever venturing
combat with either of its great foes. It is as wary as it
is curious and ferocious, and no animal is harder to approach
on the bare, open plains where it dwells. It is unusually
noisy for an antelope, continually uttering its short grunt.

Where we saw the wildebeests, at the extreme northern
end of their range, they were exclusively beasts of the open
plains. We rarely saw them even in thin thorn scrub into
which the hartebeests not infrequently ventured. Only
the gazelles were equally persistent in shunning cover and
in keeping to the grassy flats. It may be that elsewhere
the wildebeests venture more freely into thin, dry scrub.
But everywhere the animals are so conspicuous that they
make not the slightest effort to hide or escape notice.
Owing to its size—a big bull weighs from five hundred to
six hundred pounds—and its dark coloration, which seems

phy ¥ i
Ce 2 eh eed

LOITA WHITE-BEARDED WILDEBEEST, MALE

Loita Plains, B. FE. A.
From a photograph by Kermit Roosevelt

WHITE-BEARDED WILDEBEEST
From German Fast Africa In the New York Zoological Park

WHITE-BEARDED WILDEBEEST

WILDEBEEST AND HARTEBEEST 363

black at a distance, the wildebeest is more quickly seen, and
at a greater distance than any other game below the rank
of the “heavy game,” as the Boer hunters call the elephant,
rhino, giraffe, and buffalo. Its one object is to keep in such
a position that it can itself see its foes; it does not in the
least mind being seen by them. It is extraordinarily tough
and tenacious of life, paying amazingly small heed to what
would seem to be crippling wounds; and it has great speed
and endurance. Indeed, it is probably the speediest of East
African antelopes in a long run. The cowboys of the Buf-
falo Jones party were unable to run one down, although they
pursued it the greater part of a day, using relays of fresh
ponies on the Loita Plains.

Wildebeests are usually found in herds of several score
individuals, mostly cows and calves, with young animals,
and one or more herd bulls; the latter are apt to fight des-
perately with one another on the slightest provocation, and
if there is one acknowledged master bull in the herd he will
occasionally make a dash after one of the others and drive
it off at full speed. Often there are outlying sentinels in
the neighborhood of a herd. Old bulls go singly or in small
parties, either by themselves or in a mixed herd with harte-
beests, zebras, and gazelles. When fighting the bulls fre-
quently drop on their knees and plunge together, head on,
with a resounding crash. The herds feed at all hours—
wildebeests are exclusively grazers—but during the hottest
part of the day are apt to lie down, when at a little distance
they look exactly like American bison at rest on the prairies
in the old days. Doubtless they also come to drink at all
hours; however, we personally never knew them to go to
the water at night, but only in the morning or afternoon;

364 AFRICAN GAME ANIMALS

they are more watchful and wary than other game, and
probably dislike approaching a pool after nightfall for fear
of the lion. If hartebeests or zebras are with wildebeests
as they approach water, the former generally lead, the
wildebeests hanging back. We have seen this again and
again, and but once did we see the wildebeests of a mixed
herd take the lead in approaching a pool. Mr. Abel Chap-
man, in “On Safari,” has given an amusing and interesting
description of the way that a herd of wildebeests will some-
times wait about until a hartebeest turns up as leader to
take it to water. As is true of all game, wildebeest are
much less apt to detect a foe lying in wait for them in the
path they are following than to detect one endeavoring to
approach them. If the direction in which a herd is travel-
ling can be ascertained, a stunted, well-nigh leafless bush
will serve as ambush sufficient for any one who crouches
absolutely motionless in its shade. It is motion that
catches the eye of game, and, moreover, any color loses much
of whatever conspicuousness it possesses if broken light and
shade fall on it. Time and again we have thus crouched
beside or behind a bush and watched files of zebra and
hartebeest and parties of wildebeest and gazelles and water-
buck canter by within less than a hundred yards—often
not a score of yards away. They never noticed us.
Usually wildebeest are very local in their habits, the
same herd keeping within a radius of three or four miles.
At one camp where we spent a week or so we always found
one herd of wildebeests kept within a couple of miles of a
boldly shaped hill; and three or four miles off a single bull
lived by himself, occasionally joining a party of gazelles
and hartebeests but never coming near his own kinsfolk.

WILDEBEEST AND HARTEBEEST 365

When water grows scarce the herd may have to trek to it
a dozen miles each day, and if drought dries up the pastur-
age then all the wildebeests of a locality may have to shift
their ground. Under such conditions the shift may be only
for a few miles, or the animals may gather together to the
number of several thousand and travel to far-off pastures
in a great herd, like a herd of our own bison thirty odd years
ago. ‘There must be some kind of telepathy which thus in-
fluences all the animals of a given district to travel off in a
certain direction, only to halt when they have reached some
place which perhaps none of their kind have visited since
years before the oldest among them were born. Lemmings
and South African springbucks make such treks; the bison
occasionally did; and occasionally wildebeest make, or made,
such treks in addition to their ordinary shifts. The migra-
tions of East African game, both seasonal and irregular,
have as yet only been imperfectly worked out. There are
some places in which the game spend the whole year and
others where the game either shift their ground annually
or at irregular intervals.

As one approaches a herd of wildebeest they stand and
gaze, sometimes snorting or grunting, their heads held high
and their manes giving them a leonine look. The bulls
may pace up and down, pawing the ground and lashing
their tails. Then, as the hunter comes nearer, down go
their heads, and off they start, rollicking, plunging, and
bucking before they settle into a hard, level gallop. Sud-
denly they all pull up short and wheel to look back; or while
the herd is still cantering in a curve a couple of bulls or old
cows will start at full speed in the opposite direction, cur-
vetting and prancing as they pass their companions in re-

366 AFRICAN GAME ANIMALS

verse order. Then the whole herd may gallop in a curve,
strung out in line or in a mass, some of the individuals
going through queer gymnastics, kicking and flourishing
their long tails; or a bull will almost stand on its head to
toss up the dust with its horns; or two bulls may relieve
their feelings by dropping on their knees for a moment’s
furious fight, clouds of dust floating above them as they thus
curve and zigzag. Finally they conclude to leave the neigh-
borhood; then they stop their ferocious and whimsical ca-
pering, and, their heads down, away they tear with beauti-
ful, strong action.

They are very wary, and keep well out of rifle range, but
they are also very curious and hang about, just out of shot,
watching the hunter. In the same fashion they watch the
lion, going fairly close to him, as they evidently feel confi-
dent that he cannot catch them unless so close that they
cannot get under way before he is on them. Once we
came across an old bull wildebeest watching a lion and
lioness from a distance of about sixty yards; it was in the
afternoon, and the lions obviously desired the bull for
supper; but, although very much on the alert, he was not
in the least disturbed by their proximity, and evidently
knew as well as they did that at that distance they could
not hope to catch him. Some observers have suggested
that the game know when the lion is hungry and do not
fear him except at such moments, and that this accounts
for the small heed they pay to him if they see him on the
plains in the daytime. We doubt this theory. Our im-
pression is that they are altogether too accustomed to the
lion to get panic-struck about him unless he rushes at them
from ambush. They know that they can outrun him, and

WILDEBEEST AND HARTEBEEST 367

if they see him walking or trotting over the plain they know
he is not hunting them and they feel no uneasiness about
him. In Africa there is no hour of the day or night when
the game is safe from its enemies; each hunted creature is
always in possible deadly peril, and therefore each grows
accustomed to the peril, and pays no heed whatever to it
unless it is at that moment imminent. A lion passing
over the plain is no more to them than a shark passing
under a vessel’s stern is to a sailor aboard her. A wilde-
beest, zebra, or hartebeest gives itself no more concern
over such a lion than the sailor gives himself over the
shark, and for the same reason—because there is no
danger.

These same observers, by the way, speak of the plains
game as inert and stupid compared to bush game. This,
also, is not in accord with our observations. The wilde-
beest is as alert, wary, and intelligent as any game of the
woods or bush; and animals that are found both in the
forest and on the open plain, as the waterbuck, do not
differ, as far as we can see, in wariness and intelligence
from those that dwell only on the plains. We have some-
times found waterbuck more wary than hartebeest, and
sometimes less; but in only a very few instances have we
ever found individuals of either species as wary as wilde-
beest. However, it is always necessary to keep in mind
not only that animals of the same species show wide in-
dividual differences and wide differences according to lo-
cality, but also that the same individuals show astonishing
differences in conduct at different times. We have found
hartebeests and gazelles very wild in the morning and yet
in the afternoon well-nigh heedless of us even when we

368 AFRICAN GAME ANIMALS

were to windward; and we have seen a flock of the usually
tame Tommy gazelles run off in alarm, while a solitary
wildebeest bull, which was with them and was usually
far more shy than they were, displayed no emotion what-
ever about our presence.

We came across many carcasses of wildebeests that had
been killed by lions, and we never saw any more signs of a
struggle than with zebra or hartebeest. Doubtless they
are sometimes killed near water, but all those whose bodies
we saw had been killed in the open plain during the night.
Evidently the lion had either stalked them or had lain
motionless until, as they grazed or walked toward new feed-
ing-grounds, they approached him. In one or two cases
the marks showed that the lion had struck the claws of
one paw into the wildebeest’s head to hold him, a favorite
trick of the lion with heavy or formidable game. In some
cases the wildebeest had been seized on a flat, bare plain
without a vestige of cover, the grass being cropped short.
It was extraordinary that even in darkness so wary a beast
should let so big a foe approach to within a few yards.
Probably in such cases it was the wildebeest itself that ap-
proached, the lion’s sinuous length being moulded along the
ground, in the darkness, until the munching, grass-cropping
herd walked slowly within reach of his tremendous rush—
for a lion’s furious dash, whether on prey or foe, is much
more apt to bearush thana spring. Doubtless in the lion’s
hunting at night—his regular hunting time—smell counts
for much more than sight in guiding him to his prey;
details like the prey’s color are of no consequence what-
ever; his eyes are required chiefly to guide him in the
actual grapple. |

WILDEBEEST AND HARTEBEEST 369

Key To THE Races oF albojubatus

Legs lighter, drab or tawny-olive; horns more horizontal in direction, not
curved down below level of orbit; body size larger
albojubatus

Legs darker, olive-brown or sepia; horns curved downward well below
level of orbit; body size smaller mearnst

Atut WuitrE-BEARDED WILDEBEEST
Gorgon albojubatus albojubatus

Native Name: Swahili, nywmbu.
Connochetes taurinus albojubatus Thomas, 1892, Ann. & Mag. Nat. Hist.

(6), LX, p. 388.

RancE.—From the Athi Plains and south bank of the
Tana River near the Ithanga Hills, in British East Africa
southward throughout the highlands of the coast drainage
to Kilimanjaro, and thence through the Rift Valley to cen-
tral German East Africa.

A head of the white-bearded wildebeest shot by Jackson
on the Athi Plains furnished Thomas, in 1892, with the mate-
rial for his original description of this race. Notwithstand-
ing that the white throat mane of this form is a really con-
spicuous difference from the black mane of the brindled
wildebeest, it was for many years supposed to be indis-
tinguishable from that species. Speke and Grant, in 1863,
met with it on the Kigani River, opposite Zanzibar Island,
and brought back with them two heads which were assigned
by Sclater to the brindled widebeest. Later Fischer, Hilde-
brandt, Abbott, Jackson, Willoughby, and many others
shot specimens in the vicinity of Kilimanjaro under the
supposition that they were true brindled wildebeest. In
1905 Neumann described, from a specimen now living in the
Berlin Zoological Garden which came from the southern
slope of Kilimanjaro, another race which he named heck.
This he distinguishes from the white-bearded by the light
grayish-brown forehead and lighter body color. Such color
differences, however, are peculiar to the female sex and to
immature animals and are of no racial significance. Sports-
men have thought that they detected two quite different

370 AFRICAN GAME ANIMALS

species of wildebeest in East Africa associating together in
the same herds. One of these has been assigned to the
typical brindled wildebeest with a black throat mane, and
the other has been known as the white-bearded wildebeest.
In the ‘‘ Book of Antelopes” these two forms are considered
and their ranges and characteristics stated. How this error
has crept into the literature it is difficult to conceive, for
no specimens representing a black-maned form have ever
been collected in the region. The series of twenty-two
specimens in the National Museum from British East Af-
rica shows no variation from a wholly white or light-colored
throat mane. In the southern portion of German East
Africa specimens of the brindled wildebeest may occur.
The southern limits of the white-bearded are not known.
As no specimens showing intermediate coloration in the
throat mane have been reported, we are not yet justified in
considering the white-bearded a race of the brindled.

The Athi white-bearded wildebeest may be distinguished
from the Loita race by the lighter-colored legs, the more hori-
zontally directed horns, and the larger body size. The legs
are drab in color, like the body, and never as dark as the
olive-brown of the Loita race, and the body color is less
blackish, the chest, shoulders, and lower sides never assum-
ing a deep seal-brown or blackish cast. ‘The horns extend
in a horizontal plane from the head and curve downward
comparatively little from their bases. Adult male speci-
mens vary but little in body size and exceed but slightly the
females. The flesh measurements of a male from the Kapiti
Plains, shot by Colonel Roosevelt, were: head and body, 77
inches; tail, 24 inches; hind foot, 17 inches; ear, 8 inches.
The skull of this specimen has a length of 193% inches. An
average female skull measures 18%% inches in length. The
horns of adult males average 25 inches in greatest spread and
about 20 inches in length, but the record head exceeds these
dimensions by some 4 inches.

Loira WHITE-BEARDED WILDEBEEST
Gorgon albojubatus mearnst

Native Name: Masai, olengat.
Gorgon albojubatus mearnsi Heller, 1913, Smith. Misc. Coll., vol. 61, No.

175 1Pasdl

1

WILDEBEEST AND HARTEBEEST 371

Rance.—From the Loita Plains and Rift Valley in the
vicinity of Mount Suswa southward to German East Africa
and westward through the headwaters of the Victoria
Nyanza drainage to Speke Gulf.

Wildebeest from the Loita Plains of British East Africa
secured by the Smithsonian African Expedition under the
direction of Colonel Roosevelt have clearly demonstrated
the racial distinctness of the form occupying the extreme
southwestern corner of British East Africa. A specimen
shot by Doctor E. A. Mearns near the ford on the South-
ern Guaso Nyiro River has been selected for the type of
the race which now bears his name. The Loita Plains
white-bearded wildebeest differs from the form inhabiting
the Athi Plains by the decidedly darker color of their legs,
which are olive-brown or sepia in the old males and some-
what lighter in the females. The body color shows a tend-
ency to become darker or quite blackish on the chest,
shoulders, and sides. The horn differences consist in the
more downward curve of the horns from the base, the lower
margin falling well below the orbit and quite on a level with
the condyles at the back of the skull. The skulls average
smaller, indicating smaller body size.

The dorsal color is drab-gray, spotted and banded by dark
patches due to the tips of the hair becoming dark-brown as
though representing an old, worn pelage, the new only being
drab-gray. ‘This faded condition of the hair is not seasonal
but is a chronic condition in the wildebeest, and has given rise
to the color pattern and to the name brindled commonly ap-
plied toit. The limbs are darker than the body and are uni-
form olive-brown deepening somewhat on the pasterns. The
chest, fore part of belly, and lower sides are clove-brown, and
much darker than the back; the chest medially is black. The
groins, axille, and inside of the legs are drab-gray like the
back. The tail is like the back in color and furnished along
the lower surface and at the tip with long black hair forming
a terminal tuft fully as long as the tail itself. The neck is
like the back, in ground-color but shows ragged cross-
bands of dark hair bases. A mane of long black hair extends
the whole length of the nape and is continued down the
back by a black line to the rump. The throat is marked by
a mane of long buffy or whitish hair from the chin to the

372 AFRICAN GAME ANIMALS

chest. The top of the snout from the interorbital region to
the muzzle and far down on the sides to the lower level of
the eyes is deep-black, the blackness at the muzzle spread-
ing down around the mouth and uniting with the black
chin. The upper lips are whitish at the tip of the snout and
clove-brown on the sides. The sides of the head are drab-
gray, like the neck, with a lighter streak bordering the black
face blaze from the eye to the angle of the mouth. The crown
is seal-brown and shows some contrast to the black fore-
head. The back of the ears is black and united with the
dark-brown patch by a narrow ridge of color, but the rest of
the base and the inside is pale drab-gray. Some males show
a definite walnut-brown band across the forehead between
the eyes, while others are marked by short, white chevrons
similar to the specimen collected by R. J. Cuninghame on
the Loita Plains and referred to by Lydekker in his “Game
Animals of Africa.’’ The female differs in color from the male
by having the crown and interorbital region lighter-bistre or
drab-brown in contrast to the black snout and lacking the
black color on the chest, shoulders, and lower sides which are
drab-gray, in conformity with the back. The legs are also
lighter, being, as a rule, little darker than the drab body.
The throat mane is shorter and more scanty than in the male.
All of these color differences are also shared by the imma-
ture males. The newly born young, before the horns have
started to bud, are much lighter-colored than the female.
The body is at this age uniform vinaceous-buff with whitish
or cream-buff under-parts and legs. ‘The mid-dorsal line
of the nape and back is marked by a short black mane
from the occiput to the rump. The whole top of the head
from the ears to the tip of the snout and well down on the
sides below the eyes is quite uniform blackish or deep seal-
brown, in this respect resembling the adult male. The chin
is black as in the adult. The sides of the head, throat, and
neck are vinaceous-buff like the body. The ears are wholly
black on the back and whitish on the inside. The tail is
like the body in color and only black at the tip.

The flesh measurements of this race agree closely with
those of typical albojubatus, although the skull averages
¥Y% inch less in greatest length. The horns average 22
inches in greatest spread which is somewhat less than those

Mt. Lololokw!

Ae F

Mt. Varagess

ow

MAP 8—DISTRIBUTION OF THE RACES OF THE WHITE-BEARDED
WILDEBEEST AND HUNTER ANTELOPE
1 Gorgon albojubatus albojubatus 2 Gorgon albojubatus mearnst
X Beatragus hunteri

373

374 AFRICAN GAME ANIMALS

of Athi specimens. ‘The largest pair in the series of 11 males
has a width of 2414 inches and a length of 21. A series of
I5 specimens from the Loita Plains have been examined in
the National Museum which represent localities in the
Southern Guaso Nyiro drainage and the headwaters of the
Amala River near the German boundary.

HARTEBEESTS
Bubalis

Bubalis Lichtenstein, 1814, Mag. Nat. Fre., Berl., p. 154; type B. buselaphus,
the bubal hartebeest of Algeria.

The hartebeests form a very distinct genus. They are
easily recognized by their elongate heads, which have a pecu-
liarly straight dorsal profile owing to the great projection
backward of the horn bases or pedicle beyond the condyles
of the skull. No other group has such a union of the horn
bases into a pronounced pedicle from the ends of which the
horns fork. ‘The hartebeest has extremely high withers and
a narrow body. The tail is moderately long and tufted.
The ears are narrow but of fair length. The horns are alike
in the sexes and are usually curved at sharp angles outward
and then backward or inward. They are of moderate
length, usually equalling the head. The female is much
smaller in body size than the male, being usually a third less
in weight. ‘The calves are lighter-colored than their parents,
being a uniform buffy-fulvous. The skull is without definite
anteorbital fossa, having merely a shallow depression in front
of the orbit. The horn pedicle is very long and forks pos-
teriorly to the condyles, and is in breadth less than the
length from the fork to the orbit. The nasal bones are very
long and extend forward as far as the tip of the snout.
The mammez are reduced to two. The genus includes six
or seven well-marked species with their geographical forms.
Although the various species show considerable distinction,
their ranges nowhere overlap, and as a genus much discon-
tinuous distribution occurs. The fossil species described
are a Pliocene species from India and two Pleistocene
species from the Transvaal, one of which, B. priscus, had
horns very similar to cokez but was a somewhat larger
animal.

WILDEBEEST AND HARTEBEEST 375

The genus is confined to Africa, where it ranges from
Algeria and the Abyssinian highlands south to the Cape but
is absent from the Congo forest area. Arabia has been
credited with a hartebeest but without any satisfactory evi-
dence. The error, however, has gotten well established in
zoological literature and is based on the statement of Canon
Tristam concerning the occurrence of the bubal hartebeest
in Eastern Syria in the vicinity of the Dead Sea from
rumors received from the Arabs inhabiting that district.

We encountered swarms of Coke hartebeest in eastern
and middle East Africa and swarms of the far bigger and
handsomer Jackson hartebeest west of them, while on the
border line between these were here and there colonies of
the closely allied Nakuru hartebeest. In Uganda and
along both banks of the White Nile we found hartebeest
akin to the Jackson. The Swahili name for every kind of
hartebeest is kongoni. The exact relationship and geo-
graphical distribution of these species and subspecies are
discussed in the technical part of the work. The habits of
all the species are substantially the same; that is, the in-
dividuals of one species in one locality differ more in habits
from the individuals of the same species in a totally different
kind of locality than from the individuals of another species
in an exactly similar locality. We therefore discuss them
under one head. Indeed, even in aspect the chief difference
that strikes the observer is one of size; the Coke hartebeest
weighs on an average from three hundred to three hundred
and fifty pounds, while the Jackson, a far finer beast in
every way, is more than half as long again, reaching a weight
of five hundred pounds and over. There are marked horn
differences in addition, however.

Hartebeests, where not exterminated by man, range

376 AFRICAN GAME ANIMALS

over most of the continent of Africa, outside of the moun-
tains, the heavy forest, and the true desert. In individual
abundance they unquestionably surpass any other of the
genera of big game with the doubtful exception of the
gazelles, if these can be called big game; their range is
greater and they are more plentiful within it. Taken asa
whole, in the parts of Africa we traversed, hartebeests were
at least three or four times as numerous as any other horned
animal. We saw no zebra in Uganda or along the Nile;
in East Africa they were more abundant than any of the
buck except the hartebeests, but the latter outnumbered
them about three to one. There were no hartebeests on
the Northern Guaso Nyiro, where we found both kinds of
zebra, their ranges here overlapping. Why the hartebeests
were not found in this region we cannot guess. It is an
illustration of the queer and seemingly causeless and ca-
pricious discontinuous distribution of so many African game
animals. The ordinary zebra, the bonte-quagga, in its
various color forms is almost always found in company with
one or another kind of hartebeest; the hartebeests extend
over vast areas to which zebras do not penetrate; but at
this point the zebra flourishes where the hartebeests are not
found. In like fashion the big Grévy zebra is found at
different points of its range—north, south, and west—in
company with various forms of hartebeests, but they are
wanting in other places. In East Africa the small gazelle,
the Thomson, so abundant farther south, also fails to reach
the Northern Guaso Nyiro; yet a form of its big congener, the
Grant gazelle, elsewhere found in company with it, is com-
mon along thatriver. The distribution of the small gazelle is
limited to the highlands and it does not occur below an alti-

KONGONI COKE HARTEBEEST

Shot by Theodore Roosevelt, Athi Plains
Group mounted by J. L. Clark in the United States National Museum, Washington, D. C.

KONGONI COKE HARTEBEEST, FEMALE

From Nairobi, B. E. A.
Presented to the National Zoological Park, Washington, D. C., by W. N. McMillan

KONGONI FROM THE ATHI PLAINS OF BRITISH EAST AFRICA

WILDEBEEST AND HARTEBEEST 377

tude of three thousand feet, but the Grant gazelle occurs
much lower and occupies the low desert tracts. Altitude is
apparently the cause which prevents them from occupying
the Uasin Gishu Plateau lying at a height of seven to nine
thousand feet, which is above the ranges of both gazelles.
The topi is abundant in certain tracts and entirely absent
from big intervening tracts seemingly in no way different.
Such discontinuous distribution reaches its height in the
case of the square-mouthed rhinoceros.

The hartebeest is never found in the dense tropical for-
ests, whether of mountain or lowland. By preference it
haunts the treeless, or well-nigh treeless, open plains; but ac-
cording to our experience it also goes, rather more freely
than the zebra, into the sparse open woods of small, scantily
leaved thorn-trees; and both in Uganda and along the
*Nzoia we now and then found it in fairly thick vegetation,
among trees, bushes, and tall grass. It cannot too often
be said, however, that in these matters equally trustworthy
observers disagree, chiefly because animals of the same
species differ widely in their habits under different conditions
of time and place. Captain Stigand, for instance, says of
these East and Middle African hartebeests that they “ prac-
tically are never seen in bush or in shade; they seem even
to mistrust the proximity of bush.”” As a general proposi-
tion this is unquestionably true, for the hartebeest, like the
wildebeest, topi, zebra, and gazelle, is pre-eminently an
animal of the bare, open plain; but according to our experi-
ence Captain Stigand’s statement is far too broad. We
have frequently come on hartebeests among thorn-trees or
in shade, and sometimes in country where it was impossible
for them to avoid the neighborhood of thick bush. It is

378 AFRICAN GAME ANIMALS

merely a case of difference of observation, doubtless both
accurate. As regards wildebeest, our experience was iden-
tical with Captain Stigand’s, who says that they inhabit
open plains exclusively and are never seen in bush country;
but the French hunter M. Vasse particularly mentions that
in Portuguese East Africa, although they live in the vast,
bare plains they spend the hot hours in the small thorn
woods—a habit we never observed in East Africa, where the
wildebeest herds never left the plains, lying in them during
the hot hours. Along or near the edges of these bare plains
hartebeest may be found grazing in company with water-
buck and impalla; if alarmed the latter will make for the
timber, and the kongoni for the open; but we have occasion-
ally found each species spending hours in the haunts of the
other. In the Lado the waterbuck, kob, and hartebeest
were ordinarily found in precisely the same localities and
often associating together.

Normally, however, the hartebeest is found on the bare
plains, associating intimately with the other plains-dwellers
mentioned above. Its habits are so nearly identical with
those of the zebra that the two animals habitually mix in the
same herd. Where both species are numerous, such mixed
herds are almost as common as herds composed exclusively
of only one or the other species. Wildebeest and gazelle
may be found in the same herd with either zebra or
hartebeest. Sometimes such a mixed herd will consist of
numbers of every species—say a score of zebra, as many
hartebeest, and a dozen wildebeest—sometimes a single in-
dividual of one kind will go with a herd of another kind, a
wildebeest with zebra, a hartebeest with a wildebeest herd, a
beautiful Grant gazelle with a mass of zebra, wildebeest,

WILDEBEEST AND HARTEBEEST 379

and hartebeest; or, a rather comical sight, a shaggy and
ferocious-looking wildebeest bull may dwell with a band
of diminutive and delicate-looking ‘“Tommy” gazelle. We
have seen topi and hartebeest together in big herds, just as
I have seen the common zebra in herds with eland, oryx,
and its big relative the Grévy zebra.

These plains game are very gregarious; and although sin-
gle individuals, usually battered old males, are not uncom-
mon, the mass of them feel uncomfortable if alone. A herd
is harder to stalk than a single individual. Evidently each
animal feels a sense of companionship and guardianship
when with others, even though of a different species. But
of course the companionship between herds of two differ-
ent species is always intermittent, even although habitual.
They may graze together toward water, or even lie down
together; but after a time they are almost certain to sepa-
rate, whether while grazing or when frightened. Neverthe-
less, occasionally animals of different species, usually friend-
less outcasts, but not always so, will become permanent
companions.

Hartebeest may be found in herds of a hundred indi-
viduals or over or in parties of but half a dozen. Like all
other game of the bare plains, they make no effort to escape
observation, their one aim being to keep so sharp a lookout
that they may themselves see their foes while at a distance.
This is much more easy for them than for the forest-dwellers;
and because of this fact, and because of the abundance of
food for the grazers, as distinguished from the browsers of
the woodland, the dwellers on the bare grass-land are usu-
ally far more numerous, area for area, than the dwellers in
the great forests—and this is true in Africa as in America.

380 AFRICAN GAME ANIMALS

The bison and the barren-ground caribou were always far
more plentiful than the moose, just as the hartebeest and
zebra are far more abundant than the bongo. Unquestion-
ably the plains-dwellers in the absence of man can hold their
own better against brute foes, and thrive better, than the
forest-dwellers; and this continues to be true in the presence
of scattered savages in a low stage of development; but the
conditions are reversed as soon as industrial man appears,
especially when he is a rifle-bearer, and then the plains-
dwellers vanish from the land long before the beasts of the
dense woodland have grown scarce.

In many parts of East Africa the grass is so good and
the water so plentiful that the hartebeest stay perma-
nently in one neighborhood all the year round, and, more-
over, each herd or group in that neighborhood may havea
special beat not more than a couple of miles or so in diam-
eter. Ordinarily the local range is somewhat larger, but
not much, for when circumstances are favorable it is aston-
ishing to see how limited the home range of such an animal
is. If there are seasonal droughts, however, the harte-
beest may all, or nearly all, shift in a body to new pas-
turage; but when we were there there did not seem to be
such common and marked migrations among them as
among the wildebeest and oryx. If the land is very dry
and the pasturage poor the size of the home range and
the likelihood of wandering for each herd or individual
are greatly increased. In very dry wastes the wandering
may be continuous.

It is much easier to watch and study the habits of the
game of the bare plains than those of forest or bush country.
At many of our longer camps we grew to know various

WILDEBEEST AND HARTEBEEST 381

individuals and herds of the common plains game, and we
sometimes got a fair idea of the ordinary round of their
lives. It is a simple enough round, as a rule, and yet
there are extraordinary, indeed inexplicable, variations
in it under certain conditions. By far the most extraor-
dinary of these is one which concerns not merely harte-
beest, but many other African animals, including espe-
cially giraffes, eland, and oryx; namely, the power among
certain animals of all these species, in certain localities,
apparently to go not merely for days but for weeks without
drinking. In certain districts of East Africa we found
the hartebeest herds drinking at irregular intervals, but
at least twice every twenty-four hours; sometimes in the
morning or afternoon, that is, in broad sunlight, and some-
times at night. In other places they drank but once, and in
one or two places, as well as we could tell, only about every
other day. But there are regions in Africa where it seems
certain that hartebeest never drink at all for weeks at a
time, the herds being found where there is absolutely not
a drop of surface water. This we have never, personally,
seen; but on the Northern Guaso Nyiro we watched herds of
reticulated giraffe which, we are reasonably certain, did not
drink for several days, perhaps a week, at a time. Their
companions, the oryx and the big gazelle, were drinking
regularly. Of course, it is possible in this case that the
giraffe were at this time drinking at some water of which
none of the few natives knew, but we do not believe it.
At any rate, both in the Kalahari Desert and in Somaliland,
as well as in certain other tracts, there are seasons of the
year in which various animals—giraffe, oryx, eland, harte-
beest, gazelle—apparently exist without water for periods

382 AFRICAN GAME ANIMALS

so long as to seem incredible. In places, we suppose, they
dig up bulbs and melons, for it seems impossible that such
huge beasts, in the dry heat of that climate, can get on
absolutely without water or plants containing moisture,
as many smaller desert animals, such as jerboas, unques-
tionably do. At any rate, this is a matter of extraordinary
interest, which should be studied on the ground by com-
petent naturalists.

The swarming hartebeest of different species which
we studied, the herds, mixed and unmixed, the families,
the solitary bulls, lived under conditions which offered no
such enigmas to the inquirer; they were where they could
always get water; they were stationary, for we never even
came on them migrating. They lived on or near the
equator, and the seasonal changes were evidently not
sufficient to give them any one breeding-time. In different
places we found calves of different ages; and in other places,
notably on the Kapiti and Athi Plains, where the animals
were extraordinarily abundant, we found calves of all
ages, so that evidently there was no fixed time for the rut
or for breeding. Often we found large herds of bulls,
cows, and calves; often we found small parties of cows
with but a single bull, and other small parties of bulls, or
single bulls, which had evidently been unable to appro-
priate any cows. The bulls waged fierce battles among
themselves. Like the wildebeest bulls, they often fell on
their knees opposite each other before springing together,
butting and stabbing with their short horns. Their horns
do not seem as formidable as those of many other ante-
lopes; but the only case we actually came across in which
any antelope was killed in one of these duels was that of a

eae

WILDEBEEST AND HARTEBEEST 383

hartebeest. Evidently his opponent had stabbed him in
the chest, hooking him with an upward motion as they
faced each other, and, when he turned, had ripped open his
flank. Rather curiously, the only injury actually inflicted
on any man or horse of the Buffalo Jones party, in roping
lion, boar, rhinoceros, giraffe, zebra, eland, and other ante-
lope, was by a hartebeest; after it was roped the cowboys
were careless with it, thinking it harmless, and with a
sudden rush it stabbed the horse of one of them. They
doctored the wound and the horse recovered. These cow-
boys, by the way, rode down and roped the hartebeest in
fair chase. This is not a feat to which ordinary African
hunting-ponies are equal, but Buffalo Jones and his cow-
boys, Messrs. Means and Loveless, were mounted on the
best type of Western American cow-horses, big, blooded
beasts of great speed, courage, endurance, and sure-footed-
ness. They could overtake any animal in Africa. In
our experience we found the topi, hartebeest, and wilde-
beest (we did not chase the gazelles) the fleetest animals
of the plains; on our horses we could not overtake them
when they were in small parties or single, but when in
herds they interfered with one another and we could get
close enough for a shot. Hartebeest and topi were the
fastest. They were very agile, too, and would bound
easily over one another or over low ant-hills. Two or
three of our horses could overtake zebra, and eland were
not difficult to ride down. Once we crouched among some
thin, low thorn-bushes and watched long files of zebra and
hartebeest canter by within a few score yards; we were
interested and surprised to see that the hartebeest fre-
quently cantered with their mouths open, which we had

384 AFRICAN GAME ANIMALS

supposed was a proof of exhaustion, whereas they were all
entirely fresh.

Often we saw a bull hartebeest practising for a fight,
so to speak; he would get down on his knees on the bare
plain, or against the side of one of the huge ant-hills, and
butt and horn the ground, so that his face and horns and
knees would be covered with earth. Sometimes a whole
herd would stroll toward a bare bit of soil which they
evidently used as a stamping-ground, and on reaching it
would stamp and kick up the dust and roll in it; the spot
might be level or it might be the side of an ant-hill. Such
a place would be trodden into bare, pulverized dust by the
sharp hoofs. Zebra also came to these places to roll. Gen-
erally the hartebeest scattered their droppings anywhere,
like most other game; but several times we came on places
—one such area was thirty feet across—in which many
hartebeest had dropped their dung for quite a long period
of time, evidently resorting thither for that purpose. The
Grant gazelle sometimes showed the same habit, and it
seemed to be usual with the little dikdik. The herds of
hartebeest sometimes visit the salt or mineral licks like
the other game; in such places we have seen acres of ground
covered with gullies and hollows, licked out by the tongues
of countless generations of wild creatures. Like most
game, they are tormented by hosts of insects—biting flies,
loathsome bot-flies, and ticks. The ticks swarm in unbe-
lievable numbers on the Kapiti and Athi Plains; elsewhere
they are only ordinary pests. They weaken and even kill
horses; but, curiously enough, the game seem to thrive in
spite of them. We have seen rhinos, wildebeest, zebra,
and hartebeest with their groins, their armpits, the bare

WILDEBEEST AND HARTEBEEST 385

spaces under the tails, the neighborhoods of the ears (and
inside them), and spectacle-like rims around the eyes cov-
ered with thick masses of swollen ticks, like dark barnacles;
yet the animals would be fat and in good condition. The
biting flies and bot-flies annoy them much more. But
tney like. to scratch their hides. We have seen hard,
stubby trunks, and even corners of isolated rocks, worn
to a polished smoothness by the myriads of game that for
scores and perhaps hundreds of years had used them as
rubbing-posts.

Hartebeest are grazers, not browsers. Like most other
game, they draw sharp distinctions between different
kinds of pasturage. Great areas of long, thick-growing
grass will be left untouched, probably because it is sour
pasturage, while adjoining areas with different grass will
be grazed bare by innumerable wild herds. Hartebeest
are rather silent animals; they utter a kind of gasp or
sneeze as a note of curiosity, warning, or alarm. They are
both wary and silly, and the same animals, under the
same conditions, will at different times show, now the
utmost wildness, now a foolish tameness. Their high
withers and low quarters, and their long, homely heads,
carried at right angles to the neck, and therefore almost
upright, give them an awkward look; but when at speed
their even, regular, powerful action changes their whole
look. If pursued by a single dog, for which they care little,
they will play in front of him, bounding and buck-jumping.
One of their marked habits is to stand on ant-hills, espe-
cially if they have the slightest suspicion of threatened
danger. When herds are grazing or resting, whether lying
down or standing up, single animals will often be seen

386 AFRICAN GAME ANIMALS

standing motionless and alert on a neighboring ant-hill.
Such animals are often spoken of as sentinels, as delib-
erately performing, and set to perform, sentinel duty for
the herd. Much study of the herds left us in doubt as to
whether this was or was not the case; as to whether there
was more in the habit than the individual suspiciousness
and wariness of an animal concerned only with his safety
and not thinking about his companions at all. Often we
saw herds where there were one or more individuals on the
lookout, which therefore in effect acted as sentinels; but
more often the herds were grazing, resting, or lying down
without any sentinels. Moreover, if the seeming sentinel
lay down or began grazing there was normally no attention
paid by the rest of the herd and no attempt made to sup-
ply its place by another sentinel. Yet once, when the
possible sentinel descended from his post on an ant-hill, we
certainly saw another take his place; but this one may
merely have been rendered uneasy by his companion’s
movements and have risen to look round on his own ac-
count; for when, after half an hour’s motionless watching,
the beast became satisfied, descended and began to graze,
no other animal took his place or paid heed to what had
happened. When startled or puzzled the lookout animal
might give the kind of sneeze spoken of above. With both
hartebeest and wildebeest we have seen single individuals
watching, on the grazing-grounds and at the watering-
places, under circumstances that made it seem as if they
were deliberately acting as sentinels for the herd; but we
never were sure that this was the case or that the an-
imal was behaving otherwise than it would have behaved
if entirely alone. Indeed, we are inclined to think that the

WILDEBEEST AND HARTEBEEST 387

herd is protected purely by the spontaneous individual
vigilance of its members, each exercising the vigilance on
its own account and for its own safety; but of course fuller
observation may show that this is not the case, and that
different animals do take up in succession the function of
sentinels for the guardianship of their fellows.

As we have already said, there are the widest differences
of conduct among individuals of all the species of game,
and this is true, among other things, of their wariness, both
as between all the individuals of one locality and all those
of another, as among individuals of the same locality, and
as between an individual at one time and at another, even
on the same day. We have passed by herds of game which
Iet us come within easy range of them, although they were
to leeward of us, without moving; and on another day,
or even on the same day, have had them flee in panic and
terror at sight of us a quarter of a mile off, or on getting
the wind of us at a like distance. In most of the portions
of Africa through which we hunted, the beasts of prey ex-
acted a much heavier toll of life from the game than did
the white hunters; and they were in far greater and more
continuous dread of the lion than of the rifle-bearer. In
other words, in most places the game lived under substan-
tially natural conditions; there were only a few species
which had materially diminished in numbers over a con-
siderable area because of the incoming of the white man or
which, because of his presence, had materially changed their
habits. The hartebeest and zebra, for instance, were just as
they always had been. They as often from folly blundered
into the power of the lion, as into the power of the hunter.
In the daytime they often showed the same curious disre-

388 AFRICAN GAME ANIMALS

gard of both lion and hunter and, trusting in their speed
and agility—they knew that the lion, unlike the human
hunter, was not dangerous at any distance—we have seen
a wildebeest staring unmoved at a lion and lioness at a
distance of sixty yards, although they obviously had de-
signs on him, and we saw a party of hartebeest galloping
along not far from, and paying no heed to, a lion we were
pursuing. But they were always on the watch, for even in
daytime the lion, leopard, cheetah, and hunting hound
sometimes assailed them, and at night they were, and
knew that they were, ever in danger, and were subject to
continual panics; one of the stock grievances of the set-
tlers is that in their continual wild stampedes at night the
zebra and hartebeest are always galloping through the wire
fences. Speaking generally, and having in view the numer-
ous qualifications above dwelt on, the wildebeest were the
shyest, and the little Tommy gazelles the tamest, of the
common plains game. The hartebeest were less shy than
the wildebeest, and when a mixed herd came to water the
hartebeest usually came first. We have seen a single
hartebeest lead a herd of wildebeest, whereas if the
mixed herd was composed of hartebeest and zebra either
might lead. A herd of unmixed wildebeest was generally
much shyer and more wary, both when grazing or rest-
ing and when going to water, than a herd of hartebeest
or zebra.

In several different places we were able to study harte-
beest herds for some days at a time and get an accurate
idea of the routine of their daily lives. The striking facts are,
first, the simplicity and monotony of their lives, even when
in danger, and second, the amount of time spent in eating.

WILDEBEEST AND HARTEBEEST 389

Certainly three-fourths or four-fifths of the whole time
was spent in grazing or chewing the cud. Most of the
remaining time the animals were sleeping, either lying
down or standing up. There were no stated times for sleep-
ing or resting and grazing. Usually during the heat of the
day the herds would lie down, sometimes half the individuals
lying down and the others standing up; or they might stand
or lie under the thin half-shade of almost leafless, wizened
little thorn-trees. One or two, or more, might be keeping
a lookout. Yet more than once we have seen herds feeding
at high noon, especially if the day was overcast. On one
occasion a herd we had watched grazing all the afternoon,
and which we expected to see then go to water, deliberately
lay down toward nightfall, and were still lying down when
it grew too dark to see them. Probably after feeding and
moving around for a few hours the herd rests or lies down,
whether by night or day. Hartebeests may come to water
in the night-time, or at dawn or in the late evening, or in
the sunlight of the forenoon or afternoon. Ordinarily a
typical herd consists of bulls, cows, and calves of all ages.
The bulls may fight among themselves; the cow about to
calve may go off alone to drop her calf, and not rejoin the
herd for a few days; and a bull wishing to work off its
rutting rage in a safe fashion will plough along on its knees
or almost stand on its head and dig the bare earth with its
horns. There are continual false alarms, and at the mo-
ment of attack by a beast of prey the panic and terror of
the stampeding creatures reduce them to madness. But
these emotions are as short-lived as they are violent, and
soon the run becomes a canter; the herd gathers and begins
to graze without further thought of the one that has been

390 AFRICAN GAME ANIMALS

slain. Sometimes, although not often, they blunder down
wind to a drinking-pool with reedy margin; but if they have
reason to suspect that their chief enemy, the lion, may be
in the neighborhood, they are very alert, travel to and fro,
and make many halts, and often break back at a run, as
they venture near the water.

Key To THE SPEcIEs OF Bubalis

Horns wide-spread and bracket-shaped, turning at right angles to the
horn pedicle; horn pedicle short; body size smaller
coker

Horns narrowly V-shaped and diverging gradually from their base;
horn pedicle very long, extending far behind condyles
of skull; body size larger ~ lelwel

CoKkE HarTEBEEST
Bubalis coket

The Coke hartebeest is easily recognized by its wide-
spread, bracket-shaped horns and light-buffy or tawny col-
oration. It is a considerably smaller species than lelwel and
has a much shorter head with very differently shaped horns.
In fact, the two species stand at opposite ends of the harte-
beest evolutionary tree, coke1, with its short head and: wide-
spread horns, being one of the least specialized of the genus.
The horns in the Coke hartebeest project backward, in
line with the profile of the top of the head, but spread out
sharply laterally at right angles and then turn backward
again, the points usually extending parallel from the last
turn. The Coke hartebeest occupies the territory east of the
Nile watershed from central German East Africa northward
to the Tana River and Lake Baringo. An isolated race is
found in the northeast drainage area of Lake Rudolf on
the Abyssinian border. Four geographical races are recog-
nizable by difference in horn shape, but show no marked color
distinctions.

nly

WILDEBEEST AND HARTEBEEST 391

Key To Races oF cokei

Horns broadly bracket-shaped and turned sharply outward at right

angles
Dorsal coloration darker or more tawny-rufous; body size and
horns smaller coker

Dorsal coloration lighter or more buffy; body size and horns
larger kongont

Horns without a sharp right-angle turn outward, bowed out regularly
Horns wide-spread but not angulated or bracket-shaped; col-
oration darker and more tawny neumannt

Horns narrower or more V-shaped in direction; coloration lighter
and more buffy nakure

Coast HARTEBEEST
Bubalts cokei coket

Native Names: Swahili, kongon1; Taita, nose.
Alcelaphus cokeit Giinther, 1884, Ann. &F Mag. Nat. Hist., XIV, p. 426; fig. of
horns.

Rance.—Coast district of British East Africa from the
Tana River southward to Kilimanjaro and central German
East Africa, west as far as the eastern edge of the highland
plateau; absent from the moist coast belt.

The typical race was described by Doctor Gunther in
1884 from a head shot by Colonel Coke in Usagara, German
East Africa, in 1880. ‘This was the first complete specimen
of this very abundant species to reach Europe. Von der
Decken had previously, in 1862, collected the horns near
Kilimanjaro. Speke and Grant must have met with this
hartebeest, but their accounts are so vague that they may
refer to the Lichtenstein hartebeest rather than this species.

The typical Coke hartebeest is a smaller and more red-
dish animal than the highland race named kongont. It is
indistinguishable in horn shape from the latter, but the
skull is decidedly smaller, averaging an inch less in length.
The dorsal color is usually darker or more reddish, espe-
cially the crown and the snout, which are cinnamon-rufous

392 AFRICAN GAME ANIMALS

or hazel. In central German East Africa, south of the
Pangani River, the Coke hartebeest meets the range of the
very distinct Lichtenstein hartebeest, but the two species
are not known to be associated together in the same local-
ities.

Flesh measurements of specimens are not available.
The largest male specimen collected by Doctor Abbott at
Taveta measured 1758 inches in skull length, 1614 inches
in length of horns on the outside curve, and 13 inches in
spread at the tips. The individual variation, however, is
great, both in horn shape and in size of skull or body size,
so that the recognition of the race must be based on average
differences. On the plains at the base of Kilimanjaro they
are very abundant, but in the low deserts of Taru and Taita
they are far from common. Hunter has reported them as
far north as the Tana River.

Konconi HarTEBEEST
Bubalis coket kongont

Native Names: Swahili, kongoni; Kikuyu, ngonde; Masai, olsorikor.
Bubalis cokei kongoni Heller, 1912, Smith. Misc. Coll., vol. 60, No. 8, p. 5.

RanceE.—Highlands of British East Africa from the
slopes of Mount Kenia and Lake Naivasha southward and
westward to the southern shores of the Victoria Nyanza
and northwestern German East Africa.

The hartebeest inhabiting the Loita Plains district of
British East Africa was described as a new race under the
name fongoni in 1912 from a specimen shot by Colonel
Roosevelt. Sportsmen have generally considered the harte-
beest of the highlands as indistinguishable from cokei of
the coast lowlands owing to the great individual variation
in horn shape and size of head in this group of antelopes.
The kongoni, however, is decidedly a larger animal with
lighter dorsal coloration. The coloration is more purely
buffy, with very little of the reddish suffusion except on the
crown and snout, which is tawny, very seldom being as
dark as the cinnamon-rufous of the typical cokez.

The dorsal coloration is uniform tawny-ochraceous, be-
coming on the lower sides, the rump, and hind quarters

WILDEBEEST AND HARTEBEEST 393

somewhat lighter, clear buff in color, giving the rump an
indefinite lighter coloration. The forelegs in front are
tawny-ochraceous, like the body, but behind and on the inside
they are buff. The hind legs are buff, with the exception
of a darker stripe in front from the hocks to the hoofs, which
is tawny-ochraceous. The clefts of the hoofs in front are
often bordered by blackish hair. The basal one-half of
the tail is short-haired and buff, like the rump, the terminal
half being black-haired and tufted. The breast and belly
are buffy, like the rump, but the groins and axille are quite
whitish. ‘The crown of the head and the snout are tawny
and distinctly darker than the back. ‘The sides of the head
and throat are buffy-ochraceous, from which the chin is set
off by its black or dark seal-brown color. The ears are
ochraceous on the back and white on the inside and at the
base.

A male specimen from the Loita Plains measured in the
flesh, in length of head and body, 72 inches; length of tail,
22 inches; hind foot, 20% inches; ear, 8 inches. Females
are somewhat smaller than these dimensions, an average
specimen measuring 67 inches in length of head and body;
fais To, imches;) hind foot, 19 inches: ear, 734 inches.
Skulls from the Loita Plains average in length 1734 inches
in males. The largest specimen in a series of fifty from
British East Africa measures 181% inches. The horns vary
greatly in length and spread. ‘The longest male horns in
the large series in the National Museum are 1734 inches
long, the widest being 1758 inches at the tips. Ward
records a large number of specimens from British East
Africa exceeding these dimensions. His record in length
is 21 inches and in spread 1g inches. These dimensions,
however, represent freak or abnormal specimens. Average
specimens measure 16 inches in length by 14 inches in spread.
As usual, among antelopes, the record horns all belong to
young adult animals in which the points have not yet been
worn down. ‘The master herd bulls usually have the horns
very much worn at the tips and greatly shortened. The
horns of the females are much more slender than those of
the males and are shorter.

A large series, some seventy specimens, have been ex-
amined in the National Museum. ‘This material shows

394 AFRICAN GAME ANIMALS

great individual variation in horns and skull size, but
scarcely none in coloration. ‘The specimens from the Athi
Plains or the region east of Nairobi are more or less inter-
mediate between typical kongoni of the Loita Plains and
cokei of the Kilimanjaro region in size of skull. They
average considerably less than Loita Plains specimens,
although the extremes of the two series are quite equal in
dimensions.

Nakuru HartTEBEEST

Bubalis cokei nakure

Bubalis nakure Heller, 1912, Smith. Misc. Coll., vol. 60, No. 8, p. 6.

Rance.—Rift Valley of British East Africa ranging
from Lake Nakuru northward to the south shore of
Lake Baringo and eastward to the base of the Aberdare
Range.

The hartebeests inhabiting the district near Lake
Nakuru have long been recognized by sportsmen as dif-
ferent from the common kongoni owing to their less wide-
spread horns. They have usually been referred to as
hybrids between the kongoni and the Jackson hartebeest
or as representing the Neumann hartebeest described by
Rothschild from Lake Rudolf. Their recognition as hy-
brids is not consistent with what is known regarding the
habits of the species concerned or, for that matter, of the
breeding habits of mammals generally. Hybrids among
mammals are of very rare occurrence and are produced
only under very unusual or abnormal circumstances, and
never in great numbers, at certain points in the ranges of
distinct species. The kongoni and Jackson hartebeests are
nowhere known to be associated together in the same herd,
although their ranges in the Rift Valley are contiguous.
The Nakuru hartebeest resembles closely in characters the
kongoni and is as widely separated from the Jackson harte-
beest as is that form.

The Nakuru hartebeest is a geographical race of the
Coke hartebeest occupying the northwest corner of the |
range of the species. It is most closely allied to the kon-
goni, which it resembles closely in size of body and color-

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MAP 9—DISTRIBUTION OF THE RACES OF THE COKE HARTEBEEST

1 Bubalis coket coket 2 Bubalis coket kongoni 3 Bubalis coket nakure
4 Bubalis cokei neumanni

395

396 AFRICAN GAME ANIMALS

ation, but may be distinguished by its less widely spread
horns, which lack the angular bracket shape of that species.
In horn shape it approaches closely the Neumann harte-
beest of Lake Rudolf, from which if differs by lighter color-
ation and narrower horns. In the large series of specimens
of kongoni from the Loita Plains, in the National Mu-
seum, are one or two which closely approximate nakure in
horn shape, and these are connected by intermediate spec-
imens with the normal bracket-shaped horns of the typical
face:

No flesh measurements are available for comparison.
Three specimens shot near Lake Nakuru by Kermit Roose-
velt represent the race in the National Museum. One of
these has been selected as the type of the race. Two other
specimens from Lake Baringo have been examined in the
British Museum. The larger of the two males shot by
Kermit Roosevelt has a skull length of 1814 inches and
horns of the same length, but with only 8 inches spread at
the tips. The narrowness at the tips is due to the inward
curvature, the horns being widest about the middle of their
length. Herds of hartebeest have been seen on several
occasions on the Naivasha Plains, flanking the Aberdare
Range on the west, which doubtless belong to this race, as
they have from this point northward a continuous. range to
Lake Baringo.

NEUMANN HarTEBEEST

Bubalis coket neumannt

Bubalis neumanni Rothschild, 1897, Ann. &F Mag. Nat. Hist., vol. XX, p. 376.

Rance.—From the northeast shore of Lake Rudolf and
the east bank of the Omo River eastward to Lake Stefanie
and its tributary streams.

A. H. Neumann, while elephant hunting at the extreme
north end of Lake Rudolf near the village of Bumi, shot
the type specimen of the hartebeest which Rothschild
named for him in 1897. Neumann saw them only at this
particular locality. A few years later Donaldson Smith
met with this hartebeest, which he refers to as Coke’s, north
of Lake Stefanie and again on the east bank of the Omo

WILDEBEEST AND HARTEBEEST 397

River, but states that on crossing the river no more were
seen, their place being taken by the /elwel. The Neumann
hartebeest is widely separated geographically from the
British East African races of the Coke, which it closely
resembles. Geographically it is much more intimately con-
nected with the tora hartebeest, to which its horns have
considerable resemblance.

The Neumann hartebeest is known only by the speci-
mens collected by Neumann on the shores of Lake Rudolf.
The type is in Rothschild’s Museum at Tring, and another,
a female specimen, is in the British Museum. These spec-
imens have been examined. They differ from the Nakuru
hartebeest by their more wide-spread horns and darker
dorsal coloration, but their differences are remarkably
slight considering the widely isolated character of their
habitat. The horn measurements of the two known speci-
mens are: male, length, 1614 inches; spread at tips, 81%
inches; female, length, 1344 inches; spread at tips, 97%
inches.

LELWEL HARTEBEEST
Bubalis lelwel

The lelwel represents the extreme type of hartebeest
development in length of head and narrowness of horn
spread. ‘The horn pedicle in this species attains its greatest
development and projects far behind the condyles of the
skull. In body size the lelwel exceeds the other species.
The species represent the maximum in grotesqueness and
size, compared with the others of the genus. ‘The color-
ation is quite uniform and reddish in tint, although there
is in some of the races an approach to the black legs and
face blaze and dorsal stripe of the Caama hartebeest of
South Africa. This latter species, although widely iso-
lated geographically, is a closely related species in horn and
skull shape. The lelwel is primarily a Nilotic species
which has spread eastward in British East Africa, where it
is now contending in the Rift Valley with the Coke harte-
beest for territory.

398 AFRICAN GAME ANIMALS

Key To THE Races oF lelwel

Black stripe on front of legs
Black dorsal stripe or face markings wanting lelwel

A black dorsal stripe on nape and withers; head with a black blaze
or black cheek patches Insti gnis

Black markings on legs or body wanting
Horns narrow, with long pedicle
Dorsal color darker, cinnamon-rufous; head shorter
- gacksoni

Dorsal color lighter, ochraceous-tawny, head longer
rooseveltt

Horns wider-spread, with shorter pedicle kenieé

Heuciin LELWEL HARTEBEEST
Bubalis lelwel lelwel

Native Names: Djeng, Jelwel; Monbuttu, nakibbee; Niam-niam, songaroh.
Acronotus lelwel Heuglin, 1877, Reise, N. Ost. Afr., Il, p. 124, fig. horns.

Rance.—West side of the Nile from the Bahr-el-Ghazal
drainage area south to the Albert Nyanza.

Von Heuglin met with this hartebeest during his explo-
rations of the upper Nile in 1863-4. He described it in
1877, giving it the name by which it is known to the Djeng
tribe of the Bahr-el-Ghazal district. He published with
his description a figure of the horns, which he compared to
the closely allied Caama of South Africa. A few years
later Schweinfurth met with the same species on the head-
waters of the Bahr-el-Ghazal drainage and records its occur-
rence, together with a list of its native names. The lelwel
has remained in obscurity for many years, but recently it
has been re-established by Lydekker.

The Heuglin lelwel is distinguishable from the Roose-
velt lelwel of the east side of the Nile by the presence of
black stripes on the front of the legs and by its somewhat
smaller size and slightly lighter color. A series of five
specimens shot by Colonel Roosevelt in the Lado Enclave
near Rhino Camp are in the National Museum. The flesh

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-WILDEBEEST AND HARTEBEEST 399

measurements of the longest-horned male in this series
were: length of head and body, 76 inches; tail, 26 inches;
hind foot, 22 inches; ear, 7% inches. The horns are 20%
inches along the curve and 856 inches in spread at the
tips. The longest-skulled male in this series measures 2034
inches in greatest length, which is considerably less than the
longest from the east side of the Nile, but exceeds that of
any specimen of jacksoni or insignis.

RoosEvELT LELWEL HARTEBEEST
Bubalis lelwel roosevelti

Native Name: Bari, Joba.
Bubalis lelwel rooseveltt Heller, 1912, Smith. Misc. Coll., vol. 60, No. 8, p. 7.

RancE.—The lowlands bordering the east side of the
Nile from the Albert Nyanza northward to the Sobat
drainage.

A specimen shot in the vicinity of Nimule on the east
side of the Nile by Colonel Roosevelt has been selected
asthe type of this race. It is a large race, the length of
skull exceeding that of any other lelwel. It may be dis-
tinguished from the typical lelwel by the absence of black
on the legs and from jacksoni by its decidedly larger size
and lighter coloration. From niedecki of the upper Sobat
drainage, which also lacks the black leg stripes, it is dis-
tinguishable by its lighter color, distinctly larger size, and
by its longer horn pedicle.

The dorsal coloration is a uniform ochraceous-tawny
which fades gradually into ochraceous-buff on the rump,
hind quarters, legs, and under-parts. The front of the legs,
however, are a bright tawny, which deepens on the front of
the pasterns and the clefts of the hoofs to a brownish-black.
The tail is furnished with a long tuft of black hair at the tip.
The head is uniform in color with the exception of the chin,
which is deep seal-brown in color.

A male shot by Heller at Gondokoro measured in the
flesh 75 inches in length; tail, 24 inches; hind foot, 21
inches; ear, 734 inches. The longest pair of horns are

those of an immature male from the same locality meas-

uring 241% inches in length with a spread at the tips of 12

400 AFRICAN GAME ANIMALS

inches. This is the longest pair of horns of any lelwel in
the National Museum and falls but one inch short of Ward’s
record from the Nile. The horns of the older or fully
adult males are all shorter, owing to the wearing down of
the points. This race will no doubt be found to carry the
longest horns of any lelwel in conformity with its larger
body size. The longest-skulled specimen from Gondokoro
measures 2178 inches, exceeding by more than an inch
that of any other race in the large series in the National
Museum.

Ucanpa LELWEL HarTEBEEST
Bubalts lelwel insignis

Native Names: Baganda, enangazi ; Unyoro, enas.
Bubalis jacksoni insignis Thomas, 1904, Proc. Zool. Soc., p. 455.

Rance.—From the hill country of Karagwe in German
East Africa northward over the plateau region of Uganda
as far as the highlands east of Nimule.

We also owe to Jackson’s efforts the discovery of this
hartebeest. It was named by Thomas in 1904 from a
specimen secured near the Maanja River, in central Uganda.
It is one of the rarest and least known of hartebeests owing
to the extremely local character of its distribution. It has
been reported from Buddu by Gedge, from Karagwe by
Scott-Eliot and from the Mount Agora region of northern
Uganda by Powell-Cotton, but only the latter of these
records is represented by museum specimens.

The Uganda lelwel is characterized by having more
extensive black markings than any other race. Besides
having the black leg-bands it has a black dorsal stripe on
the nape and back and also a black face blaze or black cheek
patches. The dorsal stripe and the black face markings
are sometimes absent or only faintly indicated. In the
general body color and size it is identical with the Jackson
hartebeest.

Four specimens were collected by the Roosevelt expe-
dition in the Maanja River drainage near the village of
Kisingo. The flesh measurements of one of these speci-
mens, a male, were: head and body, 73 inches; tail, 21
inches; hind foot, 2114 inches; ear, 734 inches. The skull

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MAP 10—DISTRIBUTION OF THE RACES OF THE LELWEL HARTEBEEST
3 Bubalis lelwel niediecki

1 Bubalts lelwel lelwel 2 Bubalis lelwel rooseveltt
5 Bubalis lelwel jacksoni 6 Bubalis lelwel kenia@

4 Bubalis lelwel insignis
401

:

402 AFRICAN GAME ANIMALS

of this specimen measures 20 inches in length, the horns
21% inches in length with a spread of 9% inches at the tips.

Jackson LELwWEL HarTEBEEST

Bubalis lelwel jacksont

Native Names: Nandi, chemnyokoso; Turkana, etulia.
Bubalis jacksoni Thomas, 1892, Ann. & Mag. Nat. Hist., p. 386.

RanceE.—Plateau region of Mount Elgon and the Mau
Escarpment from the Lumbwa country northward beyond
the headwaters of the Turkwell River.

Jackson discovered the hartebeest which now bears his
name on his initial journey to Uganda in 1889. Until very
recently the name was applied broadly by sportsmen to
all the hartebeests of the lelwel type found in Uganda and
the Nile Valley. The Jackson hartebeest, however, is a
distinct race occupying the highland region of the Mau
and Mount Elgon. It is a darker and richer red than any
other race and somewhat smaller in size than the Nile
races. The color is quite uniform cinnamon-rufous with-
out any black markings on the legs or face. The only
contrasts in its coloration are the black chin and terminal
tuft of the tail and the white hair lining the inner side of
the ears.

A large series of specimens is in the National Musuem,
from the Uasin Gishu Plateau, collected by the Roosevelt
expedition. The flesh measurements of one of the adult
male specimens were: head and body, 72 inches; tail, 2314
inches; hind foot, 2324 inches; ear, 8% inches. An aver-
age male skull measures in length 20 inches, that of a
female 1844 inches. The horn length averages about 22
inches. Great variation in the spread at the tips is shown
in the series, which ranges from 4 to 14 inches in this meas-
urement. Such variation is due to the wide diversity in
direction taken by the horns near their tips. As a rule,
however, the points extend parallel after making the final
turn backward at right angles to the dorsal plane of the
head. Ward’s record horn length for this race is 26 inches.
The longest specimen in the National Museum series is
231% inches in length.

WILDEBEEST AND HARTEBEEST 403

Kenia LELWEL HarTEBEEST
Bubalis lelwel kenie

Bubalis lelwel kenie Heller, 1913, Smith. Misc. Coll., vol. 61, No. 17, p. 3.

RancE.—From the plateau region north of Kenia west-
ward across the Laikipia Plateau to Lake Baringo.

The Kenia lelwel was very recently described from a
specimen shot by Paul J. Rainey on the plateau region
north of Kenia some twenty miles northeast of the station
of Nyeri. The hartebeest inhabiting the north Kenia
Plateau is distinguishable from jacksoni by decidedly wider-
spread horns, shorter horn pedicle, and by somewhat lighter
coloration. From the Coke hartebeest, which it resembles
in shade of coloration, it is at once distinguishable by its
V-shaped horns, longer head, and larger body size. This
race is confined to the plateau region flanking Mount Kenia
on the north and drained by the Northern Guaso Nyiro
River. It represents the extreme western range of the
lelwel type of hartebeest. A. H. Neumann, in his “ Ele-
phant Hunting in East Africa,’ mentions shooting this
type of hartebeest on the southwestern slopes of the Lorogi
Mountains. This locality, no doubt, represents the northern
limit of the range. Westward it connects with typical
gacksoni in the region west of Lake Baringo. It occurs in
very limited numbers, is exceedingly wary, and is seldom
secured by sportsmen.

The type, which is an adult male, measured in the flesh,
in length of head and body along the curve of the back,
78 inches; tail, 2214 inches; hind foot, 2114 inches; ear,
7% inches. The skull has a length of 19% inches. The
horn pedicle has a least width of three-fourths its length from
the orbit to the forking of the horns and is much wider than
in other races of lelwel, in which the width is only two-thirds
the length. The dimensions of the horns in this specimen
are 2114 inches in length by 101% inches in spread at the
tips. Besides the type there is a younger male from the
same district, also collected by the Rainey expedition. This
specimen is very much lighter in color but agrees with the
type in the size and shape of its horns and horn pedicle.

CHAPTER XIV

BUFFALOES

SUBFAMILY Bovine

Tue subfamily Bovine, comprising the oxen, bison, buf-
faloes, and their allies, are the largest or most heavily
built members of the family Bovide. They are char-
acterized by their heavy bodies, short, thick legs and neck,
and large heads. The muzzle is large and broad, the snout
of moderate length, and the face without anteorbital glands.
The horns are broad or rounded but never ringed, and are
well developed in both sexes. The female is furnished with
four mamme. The body varies in outline from the low
withers and well-developed hind quarters of the ox and
buffalo to the high withers and weak hind quarters of the
bison. The tail is long and tufted, the hoofs large, with
well-developed false hoofs, the pelage rather short and
often scanty, and the coloration monocolor without con-
trasting color marks of any sort. The teeth are very much
broader than those of antelopes, with rectangular crowns,
and broad crests or lophs with the valleys between very
shallow or obsolete. The skull is without sinuses between
the facial bones, but has rather long nasal bones, small
anterior nares, and broad expanded tips to the premax-
illary bones. The horn-cores arise laterally behind the
orbits and extend outward and downward or backward but
never upward at an angle to the dorsal profile of the head.

In general shape the skull resembles closely that of the
404

BUFFALOES 405

brindled wildebeest, which is without doubt a near relative
of the group.

The subfamily ranges through temperate North America;
Europe, from Great Britain, within historic times, eastward
to Asia and the Malay Islands, and Africa south of the
Sahara. Members of the group have been found only as
far back as the Pliocene, but were at that time quite as
widely distributed as at present. Notwithstanding the ab-
sence of the Bovine in the older Tertiary geologic forma-
tions, they are not a modern group but rather one whose
ancestral forms are still unknown.

AFRICAN BUFFALOES

Syncerus

Syncerus Hodgson, 1847, Journ. Asiatic Soc., Bengal, XVI, new series No.
7, Pp. 709; type fixed by Hollister, 1911, Bos brachyceros, the Lake Chad buffalo.

A considerable number of distinct generic names are
to-day employed for the African buffalo by various writers.
Some of the names used do not apply, while others are too
comprehensive to be applicable to our present conception
of genera. The old Linnean genus Bos is of the latter
category. It is often used in a broad sense by writers for
all of the Bovine, but should be limited as a generic term
to the ox and its close allies. Bubalus, another name fre-
quently used in a generic sense for the African buffalo, is
properly applicable to the Indian buffalo, as the latter shows
differences in the skull and shape of the horns from the
African which are of generic weight. The name is also
objectionable on account of its close similarity to Bubalis,
the generic designation of the hartebeests. Those who
consider the African buffalo a distinct generic type from
the ox and the Indian buffalo must employ the term Syn-
cerus, of Hodgson, who proposed it in 1847 for the African
buffalo inhabiting the Lake Chad or Bornu district. The
generic character of greatest weight in this genus is the
lack of attachment of the vomer to the palatine bones.

406 AFRICAN GAME ANIMALS

Other characters are the massive horns, which are very
broad basally and cover the whole crown of the head, the
concave profile of the snout, the short, broad nasal bones,
and the broad ears. The genus is confined to Africa south of
the Sahara, where it is of universal distribution from the low
coast lands to the limits of forests on the highest mountain
areas. A single species, caffer, with its several geographical
races, comprise the genus. Three fossil species, very re-
cently extinct, from the Pleistocene age have been described.
One of these from the Orange Free State, baini, had very
much longer horns than the species now living in the same
region, but closely resembled in this respect a species,
antiguus, described from deposits of the same age in Al-
geria. A third species has been described from the Pleis-
tocene of Natal.

Nearly a quarter of a century ago the African buffaloes
were smitten by a terrible scourge, a cattle sickness of such
virulent character that as it traversed the continent, from
north of the equator to south of the Zambesi, it swept the
herds from the face of the earth. Domestic cattle suffered
equally, and a few of the big bovine antelopes suffered
slightly. Of the buffaloes in East Africa and Uganda prob-
ably not one in ten thousand was left alive! It was an ap-
palling calamity. The destruction was far more sudden
than is ever the case when man is the sole agent, and far
more complete for the length of time involved. But there
was a vital distinction. When the disease had spent its
force it vanished, and the scattered survivors were left
free to recover the lost ground. The extraordinary vigor
of natural reproductive power, of wild fecundity when there
is a vacant place for its action, was then shown to the utter-
most. The few remaining buffalo found themselves in pre-
cisely the position of the feral horses and cattle turned loose
by the Spaniards on the grassy plains of America. They

EAST AFRICAN BUFFALO, MALE

Shot by Theodore Roosevelt at Kamiti Farm
Trophy at Sagamore Hill, Oyster Bay, New York

HERD OF EAST AFRICAN BUFFALO

Upper Tana River near Fort Hall, B. E. A.
From a photograph by Carl E. Akeley

EAST AFRICAN BUFFALO

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BUFFALOES 407

had what was for their purposes a nearly vacant continent
to conquer by dint of their reproductive power. Except
where civilized man has been present, and in spite of the
presence of the native foes of the buffalo, this reconquest
has been largely achieved.

This fact shows that this animal, at least, can more than
hold its own as far as its brute enemies are concerned, and,
where the climatic conditions do not forbid, will populate
to near the limit of its food supply. Buffaloes are now com-
mon beasts in East Africa and abundant in Uganda. The
wise policy of the British Government in protecting all the
big game has aided in this recovery.

The buffalo of the White Nile belongs to the Abyssinian
species, or subspecies; while the East African buffalo is
closely allied to the Cape animal. But there is no differ-
ence in habits between the two races, although within the
limits of the same species there are marked differences be-
tween the behavior of the herds in one region and of the
herds in another.

Buffaloes are grazers and are fond of water. They are
not beasts of the desert, and, unlike the giraffe, eland, and
oryx, they cannot exist in nearly waterless regions. They
are gregarious, going in herds of from a score to a couple of
hundred individuals; and in addition old bulls are found
singly or in small parties, while occasionally a herd will con-
sist of nothing but cows, calves, and young stock. Buffaloes
are hardy animals, and are found in varying and very dif-
ferent habitats within the limits of their general range.
Neither the zebra nor any of the antelopes can adapt them-
selves to such varying conditions, provided only that there
is no dearth of water. In places buffaloes live on or near the

408 AFRICAN GAME ANIMALS

open, grassy plains beloved of the zebra and _ hartebeest;
elsewhere they live in dense forests; elsewhere they wander
in the neighborhood of some river running through waterless
flats of grass or thorn-trees; they are found in the high
mountains, where the nights are very cold; and they are
also found in the low-lying, hot regions near the coast.

In their daily habits buffaloes differ both according to the
nature of the country and according to whether they have
or have not been much hunted. In places where they live
in dense forest and are hunted, they venture into the open
only after nightfall; and where much molested they never
feed by day, so that observers have treated them as purely
nocturnal animals. But we are convinced that these ex-
clusively nocturnal habits are not natural tothem. Doubt-
less they everywhere graze as freely, or almost as freely, in
moonlight as in sunlight; and probably the twenty-four
hours are often divided into periods of alternate feeding and
resting, without much regard to light or darkness. But in
many places they feed and rest out in the open during the
day, and in other places they spend the day in thorn jungle
so thin as to afford but scanty cover.

We studied one herd on Heatley’s farm, near the Nairobi
falls. The buffaloes must have numbered over a hundred,
and Heatley had carefully preserved them; he had killed
one or two bulls, and his Boer farmer had shot another in
his garden one night, but there had been so little molesta-
tion that the animals were living practically as if there were
no men in the country. Sometimes the whole number of
animals, or nearly the whole number, went in one big herd.
More often one or two small herds split off from the main
one, and there were also outlying bulls which went singly

BUFFALOES 409

or in small parties. Their home was in a great belt of
papyrus swamp, fifteen miles long, or thereabout, and a
mile broad in places. This papyrus belt was a regular
morass of slime and water. The thick-growing papyrus
stems, with their plumed heads, were twenty or thirty feet
tall. The gloomy depths of the morass served as a secure
refuge for the buffaloes, and they had trodden innumerable
trails hither and thither through it. These trails were mere
lanes of deep mire and water, with the huge stems of papy-
rus crisscrossing over them; only the vast strength of the
beasts, their short, thick legs and brawny bodies, enabled
them to plough their way along them, or at need to shoulder
a passage through the reeds. If buffaloes were not half-
amphibious beasts they could not dwell amid such surround-
ings. While the herd was among these huge reed beds it
was practically safe from pursuit; that is, a keen hunter
would have gone in after them as a matter of course if it
had been impossible to get them otherwise, but the odds
would have been much against the man’s success, and the
danger would have been serious; besides there was no diff-
culty in getting them outside the reeds. Their station
when in the reed beds was usually marked by the attendant
cow-herons. These small white herons accompany the ele-
phant, rhino, and buffalo in flocks, frequently alighting on
their backs. They catch the grasshoppers and other insects
kicked up from the grass by the feet of their hosts. In Heat-
ley’s papyrus swamp the cow-herons evidently found the
dark cover uncongenial. The flock, which accompanied the
herd in the open as familiarly as cow-buntings accompany
cattle in our own pastures, usually perched in a body among
the papyrus tops when the herd was resting near by among

410 AFRICAN GAME ANIMALS

the bottom stems. It was a pretty sight, as the white
plumage glistened in the sun, and it generally enabled us to
know just where the herd was. There was no food for the
buffaloes in the papyrus, and evidently they liked to rest in
the open no less than to graze there. We saw them outside,
on the grassy plains, even half a mile away from the papy-
rus, grazing or lying down, at every hour of the day. We
found them grazing outside the reed beds for a couple of
hours after sunrise and then retiring to their wet and gloomy
fastness. We also saw them come out to graze in the
early afternoon, and return to the swamp shortly before
sunset. We also found them lying down in the open at
about ten in the morning, and up and grazing almost at
high noon. There was evidently no invariable routine;
and of course these buffalo could get water at any time
they wished.

On the banks of the Northern Guaso Nyiro we found that
the buffaloes came down to the river to drink every night.
Evidently they thought the strip of thick tree jungle alongside
the river too narrow to harbor them, and by dawn they were
well on their way back to the dry flats and sparse, parched
thorn jungle which covered the country away from the
river. When we were there the withered thorn-trees gave
almost no shade; and while the buffaloes rested, standing or
lying, the sun shone pitilessly on the black bodies. Follow-
ing the trails from the river, we generally found the animals
resting; but we found one herd feeding quite late in the fore-
noon and another feeding at almost noon. The scent of
the buffalo is as keen as that of the elephant or rhino, and
its sight far better, so that it is much more difficult to stalk.
We never saw anything in the nature of a sentinel; the

BUPFALOES() "7 411

herd trusted to the vigilance and the sharp senses of its
members, individually and collectively.

The Abyssinian buffaloes we encountered were in the
Lado, on the western bank of the Nile. They were living
in country much like that along the Northern Guaso Nyiro,
and their habits were substantially those of their Northern
Guaso Nyiro cousins. At one camp by a native village, we
found a herd living in the dense reed beds, through which
they had trampled a tangle of trails. This herd entirely
realized that they were safe in their.reed fastnesses, and only
came into the open country at night to graze. Yet in the
same neighborhood there were other buffaloes with entirely
different habits. These lived among the dry, scattered
thorn-trees, which, interspersed with a few other trees such
as palms, covered the surrounding country, but nowhere
formed thick cover. There were a few pools at which these
buffaloes drank. They fed and rested alternately through-
out the day and night. We found a bull grazing at mid-
day. They rested, standing or lying down, among the
nearly leafless thorn-trees, which gave scant shelter from
the sun.

Aside from man, the buffalo’s one enemy is the lion. Of
course, a crocodile may occasionally take one, or a calf or
yearling may be killed by wild hounds, but the lion is the
only beast that ever follows the buffalo as an ordinary prey.
There are localities where lions prey on buffaloes almost
solely, just as in some other places they prey almost exclu-
sively on domestic cattle. But where we were the lions
habitually preyed on other game and rarely attacked either
buffaloes or cattle. In the Lado they killed pigs and ante-
lopes; in East Africa, zebras and antelopes. A buffalo is a

412 AFRICAN GAME ANIMALS

tough and dangerous beast, and where zebra and _ harte-
beest abound lions naturally follow the easier quarry. On
Heatley’s farm a family of lions made their day lair in the
big papyrus belt which also harbored the buffalo herd.
Usually neither interfered with the other, the lion finding
ample hunting among the swarming buck of the surround-
ing plains; yet one night two lions killed a buffalo heifer
just outside the papyrus. A single lion, no matter how
large, will rarely, unless very hungry, tackle an unwounded
buffalo bull; when one is killed by lions it is usually by a
party of them, and the assailants do not always escape
scathless, it being no uncommon thing for one of them to be
killed or wounded in such a fight. A big lion will kill a
buffalo cow or young bull without much difficulty. This
is because the lion makes his assault by surprise, and at the
outset gets such an effective hold that the doomed buffalo
has no chance to exert its enormous strength. A cow with
a young calf is so on the alert that she is apt to detect the
approach of her foe; and if she does so she herself makes
the assault, without any hesitation, and may kill or drive
off the lion.

The buffalo is rightly deemed one of the most dangerous
beasts of the chase to be found in the world. In unfre-
quented places, or where it has grown accustomed to dom-
ineer over defenceless natives, it will attack unprovoked.
Near Kenia, while we were there, a cow buffalo regularly ran
amuck through the villages, killing and crippling a number of
persons before the young men slew her with spears. Shortly
after we left Africa, Messrs. McMillan and Selous made a
trip down the Northern Guaso Nyiro, and one of their porters
was charged and mortally hurt by a buffalo. On Heatley’s

nee

tnepige

EAST AFRICAN BUFFALO, MALE, AGE SIX YEARS

_From Central German East Africa
In the National Zoological Park, Washington, D. C.

LIVING SPECIMEN OF THE EAST AFRICAN BUFFALO

BUFFALOES 413

farm passers-by had twice been charged, unprovoked, by old
bulls. But the real danger comes when wounded buffaloes
are followed, especially into thick cover.

Nowadays, in Africa, buffalo have to be killed on foot,
by tracking, or by still-hunting through the country in
which they are found. Their heavy bodies and sharp
hoofs make it comparatively easy for a good tracker to fol-
low them, and although their senses are keen they are easier
to stalk than antelope, being easier to see and just as easy
to approach when seen. They are everywhere less easy to
kill than rhinos. They do not travel such distances as ele-
phants, and hence their chase does not necessitate such
wearing fatigue. The actual circumstances of the stalk
vary completely with the cover and the local habits of the
animals. Beasts that only venture from the forest or thick
jungle at night are, of course, very hard to follow successfully.
In light, open jungle, or where the beasts feed on the plains
near cover in daylight, it is not difficult to bag a buffalo.

Usually there is little danger in the first shot if taken
from a reasonable distance, although even under such cir-
cumstances there is now and then a determined charge.
Following a wounded buffalo is proverbially risky, as we
have already said. Veteran hunters differ widely in their
estimate as to which beast is the most dangerous; the
claims of lion, leopard, elephant, buffalo, and rhinoceros
have each been stoutly defended. -Our own belief, based on
all the evidence, is that when a buffalo bull does turn to
bay it is to the full as formidable as—and probably more
formidable than—a lion, and much more formidable than
an elephant, but that it turns to bay far less freely than
either. Our own slight experience supported this view,

414 AFRICAN GAME ANIMALS

although it was too slight to offer much basis for judgment
by itself. Our party killed about a score of lions, a dozen buf-
faloes, a dozen elephants, a score of rhinos, and half a dozen
leopards. The leopards were the pluckiest and most sav-
age of the lot, although, because of their small size, less for-
midable to life than the other game. It happened that no
buffalo charged us, whereas several lions charged with great
determination, and two or three of the elephants charged
without being molested. Most of the difference was un-
doubtedly due to chance or individual variation. Yet we
cannot believe that lions would have failed to charge if
placed as some of the buffaloes were, although the latter did
not charge. The first buffaloes we attacked were four bulls
grazing outside a papyrus swamp. On our firing at them
they ran, not into the swamp, but into the open. At two
or three hundred yards they halted. One then fell dead;
two others had been wounded. Yet on our walking toward
them as they stood facing us—black, ugly, formidable-
looking—they lost heart and again ran. All four were killed
without charging. Under similar circumstances it is hard to
believe that a lion would not have charged; again and again
we saw lions turn to bay and charge on less provocation.

Key To THE Races oF caffer

Horns curving downward from the base well below the level of the floor
of the skull; horns long and massive, greatest spread

40 inches or more; basal palm of horn very wide

radcliffer

Horns only curving down from the base to the level of the floor of the
skull; horns shorter and narrower across the base,
greatest spread considerably less than 40 inches;
width of palm at base less than Io inches

@quinoctialis

BUFFALOES 415

East AFRIcAN BUFFALO
Syncerus caffer radcliffet

Native Names: Swahili, mbogo or nyati; Masai, olaro; Kavirondo (Jaluo),
jut.
Bubalis caffer radcliffe Thomas, 1904, Proc. Zool. Soc., p. 371.

Rance.—From the Northern Guaso Nyiro River drain-
age and the northern highlands of Uganda southward
through British and German East Africa; east as far as the
coast, and westward to the Edward Nyanza and Lake Kivu.

The buffalo has been known to occur in East Africa
from the dawn of its history. During the early explora-
tion of the country it occurred in countless thousands from
the bush country of the coast to the plateaux of the far
interior, and was much better known than it has been since
its partial extermination by the rinderpest in 1890. Speke
and Grant recorded the buffalo as abundant everywhere
throughout the grass country. In 1904 Thomas described
the East African buffalo as a distinct race from the South
African, applying to it the name radcliffei, and basing his
description upon a specimen collected in Ankole, Uganda, by
Colonel Delme-Radcliffe during his boundary survey of the
German border. Two years later Matschie published a paper
describing several races from East Africa upon trivial differ-
ences in horn shape. The differences which Matschie has
assigned to his races are of an individual character and of
no racial value, and on this account cannot be recognized.
At least seven of his racial or specific names, as he uses them,
apply to the race here considered as radcliffei. The differ-
ences in horn shape in a single herd of buffaloes are reaily
quite wide individually when animals of the same age alone
are compared. The age difference is known by all sports-
men to be much greater, varying from the short, horizontally
directed horns of the young bulls to the great, massive, down-
curved horns of the old bulls. In “East African Game
Trails” Colonel Roosevelt, having in mind Matschie’s paper,
mentions the extensive variation in horn shape he observed
in a herd from which he shot several specimens at Kamiti
Farm, and throws doubt upon such races as Matschie’s,
based upon slight differences in horn shape. Matschie re-

416 AFRICAN GAME ANIMALS

plied to this criticism by naming two more races from this
very spot, representing the two extremes of horn shape to
be found in the herd. He cheerfully agreed with Colonel
Roosevelt that the horn variations he mentioned actually
existed in this herd, but justified the naming of the two addi-
tional races under the assumption that two distinct forms
met at this place and associated together. We, however,
contend that the variations in structure shown by individual
members of a race or species associated together or occupy-
ing the same territory must be taken as representing the
individual variation in that form. It would indeed lead
to great confusion in systematic zoology if the racial char-
acters assigned by the describers were not to be subject
to the test of individual variation as shown by geographi-
cally associated individuals. In fact, there is no other
practical way of determining the individual variation of a
race.

The East African buffalo differs from the typical race of
South Africa by the smooth or flat character of the palm of
the horn base and the larger average size of the horns in
the bull. The horn bases in the South African races are
rugose and roughened by ridges. ‘There is, however, no dif-
ference in body size, in coloration, or in the condition of the
hair covering between the equatorial race and the southern.
Both sexes are uniformly black throughout their pelage,
and the horns and hoofs are also blackish. The hair is
coarse, of fair length, and usually so scanty that the dark-
brownish color of the skin shows everywhere and adds
largely to the color effect of the animal. ‘The tail is fur-
nished with a long tuft of black hair, as in domestic cattle,
and the broad ears are margined by an irregular fringe of
rather long hair. The nursing young are lighter than their
parents, being a uniform dark-rufous color. The horns of
the female are much smaller than those of the male and
are quite different in shape, curving downward only as far
as the level of the condyles of the skull.

The largest-bodied bull in the National Museum was
shot by Colonel Roosevelt at Kamiti Farm. ‘The flesh
measurements of this specimen were: head and body along
contour of back, 108 inches; tail, 3514 inches; hind foot,
2514 inches; ear, 114% inches. ‘The skull of this individual

BUFFALOES 417

measured 1914 inches in basal length by 9% inches in
zygomatic breadth. It was, however, the smallest-horned
individual of the three males shot in this herd, the horns
having a maximum spread of only 40 inches, with a length
along the inside curve of 27 inches. The largest-skulled
specimen in the collection is one shot by Kermit Roosevelt
on the Northern Guaso Nyiro, having a basal length of
20% inches, with a zygomatic breadth of 9% inches. The
horns of a bull shot by Paul J. Rainey near the same
river exceed in spread those of any other examined. The
dimensions of this pair are: maximum spread, 44% inches;
length of horn on inside curve, 34 inches; width of palm,
measured by calipers, straight and not over the curve,
87% inches. The flesh measurements of an adult cow
shot at Kamiti by Colonel Roosevelt are: head and body,
g2 inches; tail, 28 inches; hind foot, 231% inches; ear, 10%
inches. Length of skull, 19 inches; zygomatic width, 834
inches. Horns: maximum spread, 36 inches; length on
inside curve, 23 inches; width of palm, straight, 534 inches.
The record buffalo of all Africa is a representative of this
race and was obtained near the type locality in Ankole by
District Commissioner Knowles. The horn dimensions of
this specimen are: greatest spread, 52% inches; length on
inside curve, 4834 inches; width of palm on curve of face,
1134 inches.

The East African buffalo is found throughout all the
well-watered parts of the country and as far northward in
the desert as the streams or water will allow. The Tana
River marks its northern boundary in the coast drainage.
North of Mount Kenia the buffalo follows down the Northern
Guaso Nyiro far into the desert, and occurs also north of the
river in isolated herds on the forested summits of Mount
Uaragess and the Mathew Range, reaching its northern
desert limits on the peak of Mount Nyiro, at the south end of
Lake Rudolf. From Mount Elgon the range extends north-
ward on the high plateaux as far as the Mount Agora district,
east of Nimule, at which latitude the adjacent lowlands
are occupied by the Nile buffalo. Upon the slopes of Mount
Kenia the buffalo ascends to the limits of the forest and
occasionally strays over the tundra-like moorland of the
alpine zone to the permanent snow-fields. A mummified

418 AFRICAN GAME ANIMALS

carcass lying at the lower edge of one of the small glaciers
at an altitude of 14,500 feet has been reported by several
mountaineering expeditions on Mount Kenia. The skull
of a very large bull is in the National Museum, collected
by Doctor Mearns at the upper limits of the Kenia forest
at 10,000 feet. At the present time, however, the buffalo
is known to occur only in the lower edge of the Kenia
forest, and the two high mountain records, no doubt, repre-
sent strays which wandered far beyond their normal habitat
and perished.

Nite BurFraLo
Syncerus caffer equinoctialis
y q

Native Names: Dinka, anyarr; Aluru, jobi; Bari, makorr.
Bubalus caffer equinoctialis Blyth, 1866, Proc. Zool. Soc., p. 371, fig. 2.

RancEe.—From Lake Kivu northward along the Nile
lowlands to the Bahr el Ghazal and Abyssinia; eastward as
far as Lake Rudolf and the Rift Valley of Abyssinia.

The early explorers and naturalists who met with this
buffalo in Abyssinia and the upper Nile considered it iden-
tical with the South African, owing, no doubt, to its close
similarity in size and coloration. ‘The decided horn differ-
ences were, however, detected in 1866 by Blyth, who named
the race from a specimen collected by Consul Petherick on
the White Nile. Von Heuglin, Schweinfurth, Baker, and
Emin have given full accounts of their experiences with the
buffalo of the upper Nile in the narrative of their journeys.
Lydekker has described, under the name of mathewsi, a
specimen of this race from the region lying between Lake
Kivu and the Edward Nyanza. This specimen marks the
extreme southern limits of the Nile buffalo.

The Nile buffalo is distinguishable from the East African
by its more horizontally directed horns, which never curve
downward below the level of the floor of the skull. This
difference in amount of downward curvature is well shown
by the skull when placed on a level surface, in which position
it rests on the floor of the skull, the horns hanging free of
the surface, an inch or two above it. Skulls of the East
African race placed similarly rest upon the lower curve of
the horns, the floor of the skull being raised several inches

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1 Syncerus caffer radcliffet 2 Syncerus caffer equinoctialis
419

420 AFRICAN GAME ANIMALS

above the surface. The horns are also smaller and less
wide-spread. The very largest only reach 40 inches, an
average horn spread being but 32 inches. In size and shape
they closely resemble the horns of the cow of the East Afri-
can race. A single specimen is in the National Museum,
a bull shot by Colonel Roosevelt near Rhino Camp, in the
Lado Enclave. This specimen is quite equal in body and
skull measurements to the East African ones. ‘The flesh
measurements were: head and body, 110 inches; tail, 28
inches; hind foot, 24 inches; ear, 1034 inches. Basal length
of skull, 1934 inches; zygomatic breadth, 9 inches. Maxi-
mum horn spread, 3134 inches; length along inside curve,
25 inches; width of palm, straight, 84 inches. Specimens
have been recorded by Powell-Cotton in the upper Hawash
Valley and the central plateau region of Abyssinia, near
Lake Tana. Buffalo have been shot and recorded by sports-
men frequently along both banks of the upper Nile, from
Wadelai southward to the Sobat River. A specimen shot by
Paul Niedieck in this district has a horn spread of 44 inches,
which is 3 inches in excess of the previous known record for
the race.

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