THE FAUNA OF BRITISH INDIA,
ISC' HIDING
CEYLON AND BURMA.
Published under the patronage of the
Secretary of State for India.
EDfTBD BV LT.-COL. B. B. S. SGWELf,, C.I.S., Sc.D., P.E.S.. I. M.S. (ret.).
MAMMALIA.-V01. I.
PRIMATES and GARNIVORA (in part),
Families FELID^: and VIVERRIDA2.
R. I. POCOCK, F.R.S.
With 31 plates and 106 text-figures. Illustrations
by the author, unless otherwise stated.
TAYLOR AND FRANCIS, LTD.,
RED LION COURT, FLEET STREET, LONDON, E.G. 4.
March 31, 1939.
PRINTED BY TAYLOR AND FRANCIS, LTD.,
KED LION COTXBT, IXEBT STREET,
CONTENTS.
Preface v
Introduction xv
Explanation of Plates xxvii
Systematic Contents xxsi
Class Mammalia 1
Order I. Primates 13
Order II. Carntvora 186
Alphabetical Index 459
«2
PKEFAOE.
Blanfoed's volume on the Mammals of British India,
published in two parts in 1888 and 1S91, fulfilled satisfactorily
the purpose for which it was compiled, and was for many
years the standard work on the subject. The material at
his disposal*, however, was entirely insufficient according to
modern standards, and the book has admittedly been long out
of date. Various factors, such as general increase in our know-
ledge of Indian mammals and the use of other characters
than those supplied by dried skins and skulls, contributed to
this end ; but perhaps the greatest was the new impetus
given towards the end of the last century to the study of
systematic mammalogy the world over by Hart Merriam's
introduction of his scheme for investigating the mammalian
fauna of the United States by means of intensive collecting
and accurate labelling, with measurements, dates, and altitudes,
of long series of specimens, where available, of every species
from carefully selected localities. This threw a flood of light
upon the variations of individuals of a species from the same
locality and environment with age and season or irrespective
* It is needless to repeat the sources whence he derived his informa-
tion, since they are fully acknowledged in his introduction . Incidentally,
however, he candidly admitted that, on account of the numerous
inaccuracies they contained, he referred as little as possible to the work
of J. E . Gray, who described a very large number of alleged Indian species.
This was a pity, because the inaccuracies called for correction, and Gray,
in spite of his errors, showed now and again unexpected perspicacity
in his determinations, with the result that Blanford, in dismissing his
■opinions, was not infrequently wrong. In those days, too, authors
assumed greater latitude in adopting names they preferred, and were
less strictly adherent to the law of priority.
VJ KREFACE.
of those factors, and also upon the adaptive modifications
a species may locally exhibit when distributed over a wide
geographical area with diversified physical features.
To the older school of systematic mammalogists, who adopted
the Linnsean binominal method of indicating a supposed
species by its generic and specific names, such modifications
were more or less vaguely known, and were given full specific
status if held to be important enough to warrant that rank,
or were cited as " varieties " if falling short of that standard.
But with the new method of studying mammals the term
" variety " was restricted to such individual aberrations from
the normal as are illustrated by melanism or albinism, for
example, and the term "subspecies" was invented to symbolize
the geographical and environmental variants of a species.
At the same time the trinominal system of nomenclature,
now in vogue, was introduced to designate the " subspecies,"
or " local races " as they are often called, into which most
widely distributed species can be divided.
This system is particularly useful in supplying additional
machinery for achieving the main purpose of nomenclature,
namely, the expression of the degree of kinship between
related animals. This may be illustrated by a very simple
instance taken from Blanford's volume. The three best-known
British Indian foxes were cited by Blanford as distinct species,
the hill-fox, the desert-fox, and the Bengal fox. This method
of treating them suggests that the three are equally different
from each other, and entirely conceals the close affinity between
the hill-fox and the desert-fox and the more remote kinship
between them and the Bengal fox. The facts are now expressed
by regarding the first two as subspecies of the so-called common
fox and by citing them as Vulpes vulpes montama and Vulpes
vulpes pusilla respectively, and the Bengal fox as Vulpes
bengalensis.
In theory this system appears simple, but it is by no means
always so in practice. Where a complete series of examples
of a species spread over a wide diversified area is available,
PREFACE. VU
it is the custom to select for subspecific denomination the best
differentiated local races, leaving the equally interesting
intermediate forms without the trinominal symbol. But
the number of names introduced will depend upon the judg-
ment of the systematist ; and in this respect there is often
great diversity of opinion. Actual proof, moreover, of the
intergradation of different forms is by no means always
available. In this case decision as to the specific or sub-
specific status of a distinguishable form is also a matter for
the judgment of the individual systematist. and considerable
discrepancy of view on this point may arise. At present
there is no unanimity ; and several of the forms I have treated
as subspecies in this volume may be regarded by others as
species or held to be unnecessarily named*.
Mr. Oldfleld Thomas, then in charge of the collection of
Mammals in the British Museum, was at once impressed by the
interesting results achieved in North America by Merriam
and his staff of collectors and students. He conceived the-
idea of extending the same scheme to all possible countries of
the Old World, and lost no time in putting it in practice.
When Mr. R. C. Wroughton, U.S., on retiring from office
and coming to England, offered his services as voluntary
assistant to Mr. Thomas, they speedily discovered the poverty
of the National Collection in Oriental mammals, and decided,
if possible, to inaugurate a collecting campaign in British
* The modern method has, however, been by no means an unmixed
benefit. It attracted into systematic mammalogy on both sides of the-
Atlantic amateurs to whom the determination of species was child's
play, comparable to the matching of samples of silk or wool, and -whose
sole qualification for the work was ability to detect differences in. the
colour or pattern of skins and the shapes of skulls, without the training
and judgment required to assess their value. It engendered the idea
that a difference of locality was sure to be accompanied by a difference
in colour, coat or some other part, and that if such could be perceived
it was worth symbolizing by a new name, no regard being paid to the
great possibility of age or season being the cause, thus bringing the
distinguishing feature within the range of individual variation. The
late Mr. R. C. Wroughton once humorously expressed this conception
to me by saying that two hundred miles was a good specific character !
The net result of this influx of inexperienced workers on the new lines
has been the addition of large numbers of names which, although
without real significance, have to be reckoned with and investigated by
the serious student, thus leading to waste of time and making his task,
difficult enough already, still more exacting and arduous.
Vlll PREFACE .
India, with the ultimate object of issuing a new edition of
Blanford's volume. The first and, as it proved, the wisest
-step was to get in touch with Mr. W. S. Millard, the Honorary-
Secretary of the Bombay Natural History Society. With
characteristic energy and enthusiasm Mr. Millard at once
took the matter in hand and, with the sanction of the Com-
mittee of the Society, proceeded to collect funds to defray
the cost of sending collectors to various districts of the
Indian Empire and also to interest sportsmen, forest officers,
and other members of the Society in the enterprise. Thus
was started, at first in a small way, in 1911 the Mammal
Survey of India, Ceylon, and Burma, which, owing to the
generosity of numerous subscribers and the diligence of the
employed and voluntary collectors, yielded in the following
years results which far exceeded expectations. The outbreak
and duration of the world war temporarily suspended the
activities of the Survey ; but they were later resumed, and
after April 1920, when Mr. Millard retired and came to England,
they were carried on by Mr. R. A. (now Sir Reginald) Spence,
who succeeded him as Hon. Secretary to the Society.
Donations to the fund were duly acknowledged, as received,
in the Society's ' Journal ' ; but since the present volume on
the Mammals of British India, and those that are to follow,
are based to a very great extent upon the results of the Survey,
it is fitting that the names of the individuals, Governments,
Societies, and other bodies that contributed to its success
■should be put permanently on record.
The principal subscribers were as follows : — The Govern-
ments of India, Ceylon, and Burma, of Bihar and Orissa,
Bombay, the Central Provinces, the United Provinces, Madras,
the Federated MalayStates, and Junagadh State ; the Trustees
•of the British Museum, the Royal and Zoological Societies of
London, and the Trustees of the late N. M. Wadia, CLE. ;
H.H. the Maharaja of Alwar, Sayarijao Gaikwad of Baroda,
the Maharaja of Bhavnagar, the Maharaja of Bikanir, the
Rao of Cutch, the Raja of Dhar, the Nizam of Hyderabad,
the Maharaja Tukojirao Holkar of Indore, the Maharaja of
Jodhpur, the Maharao of Kotah, the Maharaja of Mysore,
PREFACE. IX
and the Maharaja Sir Madhowrao Scindia of Gwalior ; the
Most Hon. the Marquis of Bute, Sir Dorab J. Tata, the
Hon. N. C. Rothschild, the Hon. E. S. Montagu, M.P., Sir
Ratan J. Tata, and Messrs. E. Comber, W. S. Millard, H. M.
Phipson, and Oldfield Thomas. Finally, the last of the
expeditions, sent out by the Society in 1929 to the Eastern
Ghats, where large numbers of birds as well as mammals
were secured, was generously financed by Mr. A. S. Vernay.
Of the collectors, the first to be employed by the Society
for the Mammal Survey was Mr. C. A. Crump, and when
sufficient money had been received, Mr. Oldfield Thomas
sent out to India Mr. G. C. Shortridge and, later on, Major A. W.
Mayor. In addition to these collectors, Messrs. S. H. Prater,
C. McCann, and N. A. Baptista — who were employees on the
staff of the Society — also assisted in collecting for the Survey.
In 1914 — owing to the War — Major Mayor, Mr. Shortridge,
andMr. Crump " joined up " in Prance, but Messrs. C. Primrose,
Ryley O'Brien, and H. W. Wells were still able to give their
assistance ; and when in Burma Mr. Shortridge availed himself
of the services of the late Capt. MacMillan. But the work of
the collectors was greatly helped by influential residents in
various districts, by H.H. the Maharao Sahib in Kutch,
H.H. the Maharaja Sir Madhowrao Scindia in Gwalior, Messrs.
II. Stevens and R. S. Lister in Darjeeling, the late A. P. Kinloch
in Malabar, A. H. A. Simcox hi Khandesh, and in Burma by the
late Sir Harvey Adamson, the Governor, and Sir Godfrey Fell.
Many valuable and interesting specimens were also con-
tributed to the Survey by voluntary collectors. Of these
the principal were Col. A. E. Ward in Kashmir, Col. C. H.
Stockley in the Punjab and western Himalayas, Mr J. P. Mills
in Assam, Col. J.E. B. (now Sir Ernest) Hotson in Baluchistan,
Mr. W. W. A. Phillips in Ceylon, and by Mr. J. M JD. Mackenzie,
Mr. Kingdon Ward, and Lord Cranbrook in Burma. Of
special moment were the extensive collections made by
Mr. J. M. D. Mackenzie and Sir Ernest Hotson.
The importance of the survey material was due not only
to the large numbers of specimens secured, but also to the
X PEBFACE.
accurate information supplied regarding the dates, localities,
altitudes, sex, and measurements of most of the skins. The
collections from each district were packed in Bombay and sent
to the British Museum of Natural History to be identified by
the official staff and the voluntary helpers, Mr. R C. Wroughton,
Mr. T. B. Fry, Miss Ryley, and Mrs. Lindsay. It was arranged
that the British Museum should keep the " types " of new
forms discovered*, as well as a reasonable number of other
specimens required, the remainder to be returned to Bombay
to be dealt with as the Natural History Society thought fit.
The carrying out of the scheme involved a vast amount of work
at both ends, and in this respect the services of Mr. R. C.
Wroughton in London and of Mr. N. B. Kinnear in Bombay
cannot be overestimated. The reports were published
seriatim in the 'Journal' of the Bombay Society, with the
scientific names and localities of the species and such notes
regarding habits and the vernacular names as were supplied
by the collectors.
It was the intention of Mr. Hinton and Mr. Wroughton
finally to collate the results as a basis for a new edition of
Blanford's volume. When Mr. Wroughton's untimely death
brought this scheme to a close, Mr. Hinton invited me, on
my retirement from the Zoological Society in 1923, to colla-
borate with him in the work, each of us doing such groups as
we had previously particularly studied. This I undertook,
little realizing the length of time that would be required
owing to the vast amount of material to be handled, the
numbers of skulls to be measured, the extent to which the
names and characters of the described genera, species, and
subspecies called for revision, and the necessity for studying
the mammals of all the adjoining districts of Asia before
those of British India could be adequately dealt with.
One of the first things that came to light was the general
poverty of the British Museum collection in representatives
* It was agreed that the types should be kept in London because the
climatic conditions in Bombay are not so favourable for their preserva-
tion.
PREFACE. XI
of such familiar Indian animals as lions, tigers, panthers,
snow-leopards, bears, wolves, and, as has been subsequently
proved, of nearly all the species usually understood by the
phrase " big game " *. These deficiencies had to be made
good, as far as was possible, for the first two volumes of the
Mammalian Fauna. Appeals published in the ' Journal of the
Bombay Natural History Society,' begging-letters to many
of its members, and requests to personal friends for specimens
met with immediate response, and I am especially appreciative
of the generosity of those who sent to me for the work in hand
valuable and handsome skins, as well as skulls, of the larger
Carnivora, of which the British Museum was particularly in
need. A point has been made of citing the names of the donors
where the specimens are referred to in the text ; but in addition
I take this opportunity of recording my indebtedness to the
following sportsmen not only for the animals from India,
Ceylon, Burma, and adjoining countries, but also in mam-
cases for information regarding their habits and occurrence : —
H.H. The Maharaj Kumar of Bikanir.
Col. E. M. Bailey, CLE.
The late Sir Charles Bell.
Major G. Burrard, D.S.O.
Mr. C. E. Capito, O.B.E.
Sir E. Dane, K.C.I.E., and Col. R. Dane.
Mr. A. Dunbar Brander, O.B.E.
* Several factors contributed to tMs end. For many years there was
a widespread opinion that such beasts -were of comparatively little
zoological interest and were not wanted by the National Collection.
Sportsmen also are very naturally loth to part with handsome skins
and heads, preferring to preserve them as trophies and cherished
mementoes of the chase. Finally the Survey collectors were requested
to concentrate upon the smaller members of the mammalian fauna
and to leave big game unmolested so as to avoid trespassing upon what
might be considered to be the sportsman's privilege. This decision,
if wise, was regrettable because tigers and panthers show environmental
adaptations as well as rats and mice, and because, with the progressive
reclaiming of the country from the wild and bringing it under cultivation,
the larger species will be the first to disappear, and well-preserved repre-
sentatives of them are required by the museums of the world before
that fate overtakes them. As an instance may be cited the now
irretrievable loss to the British Museum of examples of the two finest
races of lions, from the Cape and Algeria respectively, owing to the
authorities in the past being uninterested in these animals and not
foreseeing their inevitable extinction.
Xii PREFACE.
Mr. E. C. Fernando.
Mr. R. Kaulback.
Mr. N. B. Kinnear.
Major Donald Lowndes.
Mr. R. C. Morris.
Col. A. H. E. Mosse, CLE.
Mr. Fenwick Owen.
Mr. E. H. Peacock.
Mr. W. W. A. Phillips.
Major E. G. Phythian Adams.
Capt. A. H. K. Sangster.
Mr. H. C. Smith, with the assistance of Mr. P. F.
Garthwaite.
Col. C. H. Stockley, D.S.O., O.B.E.
Mr. H. J. Todd.
Col. W. R. F. Trevelyan.
Mr. H. Whistler.
I am also indebted to Mr. N. B. Kinnear for advice and
suggestions on various points, and very particularly to
Mr. W. S. Millard, a personal friend of many years, who has
consistently and sympathetically helped me in too many
ways to mention in detail *.
For permission to use many blocks for illustrations I am
indebted to the Bombay Natural History Society, and for the
loan of " types " or other interesting specimens I am under
great obligations to the Hon. Secretaries and other officials
of the Bombay Natural History Society, to Col. Seymour
Sewell, CLE., F.R.S., and Dr. Baini Prashad, who succeeded
him as Director of the Indian Museum, Calcutta, to the
Smithsonian Institution, Washington, and to the Field
Museum, Chicago.
For the purely zoological part of this volume, with its
avoidable and unavoidable mistakes, I take full responsibility,
* As an instance I may say that it was owing to his kind offices that
Col. A. H. E. Mosse secured for the British Museum the first skulls of
wild-killed examples of the Indian lion.
PREFACE. Xlll
The vernacular names of the species hare been accepted on
trust ; and since my personal knowledge of the habits of the
animals is limited to observations on captive specimens in
the Zoological Gardens of London, I have been compelled to
borrow the information on this head from the writings of
others. But the literature on this subject in the way of books
and stray notes in periodicals is now so extensive that it has
been impossible to become acquainted with it all or to acknow-
ledge in every ease the authority for statements made. It
will, I trust, be understood that the omission to quote the
observations of many sportsmen was due, not to inappreciation
of their value, but to the bar to their inclusion imposed by
the limit of time and the size of the volume.
This volume includes the Apes, Monkeys, and Lemurs of the
order Primates and the first two families, the Cats and Civets,
of the order Carnivora. It is hoped that the second volume,
containing the remaining families of the Carnivora, and some
at all events of the groups of Hoofed Mammals, will be issued
in about a year's time.
R. I. POCOCK.
Mareh 4th. 19:SS.
INTRODUCTION.
THE GEOGRAPHICAL DISTRIBUTION OF THE
MAMMALS OF BRITISH INDIA.
Fbom the zoological standpoint the greater part of British
India is a portion of the so-called Oriental Region, one of the
primary subdivisions into which the land-surface of the world
has been divided, mainly on the evidence of the average
distinctiveness of their mammalian and avine faunas.
In south-eastern Asia the usually accepted boundary of
this region is " Wallace's line," passing between the islands
of Bah and Lombok to the east of Java, and continued
northward between Borneo and Celebes, a line marked by
deep sea indicating a long-continued marine barrier which
prevented most of the highly organized Oriental mammals,
like the apes, leaf-monkeys, cats, mongooses, members of the
dog and weasel families, bears, banteng cattle, rhinoceroses,
and many others which were later comers to that part of
Asia, from reaching Celebes and other islands to the east of
the line, and similarly prevented the lowly organized marsupial
mammals, characteristic of the Australian Region, from
entering Borneo and Java. But some of the typically
Oriental mammals, like macaque monkeys, palm-civets,
deer, buffalo, pigs, and pangolins, were sufficiently early
migrants from the west to get a footing in Celebes or other
islands to the east of what is now Wallace's line, so that
there is here a blend of the faunas of the two regions.
Still less satisfactory are the boundaries of the Oriental
Region to the north, where, owing to its complete land-
continuity with south-western, central, and northern Asia,
which, with Europe and North Africa and most of North
America, constitutes the Holarctic Region, the faunas of
the two so intimately intergrade that the line defining them
can only be arbitrarily drawn roughly from Central China
along the Himalayas and thence southwards through the
deserts of north-western India to the Arabian Sea.
XVI IKTRODTTOTIOIv.
In the extreme variety of its physical features India proper,
or Hindustan, is surpassed by no country of similar extent
in the world. It is divided into three main areas, the
Himalayas in the north, the great Indo-Gangetic Plain in
the centre, and the table-land of Peninsular India in the south.
Although usually described as a continuous mass of
mountains, the Himalayas are composed of numerous ranges
lying roughly parallel to each other and Separated and inter-
sected by river valleys. The principal ranges are the
following * : — The Great Himalayan Range, lying almost in its
entirety in British India, and stretching from the southward bend
of the Brahmaputra in the east to that of the Indus in the west.
In the west of Nepal it gives off a northern range, the Zaskar,
which extends nearly as far to the west as the main axis ;
and from its southern side come two ranges, the Dhauladhar,
rising in Tehri Garhwal and dying out near Chamba, and
farther to the west the Pir Panjal, starting near the source
of the Chenab and stretching south of it, and of Srinagar as
far as the Jhelum, where it gradually decreases in altitude.
North of the Zaskar Range in Kashmir come in order from
south to north the Ladak Range, which eastward forms
part of the northern boundary of British India in Nepal and
Bhutan, the rest of it being in Tibetan territory : the Kailas
Range, which sweeps across Tibet to the north of Lhasa,
forming the northern boundary of the lower-lying open
country traversed by the Brahmaputra and, in the west,
by the upper waters of the Sutlej and Indus, an area bounded
on the south by the Ladak Range ; and the Karakorum
Range running from the Tibet boundary to the Hindu Kush,
the two forming the extreme northern boundary of British
India in the north-west. The northern slopes of the Himalayas,
pass by means of a stretch of upland grass into the high
plateau of Tibet, and it is along the Tibetan frontier that the
Holarctie element of open-country species enters the extreme
north of Hindustan. On the southern slopes of the mountain
range the conditions are different, ranging from arctic in the
higher summits to tropical below the foothills, the vegetation
passing gradually from alpine through temperate to evergreen
forests and jungles.
All the numerous rivers which intersect and form passes
through the main Himalayan Range, except in the extreme
east, are tributaries of the Indus and Ganges which give then-
name to the Indo-Gangetic Plain stretching across India
from Sind to the north of the Bay of Bengal. In the lower
Punjab and Sind this plain has low humidity and characteristic
* These particulars are taken from Major Burrard's map in his
book ' Big Game Hunting in the Himalayas and Tibet,' 1925.
INTRODUCTION. xvii
desert vegetation. Westward it extends through Baluchistan
and Persia to Mesopotamia ; but eastward, with increasing
humidity, it gradually changes its character, passing finally
into forest and into the jungly swamps and islets of the
Sundarbans about the mouths of the Ganges and Brahmaputra.
The Vindhya Hills and other northern ranges mark the rise of
Peninsular India from the Indo-Gangetic Plain. The greater
part of it is a table-land consisting of hills and grass-covered
plains intersected everywhere by the tribiitaries of its main
rivers, which cross it from side to side, the Narbada and the
Tapti in the north flowing from east to west, the Mahanadi,
the Godaveri, the Kistna, and the Kauveri in a general wajr
from west to east. Near the Carnatic and Malabar coasts-
it is flanked respectively by the Eastern and Western Ghats
and outlying ranges. The Western Ghats, supplemented by
the Nilgiri, Animalai, and other hills, have a profound effect
upon the general features of the Peninsula. They intercept the
south-west monsoon, causing exceptionally heavy rainfall,
mainly on their seaward slopes, depriving the country to the
east of much moisture and converting it into a comparatively
dry zone. Naturally the vegetation of the two areas is very
different in accordance with the rainfall ; where it is moderate-
to the east of the Ghats there are grass-covered plains, with
clumps of acacias and stunted, mostly deciduous, trees,
here and there forming copses or small forests. The seaward
slopes of the Western Ghats, on the contrary, are mostly
covered with dense forests of lofty evergreen trees, festooned
with creepers, and accompanied by luxuriant growths of
bamboo. In the Nilgiris and other South Indian hill ranges
there are wide, open, grass-covered areas broken up by thickly
forested gorges or sholas *.
Ceylon in its physical features is like Southern India, of
which it formed a part before the age of mammals. It is
divided into three mtergrading zones | : — (1) A dry zone
occupying roughly the whole of the northern, eastern, and
south-eastern parts of the island, and consisting principally
of low-lying, jungle-covered country with an annual rainfall
of from 25 to 75 inches, falling mostly during the north-east
monsoon. This zone in its physical features and mammalian-
fauna recalls the south-eastern area of Southern India.
(2) A central hill zone consisting of plateaus and mountains,
sometimes over 8000 feet, covered with virgin forest and
grassland, with a rainfall of from 80 to 200 inches, generally
* An excellent summary of the main physical features of India,
illustrated with good photographs, was given by S. H. Prater in his-
" The Wild Animals of the Indian Empire," published in the Journ.
Bomb. Nat. Trig* Soc. xxxvi. no. 1, pp. 15-21, 1933.
t See Phillip's Man. Mamm. Ceylon, 1935.
VOL. I. b
XVU1 ETTRODTTCTIOlir.
distributed throughout the year. (3) A small, low country,
wet zone to the south-west of the hill zone, consisting of
low, forest- covered hills and broken country, with a rainfall
of about 200 inches, falling mainly during the south-west
monsoon, although there are heavy storms during the north-
east monsoon. This and the hill zone tolerably closely
resemble the Malabar tract of southern India.
Burma*, which for long ages has formed a continuous
land-area with China to the north and what is now the Malay
Peninsula to the south, is less varied in its physical features
than India and differs from it in some respects. It is inter-
sected by rivers running mainly from north to south and
separated by higher and lower hill-ranges forming the water-
sheds of the tributaries that feed them. The central part,
■occupying about one-fourth of the country, with a rainfall
■of only about 20 inches in the year, and vegetation that has
been described as " scrub-forest," is called the " Dry Zone."
The greater part of the rest of the country has a much higher
rainfall, reaching as much as 200 inches in the extreme south,
■and is mostly covered with heavy evergreen forests ; but
there are extensive areas of jungle grass, up to about 12 feet
high, along the banks of the larger rivers and in other places
subject to inundations. Where the soil is poorer the grasses
grow to about half that height ; and many of the hill-tops,
at an altitude of about 5000 feet or so, are often treeless and
covered with comparatively short grass.
The general trend of the rivers and hill-ranges from north
to south offers no obstacle to the migration of the mammalian
fauna in either direction ; and the general uniformity of the
forest conditions to the north and south of the dry zone
results in corresponding similarity in the fauna throughout
the forested areas. But the forested area of lower Tenasserim
and the northern part of Peninsular Siam differs somewhat,
and some of its mammals are Malayan in type.
Of the Oriental Mammals, restricted, or nearly so f, to
the region, the following are the principal genera and species
which occur in British India : — The Asiatic tapir (Acroco&ia),
rhinoceroses (Rhinoceros, Dicerorhinu$), elephant {Elephas),
buffalo (Aiwa) %, gaur an d banteng (Bibos), nylghai (Bosdwphus)
* See E. H. Peacock's 'A Game-Book for Burma and adioinine
Territories ' (Witherby, 1933). *
t A few of the genera encroach on areas assigned to the Holaretie
Region, like the goral in Korea and the black bear in Manchuria and
Baluchistan, whereas Temminck's cat and the macaque monkeys
have outlying, isolated representatives in tropical Africa and Morocco
respectively. The smaller mammals, needing revision, are for the most
part omitted from this account.
% The American tapirs and the African rhinoceroses, elephant, and
ouHalo belong to distinct genera.
rNTBODTTCTION. xix
four-horned antelope (Tetracerus), blaokbuek (Antilope), serow
(Gapricornis), goral (Nsemorhedus), takin (Budorcas), sambhar
(Busa), cbital and hog-deer {Axis), swamp-deer and thamin
(Buc&rvus), muntjac (Muntiacus), mouse-deer (Meminna,
Tragulus), gibbons (Hylobates), leaf-monkeys {Semnopithecus,
Presbytis, and two other genera), macaques (Macaco), lorises
(Loris, Nycticebus), clouded leopard (Neofelis), marbled cat
(Pardofelis), Temminck's cat (Profelis), leopard- eat, fishing
cat, rusty-spotted cat (Prionailurus), civets (Viverra, Moscho-
thera, Viverricula), palm-civets (Paradoxurus, Paguma, Hemi-
galus, Arctogalidia), binturong (Arctictis), linsang (Prionodon),
several species of mongoose (Herpestes), smooth- coated otter
(Lutrogale), clawless otter (Amblonyx), hog-badger (Arctonyx),
ferret-badger (Helictis), panda (Ailurus), sloth-bear (Melursus),
black bear (Selenarctos) , sun-bear (Helarctos), pangolins
(Manis, Paramamis, Phatages), flying lemur or colugo (Galeo-
pterus), tree-shrews (Tupaia and others), and many squirrels
(Ratufa, Funambulus, Cailosciurus, etc.), as well as many
additional rodents and several kinds of bats.
Mixed with the above-mentioned mammals, which help
to stamp the distinctness of the Oriental Region, are many
others identical with, or closely related to, species widely
distributed in districts of central or south-western Asia,
or both, assigned to the Holarctic Region, or even in Africa.
Many, indeed, of the most prominent members of the British
Indian mammalian fauna come into this category. Some,
like the tiger, panther, jungle-cat, wild dog, jackal, and wild
boar, occur in Burma as well as in Hindustan and, except
the tiger and wild dog, in Ceylon also. Others, found only
to the west of the Bay of Bengal, but not in Ceylon, are the
lion, hunting leopard, caracal, desert-cat, hyaena, ratel, and
chinkara (gazelle), which are very nearly allied to similar
types occurring in south-western Asia and Africa, the smaller
Indian wolf, which ranges into Persia and Mesopotamia, and
the tahr (Hemitragus), of which the three isolated species
are found respectively in the Himalayas, the Nilgiri Hills,
and southern Arabia. To this series should be added the
desert fox, the wild ass or ghorkha, the Sind ibex, and the
wild sheep or gad, which spread from Persia for a short distance
into the plains and mountains of Baluchistan and north-
western India at least to the valley of the Indus.
A considerable number of the mammals, both strictly
Oriental and partly Holarctic, inhabiting Peninsular India
and the Gangetic Plains are found in the Tarai, the foothills
and the forest slopes of the Himalayas, some of them ascending
to tolerably high altitudes. Of these the most important are
the elephant, buffalo, gaur (bison), nylghai, four-horned
antelope, blackbuck, sambhar, chital, hog-deer, swamp-deer,,
62
xx iOTBODTrcTicrar.
muntjac, wild boar, ratel, tiger, leopard, jungle-cat, leopard-
cat, common palm-civet (toddy-cat), little civet, grey mon-
goose, Bengal fox, rhesus macaque, and entellus langur.
They overlap several Oriental forms not found elsewhere
to the west of the Bay of Bengal which have passed along
the Himalayas from southern China and Upper Burma,
as recorded below. But although the dominance of Oriental
forms puts this mountain range within the Oriental Region,
there is a strong Holarctie infusion from Central Asia, especially
in the western Himalayas, where typical Oriental types
gradually disappear, although a few, like the leopard-cat,
toddy-cat, yellow-throated marten, rhesus macaque, entellus,
langur, and serow, occur in the hills of Kashmir.
Of these northern Holarctie immigrants into British Indian
territory the most interesting are the Siberian ibex, the
barasingh or Kashmir stag, and the shou, these two, representing
the genus Cervus, not found elsewhere within our limits,
the musk-deer, blue sheep or bharal, the snow-leopard or
ounce, the lynx, Pallas's cat, the hill-fox, Tibetan or woolly
wolf, the red bear (Ursus), and several members of the weasel
family, including the pine-marten and Siberian mink *.
Some of these, like the snow-leopard, bharal and musk-
deer, and Siberian mink, occur apparently throughout the
Himalayan Range, the two first above the tree-line, the two
last below it ; but most of them are restricted to the western
portions, west of Nepal, although the hill-fox reaches Sikkim.
A few species, like the markhor, the hoary fox, and the ermine,
penetrate British India for a comparatively short distance
from Afghanistan and the Hindu Kush. On the open,
undulating plains to the north of the mountains, and as far
west as Ladak, are found the yak, the chiru, the Tibetan
gazelle, Hodgson's argali, the kiang, the Tibetan sand-fox,
and the Tibetan polecat.
As stated above, the mammals of the Indo-Gangetic Plain
overlap those of the Himalayas, and in the west the desert
types, which pass into Baluchistan. Southwards they
similarly blend with those of Peninsular India, especially with
those inhabiting the dry zone of this area. But, as might be
expected, the hill ranges of the western and southern parts
of the Peninsula, characterized by heavy rainfall and mostly
by luxuriant forests, have some peculiar features in their
mammalian fauna. The lion-tailed macaque, John's leaf-
monkey, the stripe-necked and brown mongooses, Gwatkin's
yellow-throated marten, and the Nilgiri ibex are typical
forms. Of these the macaque is restricted to this district,
* Characteristic Central Asiatic rodents that reach British India
are the marmots and mouse-hares or picas.
INTKODTJCTION. XXI
the leaf-monkey and mongooses are the same as, or closely
akin to, forms found in Ceylon, whereas the marten and
the ibex have no allies in any part of Hindustan but the
Himalayas.
There is no proof that Ceylon derived its mammalian
fauna from any source but India. Within the mammalian
era there is geological evidence of two unions with that country.
There may have been a connection during the Pliocene, which
admitted some early types ; but the first of the two unions
above mentioned probably occurred during the Pleistocene
Ice Age when the mammals of northern India were driven
southwards and entered the Ceylonese area (see p. xxvi).
Here, by the subsequent conversion of the area into an island,
they were isolated for a time, and some of them at least
became differentiated into the peculiar forms now characteristic
of Ceylon. The second connection took place much later,
and enduring until comparatively recently, possibly into
historic times, admitted a fresh influx of Indian species and
the northward migration of some Ceylonese forms into
southern India. Theoretically the most differentiated of
the existing Ceylonese species are the descendants of the
earlier immigrants, whereas those that are at most slightly
different from Indian species or indistinguishable from them
probably came in on the second occasion.
The mammal fauna of the island is interesting from many
points of view, not the least being the light it throws on the
period, earlier or later, of the occupation of southern India
by the species now found there. Some were sufficiently
early, others were too late to avail themselves of the land-
bridge over what is now Palk Strait. Of the better-known
Ceylonese forms that are identical with those found in S. India,
or nearly allied to them, the following are the most important :
the leaf-monkeys, macaques, loris, leopard, fishing-cat,
rusty-spotted eat, jungle-cat, little civet, palm-civets,
mongooses, jackal, otter, sloth-bear, pangolin, blaek-
naped hare, porcupine, many squirrels and other Rodents,
and Insectivores of the Shrew family, wild boar, mouse-
deer, muntjac, hog-deer, chital, sambhar, buffalo, and
elephant. Some of these forms, like the monkeys, loris,
rusty-spotted cat, palm-civets of the zeylonensis type,
mongooses, bear, pangolin, hare, porcupine, mouse-deer,
and chital, are restricted to Ceylon and India proper ; and
a few of them, namely, the leaf-monkey (Kasi), macaques,
loris, rusty-spotted cat, palm-civets, two at least of the
species of mongooses, hare, and mouse-deer, are found solely
or mainly in the southern part of Peninsular India and Ceylon.
Of the Indian species that do not occur in Ceylon it is
needless to refer to the northern types mostly restricted to
XXil INTRODUCTION.
the Himalayas. But there are many southern species,
some found far to the south in Peninsular India, which arrived
there after the final separation of the island or were excluded
from it by other factors. The principal species are : the
lion-tailed macaque, tiger, leopard-cat, caracal, hunting
leopard, Malabar civet, hyaena, wild dog, fox, marten, ratel,
tree-shrew, Nilgiri ibex, gazelle, blackbuck, four-homed
antelope, nilghai, and gaur or bison.
Although a considerable number of familiar Indian mammals,
like elephant, buffalo, gaur (bison), sambhar, hog-deer,
barking deer (muntjac), wild boar, tiger, leopard, jungle-cat,
fishing-cat, leopard-cat, small civet, common palm-civet
(toddy-cat), jackal, wild dog, smooth-coated and clawless
otters, and rhesus monkey, occur both in Burma and India,
the mammalian fauna of Burma differs from that of India
south of the Himalayas both on the negative and positive
sides. On the negative side it is characterized by the absence
of such species as the lion, hunting leopard, caracal, wolf,
hyaena, ratel, wild ass, sheep, goats, and gazelle, which came
into India from the west, and of such typically Indian species
as the nylghai, four-horned antelope, blackbuck, chital or
spotted deer, swamp-deer, entellus-langur, sloth-bear, and
pangolin. On the positive side Burma has a large number
of genera which in India are not found south of the Himalayas,
its fauna resembling especially that of the eastern part of
these mountains and of the hills of Assam and also that of
China, Indo-China, and the rest of south-eastern Asia as far
as Wallace's line. This resemblance is apparently due very
largely to the migration of typically Chinese forms westwards
along the Himalayas sometimes as far as Kashmir, and
southwards through Burma, Indo-China, and Siam to Malaya
and beyond*. The following are the principal genera and
species, with their geographical distribution, which are found
in Burma and mainly in the mountainous parts of northern
India, but not in the plain of the Ganges or southward of it
in India : gibbons, Assam to Borneo ; pig-tailed macaque,
Assam to Borneo ; Assamese macaque, Nepal to Upper Burma
and Tongking ; stump-tailed macaque, Assam to Malaya ;
leaf-monkey (Trachypithecus) and slow loris, Assam to Borneo ;
Temminck's cat, Nepal to Burma, China, and Sumatra ;
the marbled cat from Nepal to Borneo ; the clouded leopard
from Nepal to China and to Borneo ; the spotted linsang from
Nepal to Tongking ; the large civet from Nepal and China to
Malaya ; the large spotted civet from Burma to Malaya ;
* The mammalian faunas of Malaya, Sumatra, Java, Borneo, and
the Philippines have certain peculiarities which are beyond the scope
of this volume.
INTRODTrCmON. XXUL
the palm-civet (Paguma) from Nepal to Borneo ; the binturong
from Sikkim to Palawan ; the three-striped palm-civet from
Assam to Borneo ; the little mongoose from Kashmir to
Java * ; the crab-eating mongoose from Nepal to China
and to Malaya ; the yellow-throated marten from Kashmir
to China and Borneo ; the Siberian mink from Kashmir to
China and Sumatra ; the striped weasel from Nepal to Indo-
China and Lower Burma ; the yellow-bellied weasel from
Kashmir to China and Indo-China ; the hog-badger from
Sikkim to China and Sumatra ; the ferret-badger from Nepal
to China and Borneo ; the Himalayan black bear from
Baluchistan through Kashmir to China and Burma ; the
Malayan or sun-bear from Burma to Borneo ; the panda,
Nepal to China ; the two-horned rhinoceros from Bhutan to
Borneo ; the lesser one-horned rhinoceros from Sikkim and
the Sundarbans to Borneo ; the banteng or tsain, Burma to
Borneo ; takin, Bhutan to China and Upper Burma ; serow,
Kashmir to China and Sumatra ; the goral from Kashmir to
China and Lower Burma ; thamin or Eld's deer, Burma,
Indo-China, Malaya f ; pangolins, one from Nepal to China
and Burma, the other from Burma to Borneo and Celebes.
In Tenasserim, as far north as the Y6 River, the fauna
has a marked, so-called Malayan element, represented by
the tapir, Mergui to Sumatra ; mouse-deer (Tragulus),
Tenasserim to Borneo ; flying-lemur or colugo, Mergui to the
Philippines ; banded palm-civet, Tenasserim to Borneo ;
banded linsang, Tenasserim to Borneo ; the leaf-monkey
(Presbytis) Tavoy to Borneo. To these should probably be
added as essentially southern types, although spreading into
Lower Burma north and west of Tenasserim, the lar-gibbon,
which is closely related to similar gibbons spreading as far
as Borneo, and the crab-eating macaque, which under a variety
of forms reaches to the Philippine Islands.
Since most of these mammals have no allied species in
India south of the Himalayas, the contrast between the
faunas of the two areas is profound. It is, however, lessened
in a measure by kinship in a few instances. The pig-tailed
macaque, for example, has its nearest ally in the lion- tailed
macaque of Malabar ; the leaf-monkey (Trachypithecus)
is nearly as closely akin to the leaf -monkey (Khasi) of south-
western India and Ceylon, and the slow loris and the Ceylonese
and South Indian slender loris are the only genera of the
lemurs found in the Oriental Region. Similarly, the mouse-
deer or chevrotain {Tragulus) of Tenasserim and the related
* This species extends to Persia and to Gujerat in India,
t Schomburgk's deer from Siam is also a member of this fauna,
although not recorded from Burma.
XXiv INTRODUCTION.
genus (Meminna) of Peninsular India are the sole examples
of that group of ruminants occurring within British Indian
limits. The thamin or Eld's deer, and, incidentally, Schom-
burgk's deer, have their nearest ally in the swamp-deer of
parts of Northern India ; the large-spotted Burmese civet is
closely related to the Malabar civet ; and the common yellow-
throated marten is represented by an allied species in Southern
India.
The Andamans and Mcobars, a chain of mainly forested
islands in the eastern part of the Bay of Bengal, and marking
the site of an ancient land-bridge between the Pegu district
of Burma and the northern point of Sumatra, are also included
in British India, and may be granted the status of a separate
tract, characterized negatively by the absence of all the
larger typically Oriental mammals and positively by the
presence of between thirty and forty species or subspecies,
mostly rats and bats, but all of them peculiar to the Archi-
pelago, with the exception of a few, including the pig-tailed,
macaque (M. nemestrina leonina), comparatively recently
introduced by Europeans.
The islands are separated from the mainland of the Malay
Peninsula by very deep sea, indicating long isolation dating
back to a period, before the Malayan Region was occupied
by its present mammalian fauna. But the species and
subspecies inhabiting the islands belong to genera now found
•on the mainland, and are comparatively slightly differentiated.
From these facts Miller * inferred that, with the exception
•of the bats, which reached the islands by flight, the fore-
runners of the rest of the mammals were imported by the
•early native settlers and, like them, have become modified
in the interval, into the existing types. Of these, setting
aside the Bats and Rats, there is a long-tailed macaque (M. irus)
in the Mcobars related to Malayan and Sumatran races ;
a palm-civet (Paguma) in the Andamans ; a wild boar (Sits),
two shrews (Crocidura), one from the Andamans, the other
from the Mcobars, and a tree-shrew (Tupaia) from the Mcobars
which is more distinct from its mainland congeners than
are the other peculiar forms.
Reviewing the main above-described facts connected with
the distribution of mammals in British India, with others
supplied by the rodents and other orders of smaller species,
Blanford | assigned two areas of it to the Palsearctic section
* Proc. U.S. Nat. Mus. xxiv. pp. 790-1, 1902. This paper on the
Mammals of the Andamans and Nieobars has supplied me with the
information about the islands and their species given above.
f Proe. Roy. Soc. Ixvii. pp. 484-92, 1900, and Trans. Boy. Soc. cxoiv,
p. 335, 1901.
INTROBTTCTION. XXV
of the Holaretic Region. These were the Punjab tract,
including the deserts of Baluchistan, Sind, the Punjab, and
western Rajputana up to a line running roughly from Cutch
to the Himalayas nearly due north of Delhi, and the Tibetan
tract, comprising the area to the north of the Himalayas
through which the Indus and the Brahmaputra flow, and
extending westward through Ladak to Gilgit.
The rest of British India he referred to the Oriental Region,
and divided it into two subregions named, not very appro-
priately, the Cisgangetic and the Transgangetie. The Cis-
gangetic extends westwards from the ill-denned boundary
of the Punjab tract to the Bay of Bengal and the hills forming
the eastern limit of the alluvial deposits of the Ganges and
Brahmaputra and, from north to south, from the base of the
Himalayas to Cape Comorin, with the addition of Ceylon.
This subregion was further subdivided into several sub-
ordinate tracts, of which the only one that calls for special
notice is the Malabar tract, the wet zone running along the
western side of the Peninsula from the Tapti to Cape Comorin.
To this the wet zone of Ceylon is affiliated, the dry zone
of the island belonging to the tract of southern Peninsular
India of which the Kistna is the northern boundary.
The Transgangetie subregion is composed of the Himalayas
from Kashmir to Bhutan, Assam, Manipur, Tipperah, Chitta-
gong, and the whole of Burma *.
This subregion, like the Cisgangetic, is subdivided into
several tracts, of which the Andaman and Mcobar Islands
are one, but the most distinct perhaps is the Malayan tract,
including Tenasserim from the Y6 River, as Mr. H. C. Smith
tells me, not Mergui, as thought by Blanford, southwards to
Victoria Point.
Blanford sought to explain the peculiarities in the dis-
tribution of the existing mammals of British India as
resulting from the Pleistocene Ice Age and the subsequent
return of warmer conditions. At the present time the glaciers
in Sikkim do not extend below about 14,000 feet ; but there
is proof that during that cold spell they descended as low as
7,000 feet in that district, and even to lower levels in the western
Himalayas. Hence this mountain range was evidently at
that time much colder than it is now, and a considerable
area of Northern India to the south of it must have been
correspondingly affected, giving rise to conditions unsuitable
for the survival of the tropical forms that swarmed there
before the onset of the Ice Age, and were either exterminated
or driven southwards as it progressed. Later, with the
gradual return of the warmer conditions such as now prevail
* It is impossible to exclude southern China, Indo-China, Siam,
and Cambodia from this subregion.
XXvi IKXBODTTOTIOJSr.
over all the areas affected by the cold, facilities were afforded
for the reoccupation of the higher levels of the Himalayas
by such species as had survived at the base and by others
from the four main points of the compass, and for the return
to Northern India of* the descendants of the species that had
taken refuge in Southern India.
Some such theory as this affords apparently the most
plausible explanation of certain features in the distribution
of mammalian life in British India, especially of those supplied
by the wide geographical separation between some related
forms in the Transgangetic subregion and in the southern
part of the Cisgangetic subregion, which are severed by a
large tract of Central and Northern India — species and genera
of which the ancestors formerly, it may be assumed, were
continuously distributed in the northern portions of the
Oriental Region. Instances of this phenomenon have already
been cited. The most important illustrations of it are the
following : — The pig-tailed macaque, ranging from Assam to
Borneo, and the lion-tailed macaque of the Malabar tract ;
the common leaf -monkey of Assam and Burma, which also
spread to Borneo, and John's leaf-monkey of the Malabar
tract and its related Ceylonese species ; the slow loris, another
species ranging from Assam to Borneo, and the slender loris
of South India and Ceylon ; the large-spotted Burmese and
Indo-Chinese civet and the Malabar civet; the mouse-deer,
ranging from Tenasserim to Borneo, and its related genus of
Southern India and Ceylon. Finally, the common yellow-
throated marten, found practically all over the Transgangetic
subregion, and the thar of the Himalayas are represented
in some of the hill ranges of Southern India respectively by
Gwatkins's marten and the Nilgiri ibex. These species in
particular, and especially the ibex, might perhaps be cited as
cogent evidence of the truth of Blanford's hypothesis regarding
the effects of the Ice Age on the distribution of the mammalian
fauna of British India.
EXPLANATION OF PLATES.
Plate I. Faom
page
Left-hand fig. — Hoolock Gibbon (Hylobates hoolock) \ ,„
Right-hand fig. — Lar Gibbon (Hylobates lar) J
Plate II.
Upper fig. — McMahon's Rhesus Macaque (Macaca mulatto]
mcmahoni) [ ,*
Lower fig. — Common Rhesus Macaque (Macaca macaca [
mulatta J
Plate in.
Upper fig. — Assamese Macaque (Macaca assamensis) \ gg
Lower fig. — Stump-tailed Macaque (Macaca speciosa) J
Plate IV.
Lion-tailed Macaque (Macaca silenus) 66
Plate V.
A. Head of Pig-tailed Macaque (Macaca nemestrina),'
typical form
B. Head of Burmese Crab-eating Macaque (Macaca irus
aurea)
C. Head of typical Macaca irus, showing hair-growth on
the cheek
79
Plate VI.
1. The Kangra Langur (Semnopithecus entellus ajax) "|
2. The Nepal Langur (Semnopithecus entellus achiUes) > 97
3. The Tarai Langur (Semnopithecus entellus schistaceus) . . J
Plate VII.
Upper fig. — Slow Loris (Nycticebus coucang) \ ,g 5
Lower fig. — Slender Loris (Loris tardigradus) J
XXviii EXPLANATION OF PLATES.
Plate VIII. Facing
A. Face of Nycticebus coucang bengalensis 1 P^ 6
B. „ Nycticebus coucang tenasserimensis > 171
C. „ Nycticebus coucang coucang J
Plate IX.
Tiger stalking Spotted Deer 197
Plate X.
Upper fig. — Head of Indian Lion . (Adapted from engraving'
by Bennett of specimen from Hariana exhibited in the
Tower of London.) V 212
Lower fig. — The "Maneless Lion of Gujerat." (Drawn
from one of Capt. Smee's specimens)
Plate XI.
Variety of Indian Leopard from Cuddapah 223
Plate XII.
Variety of Indian Leopard from Kanara 225
Plate XIII.
A & D. " Waist " and side view of skull of Indian \ „„a
Leopard. B & C. The same of Indian Leopardess . . j
Plate XIV.
Skin of Indian Leopard with exceptionally bold pattern \ 00Q
from Pvhikhikesh / ^ y
Plate XV.
Skin of Millard's Leopard (Panthera pardus millardi) from\ 00 „
Kashmir J i6i
Plate XVI.
Skull of Snow-Leopard. A. Side view. B. "Waist.""
C. Nasals. D. Left auditory bulla, the dotted line
showing position of partition. E. The same of
Common Leopard
> 239
Plate XVII.
Snow-Leopard or Ounce (Vncia imcia) 240
EXPLANATION OF PLATES. XXIX
PLATE XVIII. Facing
Left-hand fig. — Mat skin of cub of Snow-Leopard "\ . ?
Right-hand fig. — Made-up skin of cub of Common Leopard . f
Plate XIX.
Clouded Leopard (Neofelis nebulosa) 24S
Plate XX.
Upper fig. — Marbled Cat (Pardofelis marmorata) \ ~_-
Lower fig. — Rusty Spotted Cat (Prionailurus rubiginosus) . J
Plate XXI.
Upper fig. — Temminck's Cat (Profelis temminckii tern-'
minckii). (Drawn from grey skin from Upper Chind-
win)
Lower fig. — Striped Temminck's Cat (Profelis temminckii
iristis) from Nam Tamai, Upper Burma
V 263
Plate XXII.
Upper fig. — Leopard-Cat (Prionailurus bengalensis) \ g »~
Lower fig. — Fishing-Cat (Prionailurus viverrinus) J
Plate XXIII.
Upper fig. — Indian Desert-Cat (Felis Constantino, ornata) . . \ ^a
Lower fig. — Jungle-Cat (Felis chaus) J
Plate XXIV.
Upper fig. — Caracal (Caracal caracal) ~1 „„„
Lower fig. — Lynx (Lynx lynx isabellinus) J
Plate XXV.
Cheetah or Hunting Leopard (Acinonyx jubatus) 323
Plate XXVI.
Upper fig. — Pallas's Cat (Otocolobus manul) ~\ „ , 9
Lower fig. — Large Indian Civet ( Viverra zibeiha) J
Plate XXVII.
Upper fig. — Malabar Civet (Moschothera civettina) \
Lower fig. — Little Civet (Viverricula indica) J
360
page
)> 390
XXX EXPLANATION OF PLATES.
Plate XXVIII.
Upper fig. — Indian Palm-Civet (Paradoxurus hermaphro-'
dittos)
Lower fig. — A & B. Heads of two examples of Paradoxurus
hermaphroditus hermaphroditus, showing variation in
pattern. C. Head of Burmese Palm-Civet (Paradox-
urus hermaphroditus laotum) from Mingun. D. Head
of Lesser Palm-Civet (Paradoxurus hermaphroditus
minor) from Tenasserim
Plate XXIX.
Upper fig. — Burmese Palm-Civet (Paradoxurus hermaphro-")
ditus laotum) \ . . „
Lower fig. — Masked Palm-Civet (Paguma larvata larvata) j
from Szechwan J
Plate XXX.
Binturong (Arctictis binturong) 433
Plate XXXI.
Banded Palm-Civet (Hemigalus derbyanus) 450
SYSTEMATIC CONTENTS.
Page
Class MAMMALIA 1
Order I. PKIMATES 13
Suborder 1. PITHECOIDEA. 14
Group Catarhini 15
Section 1. Astthbopomobpha 15
Family Hyiobatidjb 17
1. Hylobates 19
1. hoolock 19
2. lar 26
a. entelloides 26
Section 2. Cynomoepha ... 31
Family 1. Cebcoectheced-b} . . 32
1. Macaca 32
1. sinica 34
a. sinica 36
6. aurifrons 37
2. radiata 38
a. radiata 40
b. diluta 42
3. mulatta 44
a. mulatta 45
b. villosa 49
c. memahoni .... 50
4. assamensis 52
a. assamensis 53
6. pelops 55
5. nemestrina 58
a. leonina 59
6. blytbii 62
6. silenus 66
7. speciosa 69
a. speciosa 71
b. melanotus 73
8. irus 78
a. aurea 79
b. umbrosa 82
Page
Family 2. CoLOBn>.ai 83
1. Semnopithecus 88
1. entellus 88
a. schistaceus .... 92
b. acbilles 95
c. ajax 96
d. entellus 98
e. anchises 101
/. achates 103
g. iulus 104
h. seneas 106
i. dussumieri 107
j. hypoleucos .... 108
k. priam 109
I. priamellus 112
m. elissa 113
n. thersites 115
2. Trachypithecus 120
1. pileatus 121
a. pileatus 122
b. durga 125
c. tenebrieus 126
d. brahma 128
e. shortridgei 128
2. phayrei 129
a. phayrei 130
6. crepusculus .... 134
c. shanicus 136
3. obscurus 138
a. sanctorum 140
b. flavicauda 140
4. pyrrhus 142
a. atrior 143
3. Kasi 146
1. johnii 147
2. senex 150
a. vetulus 151
6. nestor 153
c. senex 154
d. montieola 156
XXX11
SYSTEMATIC CONTENTS.
Page
4. Presbytis 158
1. femoralis 139
a. keatii 161
Suborder 2. LEMUROIDEA . 163
Family 1. Lobisid^; 164
1. Nyctieebus 165
1. coueang 166
a. bengalensis 166
b. tenasserimensis . 169
c. coueang 171
2. Loris 174
1. tardigradus 175
a. lydekkeriamis . . 177
b. malabaricus ... 180
c. tardigradus .... 181
d. nordicus 182
e. grandis 184
Order II. CARNIVORA 186
Suborder 1. MLUEOIDEA... 190
Family 1. Feud^e. . . . 191
Subfamily Paxthesixjb 195
1. Panthera 196
1. tigris 197
a. tigris 199
2. leo 210
a. persiea 212
3. pardus 222
a. fusoa 226
b. pernigra 231
c. millardi 233
d. sindica 233
6. saxioolor 234
2. Uncia 239
1. uncia 240
Subfamily Wslisje 243
1. Neofelis 247
1. nebulosa 248
a. maorosoeloides . . 250
2. Pardofelis 253
1. marmorata 255
a. charltoni 256
3. Profelis 258
1. temminekii 260
a. temminekii 261
b. tristis 263
4. Prionailurus 265
1. bengalensis 267
a. bengalensis 268
6. horsfieldi 271
o. trevelyani 273
2. rubiginosus 276
a. rubiginosus .... 277
6. phillipsi 278
3. viverrinus 281
Page
4. Felis 285
1. eonstantina 286
a. ornata 287
2. chaus 290
a. chaus 292
6. afflnis 294
c. kutas 297
d. prateri 298
e. kelaarti 300
/. fulvidina 303
6. Caracal 306
1. caracal 306
a. caracal 307
7. Lynx 310
1. lynx 311
a. isabellirms 311
8. Otocolobus 315
1. manul 317
a. nigripectus .... 319
6. ferrugineus .... 319
Subfamily Acisosyckikjf. 322
1. Aeinonyx 323
1. jubatus 324
a. venaticus 325
Family 2. Vivebridje 330
Subfamily 1. Pmioxodoxtutm . . 332
1. Prionodon 334
1. pardicolor 337
2. linsang 339
Subfamily 2. Viterrixjb 342
1. Viverra 344
1. zibetha 346
a. zibetha 347
6. picta 350
o. pruinosa 352
2. Moschothera 354
1. megaspila 356
2. civettina 358
3. Viverrieula 362
1. indica 363
a. mayori 363
6. indica 364
c. bengalensis 367
d. deserti 368
e. wellsi 369
/. baptistse 370
g. thai 372
Subfamily 3. PAHADnxunm^ . , 376
1. Paradoxurus 379
1. zeylonensis 381
2. jerdoni 383
a. jerdoni 383
b. caniscus 386
SYSTEMATIC CONTENTS.
XXX111
Page
3. hennaphroditus 387
a. hennaphroditus. 388
6. niotitatans 392
c. scindise 393
d. Ianeus 394
e. vellerosus 397
/. bondar 398
g. pallasii 400
h. laotum 402
i. minor 403
j. senex 409
k. fusous 410
I. pallens 410
m. pugnax 410
n. sacer 410
o. puleher 410
2. Paguma 415
1. lanigera 416
2. larvata 417
o. -wroughtoni .... 418
6. grayi 420
c. negleeta 422
d. nigrioeps 424
e. tytlerii 424
/. mtrudens 425
g. robusta 426
h. janetta 427
3. Arctiotis 431
1. binturong 432
a. albifrons 433
6. binturong 435
Subfamily 4. Axctogalidiixjk . 439
1. Arctogalidia 441
1. trivirgata 444
a. leuootis 444
6. maora 446
c. millsi 447
Subfamily 5. HmuaAiixx .... 450
1. Hemigalus 452
1 . derbyanus 454
a. incursor 455
VOL. I.
MAMMALIA.
A class of vertebrated animals distinguished from the
Reptilia, its next of kin and ancestral stock, by a number of
characters of which the most important perhaps are the
presence of hairs, instead of scales, in the skin, and of mammary
or milk -glands in the female for the nourishment of the young
after birth. Other differences are the separation of the
cavity of the chest or thorax from that of the abdomen by
a complete respiratory muscular sheet or diaphragm, the
division of the heart into four chambers, and the presence
of a single great blood-vessel, the aorta or aortic arch, by
which the blood is driven from the heart to the various
organs ; these modifications of the respiratory and circulatory
apparatus being accompanied by warm blood, which normally
keeps the body-temperature at a uniform level and above
that of the surrounding medium, air or water, however cold.
In the skull the most important difference lies in the lower
jaw, which consists of a single bone jointed directly to the
skull, the subsidiary intervening bones, present in existing
reptiles, being taken into the ear, some as auditory ossicles
subservient to hearing.
To explain the technical terms used for defining the orders
and subordinate groups into which mammals have been
classified it is necessary to describe some of the external
and skeletal characters.
The normal hairy covering, occasionally represented to a
certain extent by spines, still more rarely by plate-like scales,
and lost in the Cetacea, usually consists of two elements,
the superficial (contour) hairs and the underwool. On the
head the hairs of certain definite areas are modified as tactile
bristles (vibrissas) arranged in tufts on the upper lip
(mystacial), over the eye (superciliary), on the cheek
(genal), on the. chin (submental), and on the fore- throat
(interramal). These are found in so many orders that they
may be regarded as primitive. Two additional, probably
primifcVe, organs on the head are (i) the area of moist naked
skin round the nostrils (rhinarium), which generally has
a narrow extension (philtrum) running to the edge of the
upper lip in front and dividing it into a right and left portion,
VOL. I. B
2 MAMMATIA.
and (ii) the external ear (pinna), a hollowed extension of the
skin supported by cartilage, and generally strengthened by
ridges or lobes of the same material. It varies greatly in size,,
shape, and other particulars, and may be altogether lost.
The edges of its cavity below typically meet to form a notch
(intertragal) ; just above this in front there is commonly
a lobe (tragus), and behind another lobe (antitragus) , and the
principal strengthening ridge (sujpratragus) lies longitudinally
above these within the cavity. Further modifications will
be described under the different headings where necessary.
Kg. }■—&■ Bight ear of Hunting Leopard : 6, bursa ; si, supratragus ,-
it, intratragus ; t, tragus ; in, intertragal notch. B. Bhinariuia
of Large Indian Civet, from the front : in, infranarial portion
beneath nostra ; ph, philtrum. C. Side view of head of Indian
Jackal, showing tufts of facial vibrissas : s, superciliary ; g 1 and g 2 ,
upper and lower genal ; to, mystacial ; sm, submental ; ir, inter-
ramal.
The modifications of the feet supply important systematic
characters. The primitive type of foot has five toes or
digits (pentadactyle), the inner digit being the first, the outer
the fifth. The terminal segment of each is provided above
with a compressed claw, a flattish nail or a hoof, an expanded
nail embracing the front and generally the sides of the segment.
The sole in the primitive foot is naked from the tips of the
digits to the wrist and ankle or heel, and is supplied with paad
On the digits the pads (digitals) are restricted to the terminal
segment. The rest of the pads, six in number, are on the
sole. Four of them (interdigitals) are just behind the
spaces between the digits. Frequently three of them, some-
times four, coalesce to form a large submedian pad (plantar).
The remaining two, an outer and an inner, are near the wrist
of the fore foot (carpal) and towards the heel of the bind
foot (metatarsal). When the sole is overgrown with hair the
pads may be wholly lost. The gait of a mammal walking
on the greater part of the sole is called plantigrade, on the
toes, with the heel and wrist well off the ground, digitigrade,
and on the hoofs alone, unguligrade ; but there is no sharply
defined difference between these methods.
A B
Fig. 2. — Lower surface of the feet of a Pouched Mouse (Phascogale),
an Australian marsupial mammal, to show the primitive arrange-
ment of the pads. A. Right fore foot, showing the digital pads
at the tips of the toes : 1 and 4, the internal and external inter-
digital pads ; c, the internal carpal pad ; cv, tuft of carpal vibrissae,
a primitive mammalian feature. B. The hind foot, with similar
pads ; mt, the external metatarsal,
The generative organs are also extensively used in classifica-
tion, but only recently to the extent their importance demands.
The area between them and the anus is the perinseum.
In the male the intromittent organ or penis frequently has
a thickened termination, the glans ; and it may be strengthened
by an internal, rod-shaped bone, the bacuhim. The sack
of skin into which the testes may descend is the scrotum.
In the female the generative orifice has tumid lips or labia,
b2
4 MAMMALIA.
and between these and above the urinary orifice is an organ,
the clitoris, which corresponds to the penis of the male.
In the skeleton the skull is the most important part. It
consists of a large number of bones, the lines of contact between
them being the sutures. As age advances and the bones fuse
the sutures tend to become obliterated. For convenience
the skull may be divided into two parts : (i) the facial, including
the muzzle and the eye-sockets or orbits, and (ii) the cranial,
which lodges the brain. The muzzle is made up of the lower
and upper jaws. The two halves of the lower jaw or mandible
meet in front in a symphysis ; behind they are jointed to the
skull by a condyle ; below the condyle there is usually a process,
the angular, and above it a wide flange, the coronoid, to which
Kg. 3. — Upper and lower sides of left half of skull of Indian Jackal,
■with names of bones, teeth, and other features referred to in text.
"Upper side : pmx, premaxilla ; nas, nasal ; max, maxilla ; mal,
malar bone of zygomatic aroh {zyg.); fr, frontal ; pop, frontal
postorbital process ; orb, orbit ; sq, squamosal branch of zygo-
matic arch ; par, parietal ; mast, mastoid ; oar, occipital crest ;
sag, sagittal crest ; tr, temporal ridge. Lower side : lettering as
above, with apf and ppf, anterior and posterior palatine foramina ;
pal, palatine ; m/, mesopterygoid fossa ; ham, namular process of
pterygoid; ptc, external pterygoid crest; bul, auditory bulla;
fm, foramen magnum ; oc, occipital eondyle ; poo, paroceipital
process ; ao, auditory orifice ; gl, glenoid ; fo, foramen ovale ;
afe, alisphenoid canal, marked by arrow; teeth: i, incisors; c, canine;
pm l to pm l , premolars ; m 1 , m\ molars.
MAMMALIA. 5
the great masticatory muscle, the temporal, is attached.
These are parts of the postdental portion of the mandible,
the rest being the dental or tooth- bearing portion in front.
The upper jaw contains a spacious cavity, opening in front
by the anterior wires and behind, in the back of the mouth,
by the posterior nares. The cavity is roofed above by the
nasals, enclosed at the sides by the maxillae, the anterior nares
being bordered laterally and below by the premaxillse. The
floor of the cavity, which also forms the roof of the mouth,
is composed of the premaxillse in front, the maxillse in the
middle, and the palatines behind. The palate is typically
pierced by two pairs of orifices, one, larger, in front ; the other,
smaller, behind, known respectively as the anterior and
posterior palatine foramina. The cavity of the nasal chamber
is largely occupied by two pairs of delicate, sponge-like bones,
the maxillo-turbinals in front and the ethmoturbinals behind.
On the facial part of the maxilla below and in front of the
orbit there is usually a well-defined orifice, the infraorbital
foramen, through which nerves and blood-vessels pass to the
upper lip. Usually within the front rim of the orbit is the
lacrimal foramen. From the cheek below the eye to the ear
runs a stout bar of bone, the zygomatic arch, the anterior
part of which, consisting of the malar bone, forms the lower
rim of the orbit, and its posterior part, composed of a
forwardly directed process from the squamosal, a bone forming
the wall of the lower part of the cranium above the ear,
forms the outer edge of a space, the temporal fossa, which
usually communicates with the orbit in front.
The cranial cavity is roofed in front by a pair of frontal
bones, each of which is usually produced at the side into a post-
orbital process, which sometimes meets a corresponding process
rising from the malar part of the zygomatic arch to complete
the ring round the orbit. Two roofing bones behind the frontals
are the parietals. These, with the frontals to a lesser extent,
give attachment to the masticatory temporal muscle on each
side, and the edge of the muscle is typically marked by the
temporal ridge, which extends from the postorbital process
to the back of the skull. Not infrequently the ridges creep up
to the summit of the skull with age and, coalescing, may rise
into a high sagittal crest to support the muscle. The back
of the skull is mainly formed by the occipital bone, which
encircles the orifice, foramen magnum, through which the spinal
cord passes, and on each side of this is the occipital condyle,
by which the skull is jointed to the neck. The upper part of
this bone is usually provided with a transverse curved occipital
crest for supporting the muscles of the neck. The floor of
the skull just in front of the foramen magnum is formed by
the basioccipital, and the space between this and the squamosal,
6
MAMMALIA.
to which the mandible is jointed, is occupied by bones connected
with the ear, usually a swelling, the auditory bulla, connected
with the auditory orifice, and behind this frequently a pro-
jection, the mastoid, and behind the bulla, or pressed against it,
is a process from the occipital, the paroccipital.
The base of the skull in front of the areas just described
and behind the palate is occupied by a pit, the mesopterygoid
fossa, into which the posterior nares open. The side walls of
this are formed by the palatine bones in front and the ptery-
goids behind, the latter sometimes running into a distinct
process, the hamular. In the base of the skull, adjoining,
there are typically four orifices for the exit of nerves from the
par
pop fr
Pig. 4. — Side view of skull, with mandible, of Indian Jackal. Upper
figure : lettering as in fig. 3, with iof, infraorbital foramen. Lower
figure (mandible) : cor, eoronoid process ; cond, condyle ; ang,
angular process of postdental portion ; i, incisors ; c, canine ;
pm x to %>m 4 , premolars ; m^ to ro s , molars.
brain, forming a series from the middle of the temporal fossa
backwards, and named the optic foramen, the sphenoidal fissure,
the foramen rotundum, and the last, the foramen ovale, a little
in front of the auditory bulla.
The teeth supply important characters for distinguishing
mammals of all ranks, at least from genera to orders. The
full complement of teeth in normal adult mammals is generally
considered to be 11 on each side above and below, differentiated
into four categories — 3 incisors, 1 canine, 4 premolars, and
3 molars, represented by the formula above and below for
MAMMAIJlA. 7
one side : i. | , c. j j pm. | , m. | . In only a few genera of
Oriental mammals is this number retained above and below,
the reduction affecting alike the teeth of all four categories.
In the upper jaw the incisors are the teeth present in the
premaxillse. The rest, from the canines backwards, are
lodged in the maxillse, and are sometimes collectively
called the " cheek-teeth." The premolars and molars cannot
always be distinguished by their form and structure. The
difference lies in the premolars, like the canines and incisors,
having predecessors in the milk dentition, the molars being
unrepresented in the first set. In the lower jaw the teeth are
developed in a single bone, and the incisors are identified by
their correspondence with those of the upper jaw. The
lower canine always passes or lies in front of the upper when
the jaw is closed. This position enables it to be distinguished
from the first premolar in some mammals, where the lower
-canine resembles the incisors and the first premolar is canine-
like. Similarly tusk-like teeth in the front of the upper jaw
are determined as incisors or canines in accordance with their
■emergence in the premaxdlla or maxilla.
A normal individual tooth consists of the root or roots
imbedded in the bone of the jaw and of the crown projecting
beyond it. It is developed from a pulp buried in the bone,
and during its growth the root remains open at its lower
end. In most cases the root closes and growth of the tooth
ceases by the time it is brought into use. Such teeth are
commonly short, and the condition is called brachyodont. But
sometimes the roots remain open and the tooth continues
to grow after it is in use, so that the effects of wear are made
good, at all events until old age is reached. Such teeth,
generally deeply imbedded and long, are called hypsodont.
The crowns are still more variable in structure. Like the roots,
they are composed mainly of " dentine," but they are typically
coated with a layer of hard enamel, and outside the enamel
there is in some cases a third substance, the cement.
These three successive layers are manifest in the worn molar
teeth of such animals as Elephants. The crowns of the teeth,
•especially of the posterior cheek-teeth, are typically provided
with definite cusps or tubercles varying in size and number,
and there is a theory, called the " tritubercular theory,'* by
no means always accepted, that in early mammals the upper
and lower teeth carried three cusps arranged, not in line, but
in a triangle, the upper having two outer and one inner cusp
and the lower two inner and one outer, so that they interlocked
when the mouth was closed. In the upper tooth the inner cusp
is called the protocone, the anterior of the two outers the
jparacone, and the posterior the metacone. In the lower tooth
the outer cusp is the protoconid, the anterior inner the par aconid,
8 MAMMALIA.
and the posterior inner the metaconid. These cusps can be
homologized in the teeth of many existing mammals ; but the
crown is generally complicated by accessory cusps which
may be developed, apparently, even on the cingulum, a thickened
ridge of enamel, often traceable, encircling the base of the crown.
Teeth indeed are extremely plastic organs, apparently very
responsive to diet, and may be totally dissimilar in mammals
otherwise tolerably closely resembling one another in structure * .
The rest of the skeleton may be briefly dismissed. The
vertebral column, or " backbone," extending from the front
of the neck to the tail- tip, and consisting of a series of bones
mostly freely jointed together, is divided into several regions.
The neck or cervical vertebras, seven in number in all Indian
mammals, have no ribs. A dozen or more of those that follow,
the dorsal vertebrae, carry ribs most of which are attached
below to a series of bones, the sternum. This region of the
body is the thorax. Behind the last rib-bearing vertebra
is a ribless series called the lumbar vertebras, which are
the only ones in the abdominal region. Then come a few
vertebrae welded together to form the sacrum, to which the
pelvis is attached ; the caudal vertebrae of the tail, varying
in number in accordance with the length of that organ, com-
plete the series.
To the scapula or shoulder-blade on each side of the fore part
of the thorax the fore limb is jointed, and in mammals
in which this limb is capable of movement in various directions
there is a collar-bone or clavicle, usually running from the lower
end of the scapula to the front of the sternum. The fore limb
consists of a single upper bone or humerus, jointed at its lower
end to two bones, an inner, the radius, and an outer, the ulna,,
the upper end of the latter forming the elbow prominence.
These bones at their lower ends are jointed to the wrist or
carpus, composed of two rows of small bones which give great
mobility to the fore foot. This primitively consisted of five
metacarpals jointed to the wrist and more or less welded together,
and five toes or digits, all of which, except the first, the pollex,
which has only two, are formed of three bones, the phalanges.
The bony structure, or pelvis, supporting the hind limb is
more complicated than that of the fore limb because it has
other functions. It consists of three bones, indistinguishably
welded in the adult, the upper, ilium, which is nearly
immovably attached to the sacrum, and two lower, the pubis
in front and the ischium behind. These two bones encircle
* The best instance of this, perhaps, is supplied by the Striped
Hyaena and the Aard "Wolf of Africa, which are beyond doubt nearly
related, yet have widely different premolar and molar teeth. The
Aard Wolf feeds mainly on white ants, whereas theHyasna feeds mainly
on dead mammals.
10 MAMMALIA.
a large orifice, the obturator foramen, and are not only joined
below it, but are almost always united to their fellows of the
opposite side, forming the symphysis pubis. At the point
where the three bones meet is the socket, or acetabulum, for
the hind limb, the individual bones of which correspond to
those of the fore limb ; the upper, or thigh bone, is the femur,
the two below it the tibia and fibula, representing respectively
the radius and ulna of the fore arm ; the ankle is the tarsus,
of which one of the bones, the calcaneum, forms the heel or
hock prominence, and the main body of the foot is composed
primitively of five metatarsals, followed by five toes or digits,
of which the first is called the hallux, with the same number
of phalanges as in the fore foot. In mammals with very
specialized limbs, like the Horses, Deer, and Antelopes, the
bones above mentioned, both above and below the carpus
and tarsus, may be profoundly modified.
From the primitive type, which may be pictured as a smallish,
normally hairy, long-tailed, short-limbed mammal, with penta-
dactyle, clawed, padded feet, a rhinarium, facial vibrissa,
and pinna, which had inherited from its reptilian ancestor
the capacity for running, climbing, and swimming, the mammals
in the course of their evolution have become adapted to living
under all the physical conditions the world supplies where
food was obtainable, apart from certain places, like New
Zealand, which the sea prevented the terrestrial species from
reaching. On land some have become specialized, for rapid
movement on the surface, others for burrowing beneath it ;
others, availing themselves of their climbing powers, took to
living in trees, and from arboreal species of different kinds were
evolved those with the capacity for flight, which in the case
of Bats is equal to that of birds. Prom the land, too, taking
advantage of the innate faculty for swimming, some invaded
streams, rivers, and lakes, and from aquatic species were
derived marine forms which, in the case of the Dugongs and
Whales, are as nearly completely fitted for life in the sea as the
fishes. But varied modes of life are not necessarily mutually
exclusive. In several of the more generalized mammals partly
terrestrial, partly aquatic, and partly arboreal habits are
combined.
Adaptation to the different environments above sketched
has been accompanied in mammals by a range of variation in
structure surpassing that of the existing mem'bers of the
other classes of Vertebrates. By their modifications they
have been classified into a number of different orders. Some
of these, like the Bats, Whales, Dugongs, and Elephants,
for instance, stand apart from the rest, isolated as living
animals by the extinction of ancestral linking forms. These
MAMMALIA. 11
orders are easy to define. But in other cases, owing to the
survival of comparatively primitive species, the ordinal
differences are not equally definite, and some of the differenti-
ating characters used in the following Key may seem
comparatively trivial. They are, however, accompanied
by others mentioned in the main text, where necessary.
It need only be added that, apart from the Monotremes of
Australia, the Marsupials of Australia and America, the
Edentates of America, the Hybacoidea and the TtrBUXi-
dentata (Aard Varks) of Africa, all the usually admitted
orders of Mammalia are found in British India.
Key to the Orders of Mammalia, based on those, excluding
Man, inhabiting British India.
a. Hind limbs present ; fore limbs not paddle-like ;
tail without terminal fin and usually much,
narrower than body at its base.
6. Digits provided with olaws or nails (absent on
some fingers in Bats).
c. Coat consisting of hairs or spines ; teeth
present.
d. Digits of fore limb not exceptionally
lengthened, and metacarpals not widely
separated to support a flight-membrane.
e. First digit of hind foot the largest,
opposable and prehensile, of the fore
foot (hand) sometimes the same, but
usually smaller PRIMATES.
e'. First digit of fore and hind foot, when
present, not modified for grasping.
/. Mouth not adapted for gnawing,
without a pair of specially modified
incisors above and below, and with no
infolded flap of hairy skin.
g. Body and limbs not provided with
a flight -membrane ; lower incisors
not comb-like.
h. Front teeth consisting of a pair of
tusk-like canines with a row of
typically 6 incisors between
them, of which the outer are
larger than the inner ; brain
more highly developed CARNTVORA.
h'. Front teeth very variable in
arrangement and size, some-
times approaching those of the
Carnivora, but the median in-
cisors larger than the laterals,
often very large ; brain of a
lower type INSECTIVORA.
g'. Body, limbs, and tail supporting
a flight-membrane ; lower incisors
comb-like DERMOFTERA.
/'. Mouth adapted for gnawing, with a
pair of large, median incisors above
and below and an infolded flap of
hairy skin RODBNTIA.
12
d. Digits of fore limb exceptionally long, and
metacarpals widely separated to wrist to
support flight-membrane CHIROPTERA*
c'. Hairs consolidated to form erectile plates or
scales ; no teeth PHOLIDOTA.
&'. Digits provided with hoofs.
i. Bars erect, a normal neck, at most a short
proboscis ; normal incisors present at least
in lower jaw.
k. Median axis of feet passing between 2nd
and 3rd digits, their hoofs equal and [DACTYLA.
symmetrically paired ARTIO-
k'. Median axis of feet passing through 3rd [DACTYLA.
digit, which is the largest PERISSO-
i'. Ears lying against side of neck, which is very
short ; nose and upper lip forming a long
prehensile proboscis ; front teeth repre-
sented at most by a pair of upper tusk-like
incisors PROBOSCIDEA.
a'. Hind limbs absent, fore limbs converted into
paddle-shaped flippers ; tail expanded at end
into a horizontal swimming-fin or "fluke,"
and as thick as the adjoining part of the body at
its base.
Z. Lips tumid, mobile, and prehensile, the upper
cleft ; nostrils on summit of muzzle ; back
teeth with flat crowns ; mammaa pectoral . . . SIRENIA.
V. Lips not noticeably tumid, mobile or prehensile,
the upper not cleft ; nostrils, except in the
Cachalots, on summit of head ; back teeth,
when present, with conical crowns ; mammse
inguinal CETACEA.
The increase in the number of orders over those admitted
by Blanford and the change in some of the names are due to
the dismemberment of the Colugos as Dermoptera from the
Insectivora and to the substitution for his Ungtjlata of the
three orders Aetiodactyla for the "even-toed" or "cloven-
footed " mammals, of Perissodactyla for the " odd-toed "
hoofed mammals, and of Proboscidea for the Elephants.
This is in accordance with, recent opinion. Also it is now
admitted that the Scaly Anteaters or Pangolins of Africa and
Asia are not related to the Anteaters of S. America, which,
with the Sloths and Armadillos, compose the true Edentata.
The resemblance is merely a matter of diet-adaptation. Th&
Pangolins are now called, the Pholidota. Another order,
Menotypbxa, might have been introduced by separating the
Tree-Shrews from the Insectivora.
Order PRIMATES.
As stated in the analytical Key, this order is mainly dis-
tinguished from the others constituting the mammalian fauna
of British India by the large size, freedom of movement, and
opposability of the first digit of the foot, the hallux or " great
toe," so that this extremity forms a powerful grasping organ
subservient to climbing*. In the hand the first digit, the
pollex or " thumb," may be equally large and similarly
functional, as in the Lorises, but in the Apes and Monkeys
it is not so large as the other digits, and its opposability is
useful rather for picking up food than for climbing. Asso-
ciated with the freedom of movement of the arms and legs
for climbing or other purposes are certain skeletal characters,
namely, the presence of complete collar-bones, or clavicles,
and the distinctness of the two bones of the forearm,
radius, and ulna, and of the shin, the tibia, and fibula. The
teeth do not exceed 36 in number, the formula of the com-
plete set being: i. £, c. \, pm. 3, m. 3 . There are always
two pairs of incisors above and below, and the medians
are larger than the laterals. The upper canines are always
well developed and tusk-like, at least in the male, and the lower
also in the Apes and Monkeys, but not in the Lemuroids.
The first lower premolar is larger than the rest and acts as
a sharpener to the bind edge of the upper canine, and in the
Lemuroids is itself sharp and canine-like. The back cheek-
teeth have broad crushing crowns, with four cusps, and are
more complex than those in front behind the canines.
In his classification of the Mammalia, Man very naturally
gives pride of place to this order containing himself and his
next of kin, as indicated by its name. This position is only
justified by the brain development and the intelligence that
goes with it of some of the most specialized of its members.
But this does not apply to the more " bestial " forms like the
Lemuroids, and the organization of the order as a whole is
much less specialized than in several others.
* Only one other order has a similar type of foot, namely, the
Makstjpialia of Australia and America. I long ago suggested the
possibility of this resemblance being of genetic importance. As adaptive
modifications to special methods of climbing the hallux in the Pecmates
is sometimes reduced in size. It is comparatively small, for instance,
in the Orang-utan; and in the diminutive S. American Marmosets,
which leap about trees like Squirrels, it is so small as to be nearly
funetionless, although still opposable.
14 PITHECOIDEA.
Key to the principal Subdivisions of
British Indian Primates.
a. Muzzle without rhinarium and with lips free
from the gum and protrusible ; skull with
orbit closed behind, and the front tsetli normal [p. 14.
in position and function PITHECOIDEA,
6. Skull with bony tubular auditory meatus ;
2 premolars above and below on each side . . Catabbixi, p. 15.
c. Arms much longer than the legs ; no [p. IS.
external trace of a tail Anthropomorpha.
c'. Arms and legs subequal in length; tail
usually long, sometimes reduced to a [p. 31.
stump" Cynomorpha,
b'. Skull without tubular auditory mea,tus ;
3 premolars above and below on each side . . Pzatimezxz.
a'. Muzzle with distinct rhinarium and with lips
adherent to the gum, not protrusible ; skull
with orbit open behind and front teeth [p. 163.
abnormal LEMUROIDEA,
The characters of the family Hominidse (Man), belonging
to the Anthropomorpha, are not included in this Key. The
characters of the Platyrbdqti, containing the American
Monkeys, are entered by way of contrast with, the Catarhint,
but the group is not included in the Indian fauna.
Suborder PITHECOIDEA *.
No rhinarium or area of moist skin round the nostrils,
with its extension below, as the philtrum, to the edge of the
upper lip ; the lip not closely adherent to the gum, but
protrusible.
Hands and feet with the 4th digit not longer than the 3rd,
which, more noticeably on the hand, is typically longer than
the 2nd and 4th. Foot with its 2nd digit never differentiated
from the rest by its claw-like nail. The tongue without
a sublingua or serrated lamina beneath it.
Skull with the orbit shut off from the temporal fossa by
a bony plate, leaving merely a narrow communicating cleft
below it. Lower canine not projecting forwards and ineisi-
form, the lower incisors not forming a comb-like structure
and the median upper incisors not widely separated in the
middle line.
This suborder, with the Tarsioidea, constitutes the section
of Haplorhine Primates, characterized by the structure of the
* In 1918 I proposed this subordinal name as a substitute for the
older name Anthropoidea because "Anthropoid," reasonably according
to its meaning, has been consistently applied to the man-like Apes,
and because "Anthropoid" cannot with any approach to its real
significance be applied to, e. jr., a Marmoset. But a Marmoset and a man
are alike Pithecoid or monkey-like.
CATABHIN1. 15
nose, and upper lip, by the closed orbit, and other specialized
features distinguishing them from the more primitive Strepsi-
rhine Lemurs (p. 164). Tarsius, however, which does not
occur west of Java, shows several superficial resemblances to
the African Galagos, related to the Lorises (p. 164), and was
formerly always regarded as a Lemuroid. Its survival as
a representative of the primitive stock of Pithecoids, linking
them in a measure with the Lemuroids, justifies the retention
of the Lemuroidea in the Primates instead of their
relegation to a special order, the PnosiMiiE.
The Pithecoid Primates are divisible into two groups,
the Platyrhint or American Monkeys and Marmosets, and
the Catarhtnt or Monkeys and Apes of Africa and Asia and
Man.
CATARHINI.
Distinguished from the Platyrhbsti by the presence of
a bony tube to the ear and by having only two premolar
teeth above and below on each side, the dental formula being :
i. | , e. | , pm. | , m. |== 32, instead of i. | , c. \ , pm. | ,
m. | or |= 36 or 32, the numerical equality in the latter case
being reached by the loss of the 3rd molar in the Marmosets.
In the Catarhest also the periotic bone is always perforated
by the carotid canal, marked by a conspicuous foramen,
whereas in the Platyrhtjst the artery usually enters the brain
behind the periotic. The external differences are not so
important, but the thumb in the Catarhines is more opposable
and prehensile and the nostrils are typically narrower and more
convergent at their lower ends than in the Platyrhines, which
take that name from the normally wider space between the
nostrils.
Section ANTHROPOMORPHA.
The Man-like Apes.
This section of the Catarhine Primates, comprising the
species to which the term " Apes " is now conventionally
restricted, is distinguished by the absence of all external
trace of the tail, by the great development of the arms, which
are much longer than the legs, by the thumb being more
freely jointed to the hand and emerging close to the wrist,
with its basal thickening, the " ball of the thumb," approxi-
mately on a level with the external metacarpal pad ; the
foot has its heel broader in proportion to its length, the
hands and feet are never markedly unequal in length, and the
pads on the palms and soles are poorly developed.
16
PITHECOIDEA.
These characters are associated with marked peculiarities
hoth in arboreal and terrestrial activity. In walking in the
•quadrupedal attitude the palms of the hands are never applied
to the ground, the bent knuckles of the fingers being used
instead, and the entire sole of the foot, or its outer edge,
supports the hind quarters, which, as compared with the
Fig. 6.' — A. Right band of Lar Gibbon (Hylobates lar).
B. Right foot of the same.
shoulders, are weak and light. In the young at least the
■erect bipedal attitude is readily assumed for standing, walking,
■or even running. In tree-climbing, when speed is required,
the powerful arms are principally employed for swinging
from branch to branch, the legs being too weak for long
leaps. This method of climbing has been called brachiation.
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HYLOBATEDJE. 17
To the Astthbopomobpha belong the Gorilla and Chimpanzee
of Africa and the Orang-utan and Gibbons of the Oriental
Region. They are essentially forest animals. The Gibbons
alone, constituting the family Hylobatidse, occur in the
British Indian fauna *.
Family HYLOBATIDSE.
The Gibbons.
Apes with ischial callosities invariably present and well
developed, the arms and legs, especially the arms, relatively
long, with long and narrow hands and feet and long, slender
digits, the pollex, or thumb, very long and, when closed,
fitting into a depression on the adjoining surface of the
hand, the hallux, or " great toe," also long and, when closed,
overlapping the distal margin of the sole of the foot. No
striking secondary sexual characters, the two sexes being to
all intents and purposes alike in size and skull characters,
the $ having enlarged canines, which at most are only a
little smaller than those of the <J. The skull has a larger
cranial and smaller facial portion, with weaker jaws, than in
Asiatic Catarhine monkeys.
In the almost complete absence of secondary sexual charac-
ters the Gibbons differ from all the other families of Catarhine
Primates.
Although less man-like than the other Anthropoid Apes
in mental capacity, and in some, but by no means all, structural
characters, thus coming nearer the ordinary Monkeys, the
Gibbons surpass the other Apes in physical activities, mani-
fested by their ability to walk and run with facility in an
erect attitude on the ground and by the extent to which they
have perfected the arm-swinging method of traversing the
forest. In these and other habits all the species of Gibbons
seem to be alike or to differ only in minor particulars.
On the ground they are essentially bipeds, standing and
walking on the flat sole of the foot, with the great toe pro-
jecting sideways as a support, and, when hurried, either
running with considerable speed, although with a somewhat
awkward gait and not so fast as man, or progressing with
a series of leaps, the two legs acting in unison. When thus
* Man also, forming the family Hominidas, belongs to this section.
He is chiefly distinguished from the Apes by the perfection of the
bipedal attitude, which has brought with it some modifications of struc-
ture, of which the most important is the tying up of the great toe to the
side of the foot so that it has lost its opposability to the others and
is no longer, strictly speaking, prehensile.
VOL. I. C
18 HYLOBATID-a;.
on the move they may lightly touch the ground with the
fingers to steady themselves ; but almost always they hold
the arms up, outstretched, but bent at the elbows and wrist,
and employ them as balancers. So accurate is the balance
that Gibbons can walk along the branch of a tree or, as
observed in captive specimens, even along a swaying, hori-
zontal rope, grasping it with their prehensile feet.
Equally remarkable are the means they employ and the
speed they attain when traversing forest trees. With a jump
off the upper side of a branch they launch themselves towards
•another, grasp it with upstretched hand, swing beneath it
and, letting go, are carried on to another, which is similarly
grasped by the other hand, the action being continued from
branch to branch at great speed. When arresting their
course they retain the hold of the branch and, swinging up
on its further side, alight on their feet on its upper surface.
Naturally they also climb slowly about trees after the manner
of other Anthropoid Apes, but the method above described
is the method adopted when speed is required. No other
Pithecoid Primate is capable of such arboreal acrobatic feats,
and none but man can exhibit such terrestrial bipedal
activity.
Another habit they have in common is howling vociferously
from the tree-tops for two or three hours after sunrise, then
quietly resting or feeding till towards sundown, when a briefer
concert is indulged in before retiring to rest on the branches.
Their diet consists mainly of leaves, flowers, and fruits,
varied with some spiders and insects ; and their usual method
of drinking from a pool is to dip the back of the hand in the
water and lick it off with the tongue, but occasionally they
suck up the liquid direct with their lips.
The period of gestation is seven months (see p. 31).
Mnal agreement as to the value of the characters by which
the various kinds of Gibbons that have been named differ
from each other has not yet been reached by systematists.
But the latest classification of the family by Miller (Journ.
Mamtn. xiv, p. 159, 1933) establishes the existence of at all
©vents five well-marked species which, so far as is known,
do not intergrade, and are given subgeneric rank. These are :
H. lar and H. hoolock, representing Hylobates (sensu stricto) ;
H. Uucogenys, subgenus Nomasous, from Siam and Indo-
China ; H. klossii, subgenus Brachitanytes, the dwarf Gibbon
of S. Pagi Island, west coast of Sumatra ; and H . syndactylus,
subgenus Synvphalangus, the Siamang of Sumatra and the
Malay Peninsula.
The two British Indian species belong to Hylobates (s. s.).
HYLOBATES. 19
Genus HYLOBATES Illiger.
Bylobates, Illiger, Prodr. Syst. Hamm. p. 67, 1811 ; Miller, Journ.
Mamm. xiv, p. 159, 1933 (sensu stricto as subgenus).
Type of the genus, Homo lar Linn.
Distribution. — From Assam and Burma to Siam and
Indo-China, and through the Malay States and Sunda Islands
to Borneo.
Distinguished from the species constituting the subgenera
Nomasous, Brachitanytes, and Symphalangus, as defined by
Miller, by one or more of the following characters : — Hair
on the crown directed backwards, longer laterally than mesially ;
laryngeal sae absent in both sexes and the throat not noticeably
thinly haired ; the scrotum absent, the testicles in the adult
only descending to the base of the penis, but not below it.
Skull with its facial profile tolerably strongly concave between
the top of the nasal and the summit of the prominent brow-
ridge*.
This diagnosis applies to the two British Indian species
and to the three forms agilis, leicciscics, and inulleri from the
Malay Peninsula, Sumatra, Java, and Borneo, which, with
skulls like those of lar but differing in coloration, I regarded
as subspecies of it (Proc. Zool. Soc. x. 1927, p. 722). Later,
however, Kloss (Proc. Zool. Soc. 1929, p. 117) gave reasons
for considering them specifically distinct, an opinion apparently
endorsed by Miller.
The two species occurring in British India may be briefly
distinguished as follows : —
a. Hands and feet the same colour as the aims and
legs ; black phase with a pale brow-band, but
the face not encircled with white ; pale phase [p. 19.
with the ventral surface darker than the dorsal . . hoolock (Harl.),
b. Hands and feet wholly or mostly -white above, very
conspicuously contrasted with the arms and legs
in the dark phase and always a little lighter in the
pale phase : face encircled with a white ring in the
black phase ; the ventral surface about the same [p. 26.
tint as the dorsal in the pale phase lar (Linn.),
1. Hyloljates hoolock (Harlan). The Hoolock Gibbon.
Simla hoolock, Harlan, Tr. Amer. Phil. Soe. iv, p. 52, pi. ii, 1834,
and of all recent authors under Hylobates.
Hylobates fuseus, Winslow Lewis, Journ. Nat. Hist. Soc. Bost. i, pt. 1,
p. 32, pis. i&ii, 1834.
Hylobates choromandus, Ogilby, Proc. Zool. Soc. 1837, p. 689.
Hylobates scyritus, Ogilby, Boyle's Illustr. Himal. Bot. p. ix, 1839.
Vernacular. — UlvJc (Hindi) ; MyouJc-lwai-gyau and Too-
boung (Arakan) ; Wu-wa (Shan).
* Miller mentioned as an additional diagnostic character the direction
of the hairs on the ladial side of the forearm towards the thumb. The
two British Indian species differ in this respect.
c2
20 HYLOBATID-ffii.
Locality of the type of hoolock, Garo Hills, Assam ; of
fuscus, ""the vicinity of the Himalaya Mountains " ; of
choromandus, unknown ; of scyritus, Assam *.
Distribution. — From Assam, Cachar, and Chittagong
through Upper Burma to Western Yunnan and the North
Shan States.
Coat long and thick, smooth, not raised into a tuft on the
crown of the head, and abundant round the callosities and the
genital area. Hairs on the forearm growing upwards from the
wrist to the elbow. Colour varying, sexually dimorphic in
the adult ; the male is black, with a contrasted silvery- white
brow- band and a brown or grey beard-like tuft on the genital
area ; the female is more variable and much paler, the entire
upper side and the outside of the limbs may be tolerably
uniformly golden or ruddy- buff, or brownish-buff clouded with
brown on the back or buffy-white on the head, nape, fore part
of the back, shoulders, and arms, becoming clouded with brown
on the flanks, loins, and legs ; the hands and feet may be
the same tint as the arms and legs, but usually the digits,
or at least their distal ends, are black ; the face is encircled
by white hairs, with a narrow line of white between and below
the eyes ; the ventral surface, the middle line of the throat,
and the cheeks up to the ears are darker or lighter brown,
always darker than the dorsal surface, and the brown hue
of the cheeks, often very dark, emphasises the pale band round
the face ; the hairs of the genital region just below the vulva
are brown or black, always darker than the surrounding
hairs.
There has been great difference of opinion about the incidence
of the black and pale phases in the two sexes. G. M. Allen
(Amer. Mus. Novit. no. 429, p. 5, 1930) suggested that the
species goes through colour- changes similar to those known
to occur in the Indo-Chinese species H. (Nomascus) concolor,
which I described many years ago (Proe. Zool. Soc. 1905,
p. 169). This suggestion was proved correct by McCann
(Journ. Bomb. Nat. Hist. Soc. xxxvi. p. 395, 1933), who,
from his observations in the Naga Hills, found that the young
at birth is pale greyish- white, with a yellowish tinge, gradually
darkens with age, and finally turns black before reaching
maturity. This black hue is retained throughout life by the
<J ; but in the $ at puberty the coat fades to the pale yellowish-
brown hue. McCann's observations on the colour-change in
the $> Hoolock were subsequently confirmed by Major W. B.
Shakespear (Journ. Bomb. Nat. Hist. Soc. xxxvii. p. 214,,
1934). A specimen which was at first grey gradually changed
to jet-black, and stayed that colour until about 5| years old,
* The types of choromandu.* and scyritus in the British Museum are
females in the pale phase.
HYLOBATES. 21
"when she began to turn grey and became greyish all over.
The largest black 2 I have seen had the head and body 20 in.
long, the foot 5|- in., and weighed 8 lb. ; but in the skull
the milk-canines were still in place and the last molar not
erupted.
Some individual differences in colour may be mentioned.
In three adult <£<£ from H'Kamti, Upper Burma, the chin
may be pale buffy, matching the brow-band, and strongly
contrasted with the surrounding black or dark brown or
uncontrasted. The hairs of the genital tuft, from 50 to 55 mm.
long, similarly vary from being black throughout, black
at the base, and brown distally or uniformly pale greyish-
brown and conspicuous from its generally pale hue. In a $
from Hatikhali, in the Cachar Hills, there is no white on the
chin and the hairs of the genital tuft, 58 mm. long, are brown
distally ; in one from the Kabaw Valley, Upper Chindwin,
to the west of the river, the chin is also dark, but the genital
hairs are very deep blackish-brown and aboat 45 mm. long.
But in a <J from Homalin, Upper Chindwin, on the east bank
of the river, there is some white on the chin and the genital
hairs, measuring 75 mm., are blackish at the base and largely
grey distally, and in one from Gokteik, in the N. Shan States,
the chin is brown, the genital hairs, 85 mm. long, are black
at the base, brown in the middle, and grey at the tip. The
difference in the colour of the genital hairs is probably a question
of bleaching from blackish-brown to reddish, and finally to
grey. In the 5 the prevalent colour, as shown by skins from
H'Kamti, the Kabaw Valley, and the Chin Hills, is mainly
shining buff above, brown below and on the cheeks. One
from Margharita, in the Naga Hills, stands out for the very
rich orange-ochreous hue of the upper side and the outside of
the limbs. Very different is a $ from Barahapjan, Lak-
himpur, which is not so richly coloured as the skin from
H'Kamti, being browner on the back and greyer on the
shoulders, head, and outside of the limbs.
The flesh -measurements (in English inches) and the weights
(in lb.) of some specimens are as follows : —
Head and
Locality and sex. body. Foot. Weight.
Hatikholi, Cachar Hills ; ad. <J 24$- 6i 17J
Homalin, Chindwin ; ad. <J 24* 6+ 16
H'Kamti, Chindwin ; ad. <J 2%\ 6 13*
Gokteik, N. Shan States ; ad. <J ..... . 23 6J 14*
H'Kamti, Chindwin ; ad. 2U 6 14A
Gokteik, N. Shan States ; ad. $..... . 23 6— —
Margharita, Naga Hills ; ad. ° 20 5* 14
Kulbi, Kamrup ; ad. " 18 6 —
Although the sexes overlap in size, the <J<J from these
data are a little larger on the average than the 22.
22
HYIiOBATIMI.
The skull varies individually in the height and length
of the cranial portion, the development of the brow-ridges,
which thicken with age, in the size and shape of the orbits
and of the anterior narial aperture, and in other particulars.
Kg. 7. — A. Skull of adult S Hoolook Gibbon (Hylobates haolock) from
the Upper Chindwin. B. Skull of adult <J Lar Gibbon (Hylobates
lar) from Bankaohon, Tenasserim.
As the following table of measurements shows, $ skulls are on
the average a little larger than $ skulls, and thus bear out
the conclusion established by the flesh-measurements of the
HYLOBATES.
23
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24 HYIiOBATIDJE.
slight average superiority of the $. The points in -which the
skull differs from that of H. lar, the other British Indian
species, are mentioned under that heading.
Distribution. — Pemberton's record of this Gibbon in Bhutan,
rejected by Blanford, and not subsequently confirmed, may
perhaps be true, since the Survey secured it in Lakhimpur to
the north of the Brahmaputra. In the Mishmi, Naga, Garo,
and other hill-ranges on the other side of this river it seems
to be plentiful, and west of the Chindwin, in the Kabaw
Valley and Chin Hills, it certainly occurs. But how much
farther south it goes is a little uncertain. Tickell, it is
true, said it occurs in Arakan, but he thought the Gibbon
he called the Hoolock from this district was different from
the Hoolock of Assam. In the H'Kamti State, Upper Burma,
it is equally plentiful on both banks of the Chindwin
according to Shortridge, who states that it must have
crossed to the eastern side by way of the Hukawng Valley,
above the source of the river, but that the comparatively
fiat country on the east bank, S. of H'Kamti, was unfavour-
able to its extension farther south. East of the Irrawaddy
it is found in the Kakhyen Hills, Western Yunnan, and in the
Hsipaw and N. Shan States *.
Habits. — Anderson wrote of this Gibbon as occurring in ' ' large
troops " in the dense forests on each side of the trawaddy
below Bhamo, where in the early morning the surrounding
hills echoed their howling. In the Kakhyen Hills he found
that they slept at night in the sheltered warm valleys and at
daybreak ascended the hills to a height of about 4,000 ft.,
traversing the forest with wonderful rapidity, uttering the
while their characteristic cry " Whoko ! Whoko ! " Their
diet consisted of the " essential elements of sylvian life " :
leaves, flowers, insects, spiders, eggs, and birds. From the
eagerness and method with which captive specimens in
Calcutta caught and devoured small living birds, Anderson
inferred that the Hoolock " must be a scourge to the
feathery tribe." Candler, however (Proc. Zool. Soc. 1906,
p. 187), who observed this Ape in Cachar, differs from
Anderson with respect to its diet. He thinks they feed
mainly on fruits and the succulent shoots of young bamboos
and other trees, and they will certainly eat some spiders.
But his captive specimens refused eggs and showed no
partiality for insects or small birds, merely pulling them to
pieces and possibly tasting themf.
t Anderson's record of it from Pegu was, I suspect, due to his inability
♦o distinguish it in the pale phase from H. lar.
* My own experience with Hooloeks in the London Zoological Gardens
bears out Candler's opinion. No doubt, however, they differ individu-
ally in tastes.
HYLOBATES. 25
Nor is Anderson's statement that they occur in large troops
borne out by subsequent observers. Mackenzie, who came
across them in the heavier jungles of the Chin Hills up to
5,000 ft. and in the Kabaw Valley, states that they usually
go about in parties of three or four, but sometimes singly;
once only did he notice a party of as many as sis, and in the
Hsipaw State Shortridge observed them in pairs or singly,
although he was told that where plentiful they are gregarious.
It is quite possible that their habits in this respect vary
locally and that the usual family parties may at times join
forces.
McCann (Journ. Bomb. Nat. Hist. Soc. xxxvi. p. 395, 1933)
has given a tolerably full account of the habits of this Gibbon
in the Naga Hills. He confirms the statements of Shortridge
and Mackenzie that it goes about in small family parties
consisting of the two parents and one or more young, the largest
party seen being composed of seven, of which the youngest
was a baby. He thinks there is a definite breeding season ;
and since every mature $ shot in January and early March
had a young one, he infers that the young are born in the
cold weather between November and February. He also
believes that maturity is reached in four or five, possibly
six, years, and that the young adults then leave the party
or are driven from it by the parents.
They spend most of the day in high trees, and in the early
mornings may often be seen sunning themselves on exposed
boughs, but during the hottest part of the day they frequently
descend to lower levels of the jungle. When alarmed they
.hide in the foliage, in the forks of branches or in bamboo
clumps, and their usual alarm-call is a sharp, rather harsh
bark, often repeated ; but alarm is also sometimes expressed
by a sound like a loud belch repeated from time to time.
When sleeping they typically assume a sitting posture on
a branch, with the head buried between the knees.
Their diet consists of leaves, flowers, and fruits, varied
with spiders and certain edible insects. Water for drinking
is usually taken off the foliage either direct with the tongue
or by wetting and licking the hands. In dense forests they
seldom come to the ground ; but now and again they leave
the trees and enter villages to plunder cultivated gardens,
being particularly fond of the flowers of Bombax.
Over the ground they move with exceeding rapidity for
a short distance in the typical Gibbon fashion described
above, but stop from time to time to rest.
26 HYLOBATID.ffi.
2. Hylobates lar (Linnaeus). The Lar or White-handed Gibbon.
Homo lar, Linn., Mant. Plant., App. p. 521, 1771.
Hylobates lar, Illiger, Abhand. Akad. Wiss. Berl. 181S, p. 88, and
of all recent authors.
Locality of the type, the southern portion of the Malay-
Peninsula (restricted by Kloss in 1929).
Distribution. — The Malay Peninsula northwards through
Tenasserim and the adjoining district of Siam into Lower
Bttrma.
Distinguished from B. hoohck by its slightly smaller size,,
by the hairs on the radial (inner) side of the forearm growing
downwards from the elbow to the thumb, upwards from
the wrist to the elbow externally, by having the entire hands
and feet, or at least the fingers and toes, white and contrasted
with the arms and legs, and by the absence of sexual dimorphism
in colour between the adult <J and $, which may be either
blackish or varying shades of brown, tawny buff or even
cream ; but the black phase is distinguished from that of
H. hoohck not only by the whiteness of the extremities, but
also by a conspicuous white band encircling the face. In
the pale phases these white areas are not so conspicuous,
although always visible, and the facial band is seldom thrown
into relief by the brown cheeks ; nor is the lower side noticeably
darker and browner than the upper, as in the adult $ of
H. hoohck. The rump, with its callosities, is not so concealed
by hair, and the genital tuft of the adult <J is much smaller
than in H. hoohck*. The skull is a little smaller, but has
relatively wider orbits, with the brow-ridges better developed.
at their inner ends, the upper jaw is weaker and the teeth
a little smaller (fig. 7, p. 22).
2 a. Hylobates lar entelloides Geoffroy.
Hylobates entelloides, I. Geoffrey, 0. E. Acad. Sei. Paiis, xv, p. 717,
1842 ; Arch. Mus. ii, p. 532, pi. xxix, 1843.
Hylobates lar entelloides, Kloss, Proc. Zool. Soc. 1929, p. 117.
Vernacular.— Myowk Hlwe giaw (Burmese) ; Khayoopathdn t
black phase, Khayoo pawd, pale phase (Karen) ; Ungta puttee,
pale phase, Ungta etam, black phase, Wow wow (Malay), Mawa,
(Malay at Bankachon).
Locality of the type, Malay Peninsula about lat. 12° N.
Distribution.— Northern part of the Malay Peninsula,,
Tenasserim, S.W. Siam.
* This description of the external features applies to the typical
H. lar lar from Southern Malaya and to its northern representative
in Lower Burma, but not to H. lar pUeatus of Siam and Cambodia,
which intergrades with the Burmese form.
HYLOBATES. 27
Distinguished, at least on the average, from typical lar from
the southern part of the Malay Peninsula by having the hands
and feet less extensively white by encroachment of the black
of the arms and legs below the wrist and ankle and by being
blacker and less brown on the body in the dark phase. In the
pale phase the tops of the hands and feet look soiled.
Geoffroy's type was an adult $ in the pale phase. His
coloured plate, if accurate in this respect, shows that the feet
at least were wholly white above up to the ankle-joint. The
locality of this Gibbon is about the same latitude as Tenasserim
Town, considerably to the north of Bankachon ; and in 1927 in
my paper dealing with this species of Gibbon special attention
was drawn to two <J specimens from the last-named locality,
close to Victoria Point, Tenasserim, on account of the blackness
of the wrists and ankles and the infuscation of the upper sides
of the hands and feet, serving to link in this respect typical
lar with the race H. lar pileatus which inhabits Siam and
Cambodia. On revising his abundant material of the species
two years later Kloss found that this coloration of the hands
and feet, accompanied by the general blacker hue of the back,
served to distinguish racially Tenasserim specimens from those
found farther south, and he adopted entelloides to designate
them.
In a series of skins from Bankachon, Victoria Point, Tenas-
serim (Hume and Shortridge), the darkening of the hands and
feet is very variable. In a rufous- brown $ the hairs on the
fingers and toes and some on the top of the hands and feet
are brown. In others the hands and feet may be dusky,
greyish-brown above to the base of the fingers and toes.
Even in those with the whitest hands and feet the dark hue
of the arms and legs extends practically down to the naked
skin of the palm and sole behind and to a corresponding
distance in front. But in a $ skin from Tavoy (Smith) the
hands and feet are as white as in typical lar, the black of the
limbs stopping short about an inch above the naked skin of
the palm and sole behind and to a corresponding distance in
front. In its general hue, however, this specimen is not so
brown as in typical lar, the back being black and extensively
grey on the shoulders.
From available material it appears the percentage of skins
in the black phase is less in Southern than in Northern Tenas-
serim. Thus in 22 skins from Bankachon, 4 <J<J are black,
11 &J and 7 $$ are pale. On the other hand, 1 $ from
Tavoy is black ; in 4 from the Taok Plateau (Vernay), 1 <J
and 2 $$ are black, 1 $ pale ; in 2 £,$ from 17 miles east of
Lakya (Vernay) one is black, the other pale, and 1 $ from
the Haungtharaw River, Kawkareik (Davison), is pale. Of the
28 HYLOBATIDiE.
8 skins from these localities, 5 are black and 3 pale, 2 $£
being black, 2 pale, and 3 $$ black and 1 pale. In Amherst,
however, Tickell says pale specimens were more numerous
than black.
The only newly-born young I have seen was collected by
Shortridge at Bankachon. It is darkish brown in colour,
paler than the normal dark phase, and darker than the pale
phase of the adults. A rather older black young one from
Amherst was figured by Tickell ; and two half-grown specimens
from Bankachon are in the normal pale phase.
The extremes in size and weight are supplied by the long
series of skins from Bankachon. The largest and smallest
$ and $ are entered below, with the average length of the
head and body. The average of the $ includes three from
the Taok Plateau, which fit in with the Bankachon females.
Intermediates between the extreme weights quoted have been
recorded.
Head and
Locality and sex. body. Foot. Weight.
Bankachon ; ad. $ 23 5fc 16f
Bankachon ; ad. <J 21J 6 13}
Average of 10 ; ad. <J 22+ — —
Bankachon ; ad. °. 22£ 6 15
Bankachon ; ad. $ 20 5f 10
Average of 7 ; ad. $ 21 \ — —
The sud joined table of skull-measurements shows that
although the total and mandibular lengths of the largest
may equal those of the smallest skulls of H. hoolock, the
averages are decidedly less. On the other hand the lengths
of the condylobase and of the upper cheek-teeth are in all cases
less. Also it seems evident that the skulls of H. lar, both
<? and o, are individually much more variable than of H. hooloch
despite the comparatively restricted geographical range of
E. lar entelloides.
The young are born in the early part of the cold weather,
as in the case of the Hoolock, and are dependent on the mother
for seven months.
Habits. — This Gibbon, Tickell tells us, is found in great ab_ m-
dance in all the forests skirting the hills of Tenasserim,but not
above 3,500 ft. It habitually lives in troops of from eight or
ten to twenty individuals of all ages and both sexes. Only
occasionally is an old male seen by itself. In indulging in
early morning concerts in the tree-tops, one troop calling
to another in a distant part of the forest, and in remaining
silent, feeding and resting, during the greater part of the day
afterwards, also in its instinct to hide in the foliage or behind
a thick branch when danger threatens, it resembles the
Hoolock ; but according to Tickell it is much less active on
HYLOBATES.
29
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HYLOBATIDiE.
the ground than that species and can only shuffle along on its
legs in a half-erect posture, using its arms for support*.
According to Shortridge this Gibbon is plentiful near
Victoria Point, where, although not avoiding the neighbourhood
of human habitations, it seldom leaves the forests and is
probably rarely destructive to ground crops. It is particularly
noisy in the early morning and evening, the call consisting
^gs&'lti'
it. ;*t =r_ „. >*
Pig. 8. — Lar Gibbon (Hylobates lar entelloid&s), black and pale phase,
showing method of climbing by brachiation.
of a series of rapid whoops ascending and descending through
several octaves, the sound, although somewhat resembling
that of a siren, being one of the most musical and striking
* This statement must be taken with reservation. I have seen
Lar Gibbons in the Zoological Gardens that ran and walked erect with
uplifted arms, and differed not at all from Hoolocks infthose respects.
HYIiOBATES. 31
to be heard in tropical jungles. He confirms Tickell's state-
ment that the ape is nearly always found in parties of about
a dozen, although several parties may sometimes mingle
temporarily.
From observations on captive specimens of this Gibbon
in Rangoon Sir S. M. Robinson discovered that the period
of gestation is seven months as in ordinary Monkeys, not
nine months as in the Chimpanzee. At birth the young one
had very little hair on the head and a wizened, old-looking
face ; but in a short time the hair grew and the skin of the
face filled out. It clung to the hair of the mother, who held it
in place with her arm, thigh or foot. In another case, recorded
by Mr. Ogilvie, the <$ stayed with the $ after the birth of the
young one, which was carried by the $ until it was over two
years old. During that time it was never seen to take any
food but its mother's milk.
Section GYNOMORPHA.
To this group of Catarhine Primates belong the Monkeys
of Africa and Asia. The tail, in British Indian species, is
generally long or longish, but may be reduced to a stump
1 in. or less in length. The arms and legs are subequal in
length, the hands are much shorter than the feet and have the
thumb weak, tied closely to the palm and emerging from the
middle of its inner edge ; the heel of the foot is narrower
in proportion to its length, and the pads of the palms and soles
are comparatively well defined.
As in the Apes, or Aetthbopomobpha, these structural
features are correlated with the habitual method of progression
both in trees and on the ground. In running or walking the
attitude is essentially quadrupedal. The fore and hind quarters
are nearly on a level, and part or the whole of the palm of the
hand is applied to the ground, but the heel of the foot is raised.
The erect bipedal attitude can be assumed for the purpose
of looking around, but walking on the legs is difficult and only
occasionally resorted to. In climbing, rapid progression
through the trees is effected by a series of leaps, the propelling
power lying in the strong, springy hind quarters and legs.
It is on account of their resemblance to typical quadrupedal
mammals in their attitude and way of moving on the ground
that these monkeys are called Cynomobpha or Dog-like ;
and in their method of climbing they do not differ essentially
from ordinary arboreal mammals. They are clearly less
highly organised than the Apes ; but are much more varied
in their habits, some being essentially forest dwellers, whereas
others live mainly on the ground, often on rocky hill-sides.
32 CEBCOPITHECIDiE.
The Cynomobpha are divisible into the following two
families, both of 'which are represented in the British Indian
fauna : —
a. Check-pouches present, the stomach simple [p. 32.
and undivided Cereopitheeidee,
a'. Cheek-pouches absent, the stomach complex
and sacculated Colobidae, p. 83.
Family CERCOPITHECIDJE.
To this family belong the Baboons (Papio) the Mangabeys
(Cercocebus), the Cercopithequ.es or Guenons (Cercopithecus)
and other genera restricted to Africa, and the Macaques
(Macaco), which are mainly found only in southern and
eastern Asia, with one isolated species in Morocco, whence
it has been introduced to Gibraltar. The British Indian
species in this volume are, for the sake of convenience, all
assigned to Macaco, ; but they fall into several natural groups
for which generic or subgeneric names have been proposed
and are sometimes adopted The characters on which these
are based are mainly the structure of the genital organs in
the male, the length of the tail, the direction of hair-growth
on the head, and a few other features of less importance.
As in most Monkeys, the external characters supply a much
better basis for the determination of the species than the skulls,
which are individually very variable, and. intergrade in some
cases in a remarkable way.
Genus MACACA Lacepede.
Macaca, Lacepede, Tabl. Mamm. p. 4, 1799 (reprinted Mem. Inst.
Hat. Sei. iii, p. 490).
Macacus, of most subsequent authors.
Type, of the genus, Simia inuus Linn.
Distribution. — Apart from the typical species, isolated and
indigenous in Morocco and Algeria, restricted to south-
eastern Asia roughly from Kanristan to Pekin, omitting the
Tibetan Plateau, and thence to Borneo and the Philippines.
In all its external features this genus is very variable.
Its nearest ally appears to be the African genus Cercocebus,
the Mangabey. From this it may be distinguished by the
absence in the skull of the deep pit on the side of the muzzle
just below the orbit.
In many species of Macaques the hair on the crown radiates
from a central whorl and is differentiated by its direction,
sometimes by its length, from the hairs on the sides of the
crown and behind the brows to form a definite " cap." In
British Indian species this arrangement occurs in M. silenus,
MACACA. 33
M . nemestrina, M. speciosa, M. radiata, and M. sinica, the
" cap " reaching its maximum of development in the last
two, which take their trivial names, " Bonnet " and " Toque "
Macaques, from that feature. The arrangement is also found
in, perhaps, about 25 per cent, of skins of M . assamensis.
But in M. mulatto, and M. irus the hair slopes backwards
from the brow without a parting or whorl, and the " cap "
is similarly absent in the S. Chinese M . cyclopis, the Japanese
M. fuscata, and the " Gibraltar Ape," M . sylvana. Probably
the " cap " is a later development in evolution than the
straight-haired arrangement.
Key to the Identification of the Species of Macaea.
a. Colour black, relieved by a ruff of long
greyish hair extending on each side of the
face from the temples to the throat sihnus (Linn.), p. 66.
a'. Colour paler, no such ruff on sides of head.
b. Tail inconspicuous, reduced to a stump
seldom over 1 in. long speciosa Cuv., p. 69.
b'. Tail always conspicuous, shortish or long.
c. Tail shortish, up to about half the
length of the head and body and to
about twice the length of the foot . . .
d. Tail thin and short-haired, carried in
an arch ; hair on crown short, [p. 58.
radiating to form a definite cap . . nemestrina (Linn.),
d'. Tail normally hairy, not carried in
an arch ; no definite cap on crown.
e. Hind quarters orange-red,brighter [p. 44.
than fore quarters nmlaita (Zimm.),
e'. Hind quarters never red, and
typically duller than fore [p. 52.
quarters assametisis MeOl.,
c'. Tail almost as long as the head and
body or longer, about four times the
length of the foot.
/. Hair on crown short, sweeping
backwards from brow irus Cuv., p. 78.
/'. Hair on crown long, with median
whorl or parting, forming a "bon-
net."
q. Haii' on cheek growing upwards
from throat to crown without [p. 38.
whorl; forehead more exposed . . radiata (Geoffr.),
jr'.Hair on cheek forming a definite
whorl ; forehead more covered. . sinica (Linn.), p. 34.
In this key the species are not arranged altogether in
accordance with their affinities. The natural groups into
which they fail and the names that have been given to them
are as follows : —
Silenus, Goldfuss, 1820 ; type, silenus. Here also, apparently,
belongs nemestrinus, which has several characters in
common with silenus, despite the differences set forth
in the key.
VOL. I. D
34 OEECOPITHECID^;.
Lyssodes, Gistel, 1848 ; type, speciosus. This is an isolated
species, distinguished mainly by the abnormal external
genital organs of the <J.
Rhesus, Lesson, 1840 ; type, mulatta. M. assamensis may
also be provisionally referred here.
Cynamdlgus, Reichenbach, 1862 ; type, irus. This species
also stands alone, the length of the tail, in which it resembles
the others entered under e', being merely a primitive
character.
Zati, Reichenbach, 1862 ; type, radiata ; also contains sinica.
These two are nearly alike in the mode of hair-growth
on the head, and differ from the rest in the peculiar
structure of the cj genitalia.
The type-species of Macaca, an earlier name than any of
the others, is apparently most nearly related to Rhesus,
but differs in its tail, which is generally wholly suppressed
and is at most just detectable.
3. Macaca sinica (Linnaeus). The Toque Macaque.
Simia sinica, Linn., Mant. Plant., p. 521, 1771 ; Sehreber, Saug. i,
p. 108, 1775.
Macacus sinieus, Kelaart, Prodr. Faun. Zeylan. p. 8, 1852.
Cynamolgus (Zati) sinieus and audeberti, Reichenbach, Vollst. Nat.
Affen, pp. 130 and. 132, 1862.
Macacus pileatus, Blyth, Cat. Mamm. As. Soc. p. 9, 1863 ; Anderson,
Zool. Res. Yunnan, p. 91, 1878 ; Blanford, Mamm. Brit. Ind. p. 24,
1888 (not pileatus Kerr, 1792, or of Shaw, 1800, or Reichenbach,
1862). ,
Macaca sinica, Hinton & Wroughton, Journ. Bomb. Nat. Hist. Soc.
xxvii, p. 813, 1921 ; Pocock, Journ. Bomb. Nat. Hist. Soe. xxxv,
p. 281, 1931 ; Hill, Ceyl. Journ. Sci. xvi, p. 316, 1932 ; Phillips,
Man. Mamm. Ceyl. p. 5, 1936 *.
Vernacular. — Red Monkey ; Rilawa (Sinhalese) ; Kurangu
(Tamil); Sirra Kurangu (Jaffna Tamil) .
Locality of the type unknown.
Distribution. — Cbttlon.
The smallest of the British Indian species of Macaca,
with the tail subcylindrical, smooth-haired, and long, longer
than the head and body ; the hairs on the crown long, radiating
* The confused synonymy of this species and of its near ally M . radiata
of Southern India was cleared up in 1921 by Hinton and Wroughton,
who showed that the " Bonnet chinois " of BuSon, upon which Simia
sinica Linn, was based, was the reddish Ceylonese Bonnet Macaque,
and not the Indian form as described by Anderson and Blanford.
Also that the description of pileatus given by Kerr (Anira. Kingd. p. 69,
1792), an author ignored by Blanford as not being " binomial," most
emphatically does not apply to any Macaque, nor, indeed, to any
monkey as yet identified. The same is true of pileata Shaw (Gen. Zool. i,
p. 53, 1800), to whom Blanford assigned the name. In this monkey
the hair on the crown forms an upstanding rounded tuft, the limbs-
are black, and the body brown. Hinton and Wroughton's conclusions,
were confirmed by Miller (Anat. of Rhesus, p. 4, 1933).
MACACA.
35
from a central whorl, those in front of the whorl forming a
definite frontal fringe, with their tips nearly reaching the
black eyebrows ; the hairs on the cheek short, not forming
bushy -whiskers, those on the temples and close to the face
directed backwards and meeting the upward and forward
stream in front of the ear to form a distinct whorl on the cheek.
The penis large, with a long, swollen glans which has a well-
developed " corona." No periodic catamenial swelling affect-
ing the tissues round the genital orifice in the $.
General colour above variable, sometimes brownish or
olivaceous, with paler buffy speckling, but usually more or
less ochreous or reddish, sometimes bright and ferruginous,
the limbs externally typically paler than the back, but the
upper thigh brighter, the tail above darker, with a blackish
tinge ; the underside, inner side of the limbs, sides of the
neck, and cheek white or whitish, sometimes with a pale area
Fig. 9. — Upper and side views of head of Toque Macaque (Macaca
sinica) to show typical hair-growth on the crown and cheeks.
(Drawn from skin of the type of inaurea from Cheddikulam.)
passing over the ear from the temple. Hairs on the crown
very variable, sometimes directed forwards as far as the
brows, but sometimes there is a shortish area behind the brows,
with the short hairs directed sideways or backwards.
The skull of M . sinica is small for the genus, and in the adult
cj rather low for its length, due to the produced jaws ; the
brow is prominent owing to a pronounced depression behind it ;
the temporal ridges are moderately well developed, sometimes
meeting in the middle line ; the occipital crest is moderately
salient, and there is a distinct constriction of the muzzle
caused by a depression behind the canines and below the orbit*.
* Phillips published an excellent photogravure of the skull of an
adult cj of this species, said to be §■ nat. size. It is, however, reduced
by considerably more than \. I have no material sufficing to show
if there is any cranial difference between the two races of M. sinica
Probably there is not.
d2
36 CEECOVITHECIDJE.
3 a. Macaea sinica siniea (Linnaaus).
Macaco, sinica sinica, Pocock, Journ. Bomb. Nat. Hist. Soc. xxxv,
p. 285, 1931 (restricted) ; Phillips, Man. Mamm. Ceylon, p. 5,
1936.
Macaea sinica inaurea, Pocock, Journ. Bomb. Nat. Soc. Hist,
xxxv, p. 286, 1931 *.
Locality of the type -unknown ; of inaurea, Cheddikulam,
N.P.
Distribution. — " The low-country dry zone " from the extreme
north to the extreme south of Ceylon (Phillips).
The " bonnet " uniformly coloured throughout, the hairs
drabby-greyish, with paler buffy tips. The general colour is
individually and seasonally very variable; usually in the
fresh coat the hairs of the upper side are dark greyish or blackish
for about two-thirds of their length, with the tips ochreous
or reddish, the shoulders and outside of the thighs being
brighter than the back, but the tips may be dull buffish, so that
there is no bright tint in the pelage except a little on the
outer side of the thigh. Sometimes the dorsal tint is more
uniformly brown without bright speckling. This may be
due to the breaking off of the pale tips of the hairs when dead
before the moult. Skins, for instance, from Maha Oya, E.P.
(Aug. 12 and 17), have the coat on the back short, coarse,
and dark olive-brown in hue, quite unlike normal fresh-
coated skins from the Eastern Province ; but beneath the
old brown hairs the newly erupting coat shows the characteristic
rich ochreous speckling. The hairs of the frontal fringe vary
from about 45 to nearly 70 mm., and of the shoulders from
about 45 to 55 mm.
The following are some measurements (in English inches)
and weights, several being Phillips's records : —
Head and
Locality and sex. body. Tail. Foot.
Phillips's largest, loc. ? ; ad <J 21 22£ 5
Mankeni, E.P. ; ad. 5 19i 23f 5J-
Oheddikulam, N.P. ; ad. g 18f 22£ s|
Mankeni, E.P. ; ad. g 17§ 21f 5
Phillips's average of 6 ad. gg 19 21 J 5
Phillips's largest ad. $ 17 22£ —
Wellawaya, Uva ; ad. $ 16f 22 4f
Phillips's average of 2 ad. £$ 16|- 21 4f
* I here accept Phillip's decision to the effect that dull and bright
tinted specimens in fresh coat occur mdiscrirnioately throughout the
dry zone, although the material at my disposal suggested that bright
reddish specimens are dominant at Mankeni and Maha Oya in the E.P.,
and at Wellawaya and Tellulah in Uva, and duller, olivaceous specimens
in the N.P., the two meeting at Kala Oya in N-W.F. This opinion was
expressed by the racial name inaurea given to an adult g in perfect
coat (Nov.) collected at Cheddikulam, north of Adam's Bridge, N.P.
On the label of this skin, which closely resembles some skins of the
MACACA. 37
The weight of the larger $ from Mankeni was 12 lb., of the
smaller 10 lb., the one from Cheddikulam being lOf lb., the
same as Phillips's average of 4 $g ; Phillips's largest $ was
9| lb. and his average of 2 $$ 7| lb.
This Toque Macaque is found in the jungles of the low-
country dry zone about as far south, according to Phillips,
as lat. 8° 1ST. in the west and central portion of the island
and on the eastern side to the extreme south. Major E. W.
Mayor, who collected many specimens both in the northern
and eastern parts of its range, found them shy and very difficult
to shoot. Phillips reports that troops composed solely of
young individuals have been observed on several occasions.
The rest of its habits are no doubt the same as in the next,
better-known race.
3 b. Macaea siniea aurifrons Pocock.
Macaca siniea aurifrons, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxv, p. 286, 1931 ; Phillips, Man. Mamm. Ceylon, p. 8, 1936.
Locality of the type, Rayigam ELorale, Western Province.
Distribution. — " The low-country wet zone and the central
hill -zone " of Ceylon (Phillips).
Distinguished from typical siniea by the colour of the
" bonnet," of wlieh the hairs of the anterior part or fringe
are uniformly reddish or buffy throughout and those of the
posterior half dusky greyish at the base, reddish at the tips.
The general colour of the upper side is also rather brighter
red on the average, and a darker appearance is imparted to
the pelage by the partial exposure of the deeper black base
of the hairs owing to the looseness of the rather longer coat.
The type of this race is an unusually richly red specimen
with the hairs of the frontal fringe 75 mm. and of the shoulder
58 mm. In another the same measurements are 72 and 60 mm.
The following measurements (in English inches) and weights
(lb.) are taken from Phillips's volume : —
Head and
body. Tail. Foot.
Largest $ 19$ 24i 5£
Average of 3 <?<J 18£ 23" 5+
Largest 9 17£ 18J 4£
Average of 4 $? 16£ 20 4J
The average weight of 2 gg was 9| lb., of 2 ?? 7£ lb.
These particulars suggest that this race may be a trifle smaller
on the average than typical siniea ; but the data are insufficient
Indian species, M. radiata, except that there is a little red on the outside
of the thigh, the collector, E. W. Mayor, who two months earlier had
secured a fair number of skins in the E.P., wrote : "Fur seems darker
here," i. e., at Cheddikulam, where he saw living specimens.
38 CEKCOPITHECID-SE.
and there are not enough adult skulls available to throw-
light on the point. (For skull-measurements of M. sinica
see p. 43.)
According to Phillips this race of the Toque Macaque is
found from the highest mountain peaks to the seashore, and
on the lower hill-slopes of the northern and eastern portions
of its area it everywhere blends with the typical dry-zone
lowland race. It occurs both in the forests and in the ' ' sholas "
or isolated patches of jungle in the grass-covered hills. Like
other Macaques it is gregarious, its troops consisting of about
twenty or more individuals of both sexes and of all ages.
It is mostly active by day, moving about and feeding both in
trees and on the ground, its diet consisting of fruits, berries,
flowers, seeds, and shoots as well as of insects, grubs, and
spiders. Often also it raids plantations of coco-nut and
cardamom, committing considerable damage. In districts
where it is persecuted on this account it is wary of man,
but otherwise pays little heed to him. It has, however
a great dread of leopards, pythons, and crocodiles, the latter
especially when it approaches water to drink. Sometimes,
however, it takes to the water to hide and is a good swimmer,
both at the surface and below it. Like the Wanderoos (Leaf-
Monkeys), with which it occasionally associates, it has the
habit of hiding in the foliage when frightened.
4. Maeaca radiata (Geoffroy). The Bonnet Macaque.
Gercocebus radiaius, Geofiroy, Ann. Mus. Nat. Hist. Paris, xix, p. 98,
X912 ; F. Cuvier, Hist. Nat. Maram. folio 1, no. 33,
Maeaous radiatus of Blyth, Jerdon, and other earlier writers on
Indian Mammals.
Maeaca radiata, Hinton& Wroughton, Journ. Bomb. Nat. Hist. Soe.
xxvii, p. 814, 1921 ; Pocock, Journ. Bomb. Nat. Hist. Soo. xnv,
p. 276, 1931 ; Hill, Ceyl. Journ. Sci. xvi, p. 314, 1932.
Macacus sinicus, Anderson, Zool. Res. Yunnan, p. 90, 1878 ; Blan-
ford, Mamm. Brit. Ind. p. 23, 1888 (not of Linn.) *.
Vernacular. — Bandar, Bandra (Hindi and Dekhani) ; Mdkad
Lai manga (Marathi) ; Makadu, Wdnar, Kerda (Mahr.) ; Mang,
Kodaga, Koti, Edpi, Maungya, Kemp Manga (Kanarese) ;
Koranga, Vella Manthi (Mai.) ; Kurangu or Corongv, (Tamil) ;
Mucha (Coorg) ; Kodan (Toda).
Locality of the type unknown ; apparently a menagerie
specimen, according to Miller.
Distribution. — Soxttheen India, at least as far north on the
west as Satara, and on the east not north of the G-odaveri
River.
Distinguished from M . sinica by its larger size, as shown
by the flesh and cranial measurements, by the hair-growth of
the top and sides of the head, and by the generally duller colour.
* For the synonymy of this species see above under M . sinica.
MACACA.
39
The arrangement of the hair on the crown is typically
fundamentally the same as in M . sinica. It always radiates
from a central whorl and is long posteriorly and laterally,
but anteriorly it is always shorter, so that the fringe overlaps
a smaller area of the forehead, and the bonnet, instead of being
circular, is subelliptical. Usually the tips of the hairs of the
fringe extend about half-way between the whorl and the
eyebrows, and the short hairs behind the brows extend
sideways from a median parting*, but the arrangement
is very variable. Sometimes there is a whorl far back on the
forehead, whence the short hairs radiate at the hinder end
of the medium parting ; sometimes the hairs of the centre
of the fringe in front are only a few mm. long, and the lateral
hairs extend like a tuft on each side ; sometimes the hairs
behind the brows are directed backwards, imparted and long,
those in the centre overlapping the whorl on the crown and
ITig. 10. — Upper and side views of head of Bonnet Macaque (Macaca
radiata) to show typical hair-growth on the crown and cheeks.
(Drawn from skin from South Coorg.)
the mesially abbreviated fringe in front of it. The short
hairs on the cheek grow upwards from the throat and lower
jaw to the level of the top of the ear or thereabouts without
forming the whorl described in M . sinica.
The coat and colour are variable seasonally and individually
independently of the season. Typically the winter coat
is deep brown or olive-brown above, with little, if any, pale
speckling, and the hairs of the bonnet are dark distally,
paler at the base, the arms are greyer than the back, the legs
greyer than the arms, the tail is blackish above in its basal half,
brown distally, and the underside is whitish-grey ; but in
spring and early summer the long hairs of the upper side,
up to about 70 mm. on the shoulders, fade to a buffy-grey
tint, and become dull, lustreless, shaggy, and harsh to the touch.
Occasionally, however, there is a distinct yellowish speckling
* Described by Hinton and Wroughton and illustrated by Hill.
The hairs of the middle of the fringe vary, however, from about 25 to
«0mm.
40 OEKCOPITHECIDiE.
in the dorsal pelage recalling that of M . sinica, and in one
skin there is a reddish tinge in the hairs of the flanks, shoulders,
and sides of the neck. Some skins are very like the skin of
M. sinica from Cheddikulam, except that they never have
the brighter hue on the outside of the thigh*.
The skull of N. radiata is larger than that of M . sinica,
and has the brows less prominent owing to a shallower
depression behind them ; and this is associated with relatively
narrower width across the orbits, with on the average at
least weaker temporal ridges and a shallower depression on
the muzzle behind the canines and below the orbits. These
features indicate less muscular development.
4 a. Macaca radiata radiata (GeoSroy).
Macaco, radiata radiata, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxv, pp. 276-8, 1931 {sensu atricto) ; Osman Hill, Froc.
Zool. Soc, Syst. 1937, p. 212.
Locality of type unknown.
Distribution. — As above, but replaced in Travaneore by
the next race.
General colour as described above under the species,
typically olive or greyish-brown on the upper side, sometimes
speckled; bonnet rather darker, greyish- white on the under-
side, with areas of dark pigment in the skin emphasizing the
pale hue of the hairs.
The following are the flesh-measurements (in English inches),
indicating the range in size : —
Head and
Locality and sex. body. Tail. Foot.
Bellary : ad. <J 23f 27 6£
Shevaroy Hills ; ad. <? 21f 24| 5|
S.W. Dharwar ; ad. £ 2l| 22J 5
N. Coorg ; ad. $ 2\\ 20 5
Satara ; ad. (J 20| 25| 6
Dharwar ; ad. $ 20| 22§ —
Dharwar ; ad. $ 19| 22| 5
Dharwar ; ad. ? 18J 2o| 4|
* Osman. Hill, who compared living and injected examples of M . sinica
and M . radiata, says that M. radiata differs in having the face muddy-
flesh tinted, with soft hairs, the edges of the lips and the entire ears
unpigmented, and the ears more prominent, whereas in M . sinica the
face is pale flesh, with bristly hairs, the edges of the lips and the ears-
pigmented, and the ears less prominent. His figures, however, con-
tradict the statement regarding the prominence of the ears, and in
dried skins of radiata the ears may be pallid or dark greyish-black..
As a further difference he says that M. sinica has a well-marked tract
of white hairs passing over the ears from the upper cheek to the side
of the neck, the same tract in M. radiata being dark. This tract is.
very variable in distinctness in made-up skins of M. sinica, and in
a skin of M . radiata from Mysore there is a distinct pale tract of light
hairs on the area in question. In the texture of the hairs of the face-
I can find no constant difference.
MACACA.
41
The tail is very variable. It may be as much as about
5 in. longer than the head and body or 1 in. shorter. The
weights of the adult <J range from 13 lb. (N. Coorg) to 19| lb.
(Bellary) ; of adult $ from 7 to 8 lb.
A few isolated specimens, e. g., one from sea-level below
the Gersoppa Falls in Kanara and. another from Cumbum in
Madura, which are aberrant in colour, suggest the possibility
of the inclusion of more than one race in typical radiafa as
above defined. Setting these aside, this Macaque occurs in
Western India in Satara, Dharwar, 2,300 ft. ; Kanara, 2000, ft. ;
Mysore, Coorg, 2,000 to 3,555 ft. ; the Nilgiri Hills, 5,120 ft. ;
Cochin, 1,500 ft. ; the Palni Hills, 3,000 to 5,500 ft., and in the
Eastern Ghats at Salem, the Shevaroy Hills, 4,500 ft., and
Kurnool. It is even found "in populous towns, where it
pillages the shops of fruit and grain." In Cochin, at least, it
is commoner, according to R. O'Brien, in the plains than in the
Fig. 11. — Skull of adult S Bonnet Macaque (Macaca radiata) from
Dharwar. X £.
hills. Shortridge reports that in Dharwar it is very plentiful,
going about in large parties of from 20 to 30 individuals.
On the banks of a stream he saw troops of them mixing
apparently on perfectly friendly terms with Langurs, but when
alarmed the two species separated. Near the Hindu temples
at Gaday, where it is protected, large numbers are said to have
died of bubonic plague. At Vijayanagar in Bellary it is plentiful
in the vicinity of Hampi Temple, and it is sufficiently plentiful
throughout Coorg to be a great nuisance to the owners of
coffee estates by destroying large quantities of ripe berries.
Its habits no doubt are the same everywhere, and similar
to those of M . sinica. In the Nelliampathy Hills, according
to Kinloch, it occurs in moderate troops on the northern
cliffs and in the bamboo jungles on the south slopes, but not
in the evergreen forest of the plateau.
42 CEECOPITHEOID-a!.
4 b. Macaca radiata diluta Pocock.
Macaca radiata diluta, Pocock, Jaunt. Bomb. Nat. Hist. Soc,
xxxv, p. 278, 1931 ; Hill, Proc. Zool. Soc, Syst. 1937, p. 212.
Locality of the type, Boothapaundy, on the Ghats north
of Aramboly in Travancore.
Distribution. — Tbavancore.
Distinguished from typical radiata by its colour, which,
although very variable, is paler, the fore part of the bonnet
tending to ochraceous, and the same colour often present on
the shoulders, lower back, buttocks, and thighs, the underside
varying from dirty white to buff.
This description is taken from the account given by Hill,
who saw many living specimens. I described the race on the
evidence of two skins, and distinguished it by its much paler
winter coat, buffy throughout above, the underside being
cream, with the skin pallid, not blackish. The flesh-measure-
ments of the larger of the two specimens collected by
E. S. Pillay are : —
Head and
Locality and sex. body. Tail. Foot.
Boothapaundy ; ad. ? 16f 20$ —
Boothapaundy ; ad. £ 13§ 21 —
These dimensions suggest a smaller race than typical
radiata ; but the skull of the smaller specimen, the type, is
as large as $ skulls of radiata.
Blanford referred to a specimen from Travancore with
" unusually long top-knot." In the type of diluta the hairs
of the forehead are long, and sweep backwards from the brow
to form with the front hairs of the bonnet a kind of upstanding
crest just in front of the whorl on the crown. Presumably
this feature is inconstant, since Hill does not refer to it.
According to Hill this Macaque is quite common in Travan-
core and is distributed throughout the district. He came
across two troops associated with Hindu temples, one a few
miles south of Alleppey, on the road to Quilon, the other
five miles south of Aramboly.
Although the $ skulls of radiata intergrade in size with those
of sinica, the general superiority of the skulls of both sexes
of radiata is shown by the following averages : —
Average total length of 8 ad. <^ skulls of radiata = 121 mm.
6 ad. ? „ „ = 104 „
,, „ 4 ad. <j „ sinica = 115 „
„ » 4 ad. $ „ = 98 „
MACACA.
43
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44 CERC0PITHEC1D.3B.
5. Macaca mulatta (Zimmermann). The Rhesus Macaque.
(For synonymy and bibliographical references see under the
subspeoiflo headings.)
A medium-sized or tolerably large Macaque, with the tail
uniformly hairy throughout, typically pendulous, not carried
in an arch above the level of the buttocks, tolerably short,
usually about half the length of the head and body, typically
considerably longer than the hind foot, but in some Chinese
races hardly exceeding it. Hair of the crown directed back-
wards from the brows, without a whorl or parting, and not
forming a definite crest or " cap " ; no definite crest on
the cheeks or temples, but usually a whorl low down on
the cheek. Colour very characteristic, the head, shoulders,
arms, and fore back generally olivaceous in hue owing
to the hairs being blackish or grey, but speckled with
duller or brighter buff annulations ; but the loins, rump,
and the base of the tail are orange-red in varying intensities ;
the base of the hairs is grey or whitish, not annulated. The
skin of the face, hands, and feet is pallid, although the face is
sometimes suffused with red. The penis is tolerably large,
with the glans elongated, about twice as long as its posterior
width, expanded distally, with the left lip of the orifice much
larger than the right and the orifice terminal. There is no
noticeable catamenial swelling in the female, but the skin
of the rump is often bright red.
The skull of M. mulatta * is much more variable in size and
shape than the skull of M . radiata, and has the sagittal crest
and the temporal ridges typically less well developed, and there
is scarcely a trace of constriction behind the canines on the
muzzle. But in skulls of about the same length in the two
species the skull of M. mulatta has the zygomatic and trans-
orbital widths a little greater, giving a more robust aspect
to the skull. The facial profile may be about the same as
in M. radiata, the interorbital septum and the nasals forming
a relatively shallow, evenly curved concavity ; but in some
skulls the septum is more steeply and the nasals less steeply
sloped, so that the concavity is deeper, giving a " snub-nosed "
aspect to the face. I have not observed this condition in any
skull of M . radiata.
Distribution. — Fxova. Kafiristan and Chitral through the
Him ala yas and Noethben India eastwards to Bubma,
Siam, Indo-China, and Hainan, and northwards through
China to the latitude of Pekin.
Three races are found in British Indian territory. In
* As pointed out by Hinton and Wroughton, the drawings published
by Blanford, (p. 12) to illustrate the skull of M. rhesus {=imdatta) were
taken from a skull of M . assamensis. Anderson put Blanford wrong.
PLATE
MAMMALIA.
Upper fig.— McMahon's Rhesus Macaque (Macaca mulatto mcmahoni).
Lower fig.— Common Rhesus Macaque (Macaca mulatto mulatto).
MACACA. 45
addition to these, M. m. lasiotus Gray inhabits Szechwan,
M. m. sancti-johannis Swinhoe (—littoralis Elliot) occurs in
the neighbourhood of Hong-kong, and M. m. tchelimsis
M.-Edwards near Pekin.
Since the three British Indian races merely differ slightly
in size, thickness of coat, and some cranial features from each
other, it is needless to epitomize their distinctive characters
in an analytical key.
5 a. Macaea mulatto mulatta (Zimmermann).
Cercopithecus mulatta, Zimmerman*!, Geogr. Gesch. Mensch. ii,
p. 195, 17S0 (based on. the " Tawny Monkey," Pennant, Syn.
Quadr. p. 120, 1771).
Simia (Cercopithecus) fulvus, Kerr, Anim. Kingd. p. 73, 1792.
Sirnia rhesus, Audebert, Hist. Nat. Singes, p. 5, 1798.
Simia erythrsea, Shaw, Gen. Zool. i, p. 33, 1800.
Macacus (Pithex) oinops and nipalensis, Hodgson, Journ. As. Soc.
Beng. ix, p. 1212, 1840.
Macaea mulatta, Hinton & Wroughton, Journ. Bomb. Nat. Hist.
Soc. xxvii, p. 668, 1921 (where the involved synonymy of this
Macaque is unravelled).
Macaea mulatta mulatta, Poeoek, Journ. Bomb. Nat. Hist. Soc.
xxxv, pp. 533-9, 1932.
Vernacular. — Bandar (Hind.) ; Markat (Beng.) ; Myouh Sat
(Burmese) ; Leng Leng (N. Shan States).
Locality of types of mulatta and fulvus, both based on the
" Tawny Monkey " of Pennant, " India " ; of rhesus and
erythrsea, based on " Macaque a queue courte " of Buffon,
Hist. Nat. Suppl. vii, p. 56, 1789, unknown ; of oinops and
nipalensis the Nepal Tarai. This locality, the first definite
district of India assigned to the species, I selected in 1932
as the type-locality of mulatta.
Distribution. — The whole of Nobthern India from Nepal,
Bhutan, N. Kamrtxp, and Assam as far south at least as the
Tapti Kiver in the west and not south of the Godavari in the
east ; also the whole of Burma from H'Kamti and Homalin
in Upper Burma to the south of Prome. Evidence for the
occurrence of this race in Siam is supplied by a specimen
described as M . siamica by Kloss ; in Indo-China by specimens
collected by Delacour and Lowe in Annam and Tong-king
and in Hainan by specimens which Elliot described as
Pithecus brevicaudus.
The smallest of the three British Indian races of M. mulatta,
both in size of the body and the skull, which has on the average
a weaker muzzle and less pronounced brow-ridge and the
temporal ridges farther apart. The winter coat is also less
woolly and shorter, from 2 to 2| in. on the shoulder.
The colour in the fresh, unfaded coat is very variable
individually, irrespective of age, sex, locality, and season,
46
CEECOHTHECID-a] .
in accordance with the brightness and extent of the paler and
darker annulations and the depth of hue of the bases of the
hairs of the head and fore quarters, and the duller or more
brilliant orange of the hind quarters. The variation is well
illustrated by two adult <J examples collected on 21 November
and 25 November at Rajapara, S. Kamrup, 600 ft. In one
the head, nape, and shoulders are yellowish-olive, the annula-
tions of the hairs being clear yellowish-buff, with the bases
ashy-grey ; the back is brighter and yellower, turning to orange
on the loins, and nearly fiery red on the outside of the thighs,
the basal portion of the hairs on the red areas being white,
the distal portion red and not annulated. The tail is like the
Pig. 12. — Skull of subadu.lt <J Bhesus Macaque (Macaca mulatto, mulatto,)
from. Bhutan Duars.
rump at the base, greyish-olive terminally. The arms are
grey, with yellowish-buff speckling ; legs washed with bright
yellow, paling from the thigh to the foot, which is greyish-
yellow. Underside mostly white, slightly red on the belly.
The other is much darker, the head, nape, and shoulders being
deep greyish-olive, the annulations of the hairs being pallid,
greyish-buff, with the bases deeper smoky grey ; the back is
a more yellowish-olive, with the lumbar region like the head
and nape of the first specimen ; the buttocks and outside
of the thighs are not nearly so fiery or so extensively red in
hue. The tail is much darker ; the arms also are darker,
with grey speckling and blacker hands.
Occasionally the colour of the fresh coat is rich rusty red
all over the dorsal surface, although brighter on the rump.
One red specimen of this type was secured at Dangs in Surat
MA.CACA. 47
together with, examples more normally tinted. Another,
even redder, came from Baska Nadi, N. Kamrup, 2,000 ft.,
collected on 5 January. This specimen differs strikingly
from the two from S. Kamrup recorded above.
The colour, as well as the length and texture of the coat,
also varies greatly in accordance with the season. In the
winter, from about November to January, the coat is perfect
in colour and soft in texture, the tips of the long hairs tolerably
smoothly overlapping, so as to conceal the grey basal portions.
Through March, April, May, and June the coat gradually
loses its lustre, softness, and colour, the hairs, owing to bleaching,
tend to become more uniformly tinted, buflSsh- or brownish-
grey, and to adhere loosely, in long tufts or patches, displaying
the grey hue of their basal portions. The coat is then shaggy
and streaky, and decidedly harsh to the touch, the terminal
portion of the hairs subsequently breaking off, seemingly
from brittleness. Before this dead coat is shed the new coat
begins to erupt beneath it ; and in May or June, as a rule,
the old coat begins to be shed, clearing first on the crown,
the hands and feet, and the tail, the tail, when its long
hairs have dropped, closely resembling that of the Pig-tailed
Macaque (M. nemestrina Linn.). Even as late as July, or
even August, remnants of the old, dead, uniformly greyish
coat may be found on parts of the body. Similar variations
in the colour, independent of the season and in accordance
with the seasonal condition of the coat, occur also in the other
two British Indian local races of this species.
The moult is not exactly coincident in time, even in the
same locality, and no doubt varies still more with altitude
and latitude in accordance with the onset of warm weather.
The following are the flesh-measurements (in English
inches) of some specimens of M . m. mulatto, : —
Head and
Locality and sex. body. Tail. , Foot.
Chindwin ; ad. <J 23 8£ SJ
Mingun, Upper Burma ; ad. $ 22 8 6|
S. Kamrup ; ad. (J 21£ 9 5£
Nepal Tarai ; ad. 3 2l| 104, 64
Gorkna, Nepal ; ad. $ 20| 10 64
Bhutan Duars ; ad. 3 19 8| 6~
Darjeeling ; ad. $ 20+ 8| 54.
Mishmi Hills, ad. $ 19J 11| 5|
Sohagpur, C. India ; ad. $ 18| 8f 5f
The weights of the $ specimens from Mingun and S. Kamrup
were 15 and 16J lb. respectively, of the $ from Darjeeling
11J lb. ; but a $ of about the same size from the N. Shan
States was 13 lb.
About the special habits of this race of mulatto there is
little to report. Blanford described it as the common monkey
48
CEECOPITHBCID^!.
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MACACA. 49
of Northern India as far south, as the Godavari. Although
not regarded as sacred by the Hindus, it is left unmolested by
them. This no doubt explains its general fearlessness of man,
which he reports, and its occasional occurrence in native
villages. Otherwise it is most usually seen near cultivated
grounds, especially by tanks. In the wilds, he adds, it is
commoner amongst trees by streams than in forest jungle.
Crump, on the other hand, stated that at Singbhum in Bengal
it was very shy and difficult to approach owing to the thick
undergrowth it frequented. Apparently its habits vary
locally. In Burma Shortridge found it fairly plentiful at
Hsipaw in the North Shan States, often near paddy-fields and
other cultivated tracts, where small parties of it were observed.
But here, too, it was shy of man, possibly because it is some-
times eaten by the Burmese, as recorded by Mackenzie, who-
stated that 20 miles south-west of Kindat it was common,
especially near cultivation, although big herds of it lived in
the jungle.
5 b. Macaca mulatta villosa True.
Macacus rhesus villosus, True, Proe. U.S. Nat. Mus. xvii, p. 2, 1894.
Piihecus villosus, Elliot, Rev. Prim, ii, p. 200, 1913.
Macaca mulatta villosa, Pocoek, Journ. Bomb. Nat. Hist. Soo.
xxxv, p. 539, 1932.
Vernacular. — Wandar, Puriz, Punj or Ponj (Kashmir)-
Gye (Hokul).
Locality of the type, Lolab, at the northern end of Wular
Lake, about 40 miles north-west of Srinagar in S. Kashmir.
Distribution. — Southern Kashmir, Upper Pottjab (Murree,
Dharmsala, and Kangra) ; also Ktjmatjn, where it blends
with typical mulatta.
Distinguished from typical mulatta by being heavier,
slightly larger on the average, with the winter coat a little
longer, from about 65 to 90 mm. ( =2 J to 3 J in. ) on the shoulders,
and fuller, and the skull also a little longer, with on the average
a deeper muzzle, more pronounced brow-ridges, and the
temporal ridges closer together, indicating greater muscular
development.
The following are the flesh-measurements (in English
inches) of some specimens of M . m. villosa : —
Locality and Bex.
Murree ; ad. 3
Kangra ; ad. $
Knmaun ; ad. $
Kumaun ; ad. 3
Nairn Tal ; ad. 6"
Naini Tal ; ad ?
Kumaun ; ad. $
VOL. I.
Head and
body.
Tail.
Foot.
25
12
7
23
10
6J+
22
Hi
6*
m
9*
6|
20J
9
6*
19
10
5*
18£
8*
H
50 CEBCOPITHECID^:.
The weight of the three $ specimens from Kumaun and
Naini Tal, in order, were 22, 23, and 17| lb. respectively, and
of the two $ specimens 14J and 13 lb.
Although the (J from Murree is a little larger than the
cj examples of typical mulatto,, the $ and $ specimens from
Kangra and Kumaun are about the same size as in the typical
race ; but the weights are greater, suggesting more sturdily
built monkeys. The dimensions given by True of two <J
specimens from the type-locality, Lolab in Kashmir, as head
and body and tail 23J+9J and 22+11 respectively, agree
closely with those of the <J specimens from Kangra and
Kumaun.
True records that the original examples of this Rhesus
frequented the pine-forests at Lolab at an altitude of 7,500 ft.
The example from Patriata, Murree, in the Upper Punjab,
was shot at 7,150 ft. At Dharmsala, 4,500 ft., and in
Kangra from 2,450 to 5,000 ft., it is, according to Wells, fairly
common, frequenting rocky hill-sides and cliffs bordering
streams, but is very difficult to catch owing to its shyness of
man from being driven away by the owners of the cultivated
fields and gardens it habitually raids. In Kumaun, where it
occurs at altitudes ranging from 3,700 to 1,100 ft., it is more
abundant, Crump tells us, than the Langurs (S. entellus
schistaceus or hector) in the outer ranges of the hills, where,
during cold weather, it is found in large numbers up to 4,000ft.
It sometimes associates with the Langurs by day ; but the
two species separate at nightfall when retiring to rest.
5 c. Macaca mulatta mcmahoni Pocock.
Macaca mulatto, mcmahoni, Pocoek, Journ. Bomb. Nat. Hist. Soo.
xxxv, p. 544, text-fig. of skull and pi., 1932.
Locality of the type, Kootai in Lower Chitral, between the
Bashgal Valley in Kafiristan and the Chitral Valley, 3,600 ft.
Distribution. — Kafiristan and Chiteal.
The largest of the British Indian races of M. mulatta and,
on the available evidence, differing also from vilhsa by its
longer coat, rather duller colour, and bigger skull, which has
the brow-ridges higher and more prominent.
Only two specimens of this Macaque have been described
in detail. The type, killed at the beginning of February,
"before the long, loose coat had reached its full length, has the
hairs on the shoulder about 100 mm. (4 in.) long. In the
general darkness of the hue of the fresh coat the skin differs
strikingly from skins of typical mulatta at the same time of
the year, but closely resembles that of the skin of villosa
from Murree, shot in the middle of June, with the winter
coat deteriorated and in process of moulting. The skull,
JIACACA.
51
although larger than the largest skull of villosa, is not quite
fully developed. The other specimen, a $ from Chitral,
presented by Sir H. McMahon to the Zoological Gardens,
where it died in January, has the coat shorter, only about
60 mm. on the shoulder, but full and longish everywhere.
The colour is a little brighter than in the g, but is, on the
average, more dusky olive on the fore quarters than in villosa.
Although Capt. Raverty may possibly have been the first
to record the occurrence of this Macaque in Kanristan when
he wrote (Journ. As. Soc. Beng. xxviii, p. 332, 1859), " In
some of the warmer parts of Kanristan, in the densely wooded
Pig. 13. — Skull of subadult <J of McMahon's Rhesus Macaque (Macaca
mtdatta mcmahoni) from Chitral.
districts, monkeys of the largest kind are found, but are
not very numerous," I am now inclined to think the
species he referred to may have been a Langur. The credit,
however, of discovering this race of M . mulatto, in the Chitral
district belongs to Sir Henry McMahon (Journ. As. Soc.
Bengal, lxx. pp. 4-5, 1901), who reported, "Monkeys are
to be found in the lower end of the Chitral Valley. Capt.
Gurdon saw a troop of them at Mirkandi, on the bank of the
Chitral Biver, only 4,000 ft. above sea-level. The general
characteristics seem to be those of Macacus rhesus, but all
the specimens I have seen have, instead of a tapering tail,
a tail of about 8 in., coming to an abrupt end as if cut off like
e2
52 CEBCOPITHEOIDiE.
a fox-terrier's." Capt. H. Fulton (Journ. Bomb. Nat. Hist.
Soc. xiv, p. 758, 1909) subsequently observed this Macaque
in Chitral and, identifying it as Macacus rhesus, recorded
it as " Very plentiful at the lower end of the Chitral Valley
in summer on the right bank of the river. They come up as
far as the Utzun Valley in troops at 5,000 ft., and probably
come over the pass into Utzun from Kafiristan Valley."
6. Maeaea assamensis McClelland. The Assamese Macaque.
Macacus assamensis, McClelland in Horsfield, Proc. Zool. Soc. 1839,
p. 148.
Macaca assamensis, Hinton & Wroughton, Journ. Bomb. Nat. Hist.
Soc. xxvii, p 669, where the involved synonymy of this species,
due to Anderson's confusion of it with M . mulatta, is unravelled.
(For synonymy and bibliographical references see under the
subspecifie headings below.)
Distribution. — The Himalayas, at tolerably high altitudes,
from Mussooree through Nepal, Slkkim, Bhutan to Assam,
thence southwards to the Sttndaebans ; Uppeb Bxtema,
N. Siam (?), and Indo-China, Annam, and Tong-king.
Distinguished from M . mulatta, which it closely resembles in
size and in the length and hairiness of the tail, by the entire
absence of orange-red hue on the bind quarters, by the growth-
direction of the hair on the head, and by some cranial and dental
characters.
The coat is soft and full, wavy or smooth, but without
much underwool, and on the shoulders, before the spring
moult, may reach a length of about 85 mm. (over 3 in.).
The general colour above varies individually from dark brown
to lighter yellowish-brown, but the shoulders, nape, crown, and
arms are almost invariably brighter and yellower than the
hind quarters, legs, and tail, which is exactly the opposite
of the condition seen in M . mulatta ; the hairs, however,
are scarcely visibly speckled with annulations anywhere ;
the arms, legs, and tail are approximately the same tint as
the adjoining parts of the body; round the face the hairs are
black, but to a very variable extent ; the cheeks are typically
grey, with the hairs pigmented at the tip ; the underside
and the inner sides of the limbs are also pale, mostly greyish-
white, with a brownish tinge sometimes on the abdomen ;
but the underside of the tail is hardly paler than the upper.
The growth-direction of the hair on the head, not recorded
in the living animal, is very variable in made-up skins, but
typically, at all events, there is a short median parting just
behind the brow-ridges, whence the hair diverges to right and
left and sweeps round the sides of the crown above the ears,
the rest of the hair on the crown being directed backwards.
But in some skins the crown exhibits a distinct whorl, whence
the hair radiates in all directions, forming a kind of " cap "
MMALIA.
PLATE III.
Assamese Macaque (Macaca assamensis)
^^^^^J^^lJ^^m^^:,
«.<
tP: *",
T,
\;}' 3V'*" ''.VA
Stump-tailed Macaque (Macaca speciosa).
MACACA.
53
-with, well-defined anterior edge much resembling that of
M . nemestrina and M . silenus. On the fore part of the cheek
the hairs typically sweep backwards from the face and, meeting
the forwardly-directed hairs from the sides of the neck and
hind cheek, form with them, a low crest below the level of
the ear and in front of it. The condition, however, varies in
made-up skins.
The skulls, at least those of adult <J specimens, may nearly
^always be distinguished from those of M. mulatto by the
following features : — There is a strongly developed, shelf-like
occipital crest, and associated with it almost always a sagittal
crest which starts from the middle of the frontal bones ;
the margins of the orbits are thicker, the muzzle more ' ' pinched : *
behind the canines, and the posterior nares and mesopterygoid
Fig. 14. — Skull of adult cJ Assamese Macaque (Macaca assamensis pelops)
from Tonglu, Darjeeling. x J. (In its facial profile this skull
is intermediate between the two adult cJ skulls of the typical
race from the Mishmi Hills described on p. 56.)
fossa narrower. Also the canines are considerably larger,
but the molars are relatively, if not actually, smaller. Some
skulls approach those of M. mulatto, in poorer muscular develop-
ment, and the race of M . mulatto, which comes nearest to
M . assamensis, is the large Chitral form, mcmakoni.
6 a. Macaca assamensis assamensis McClelland.
Macaca assamensis, McClelland in Horsfield, Proc. Zool. Soc. 1839,
p. 148 ; Anderson, Zool. Res. Yunnan, p. 64, 1879 (description
of the type).
Macacus assamensis, Blanford, Mamm. Brit. Ind. p. 15, 1888, and
of other authors (in part.).
Macaca assamensis coolidgei, Osgood, Field Mus. Nat. Hist., Zool.
xviii, p. 202, 1932.
Locality of the type of assamensis, Assam, " possibly from
54 CERCOPITHECID.E.
the Mils to the northward," according to Blanford ; of
coolidgei, Hoi Xuen, Annam.
Distribution.— Assam, the Mishmi and Naga Hills ; Upper
Burma ; ? the Sttndarbaits ; Tong-king and Annam.
Tail short, usually considerably less than a third, and
always much less than half, the length of the head and body,
and only a little longer than the foot.
Anderson gave the measurements of McClelland's type,
a mounted specimen, as : head and body 26| in., over the
curves ; tail 9| in., probably including the terminal hairs.
These dimensions, making allowance for the method of measure-
ment, agree tolerably well with those of the two adult <J
specimens from the Tebang River, 2,000 ft., in the Mishmi
Hills (Wells) entered below. Unfortunately no flesh-measure-
ments were taken of a considerable number of examples from
Mokokchung, 6,000 ft., in the Naga Hills, but on the dried
skins the tails are obviously short. The young specimen
in the table from the Adung Valley, 6,000 ft., Upper Burma,
lat. 28° 10' N., long. 97° 40' E., has the tail rather more
than one-third the length of the head and body, but only
a little longer than the foot. It was collected by Lord Cran-
brook. The measurements of the two $ specimens from the
Mishmi Hills suggest also that this race is a little larger
than the next ; but the evidence is not sufficient to establish
that point.
There is nothing distinctive in the coat, which varies
seasonally from about 40 to 80 mm. ; the colour also is in-
dividually variable to a certain extent, darker or lighter
brown in accordance with the amount of yellowish- buff in
the pelage ; but an unmeasured immature specimen collected
on the Tuzu River north of Tamanthi, on the Upper Chindwin,
460ft. (McMillan), is unusually brightly tinted, almost reddish-
oehreous, on the head, nape, and shoulders, with a yellowish
wash over the hind quarters and legs. This may be a juvenile
.character, but it is not certain that the specimen belongs to
this race.
Farther information is also required about the representa-
tives of this species that occur in the Sundarbans, whence
Anderson had at least one specimen (Proc. Zool. Soc. 1872,
p. 529), as well as several examples of M. mulatto. In this
low-lying district, the exact spot being about 50 miles east of
Calcutta, might well occur a race distinct from typical
assamensis, which is found at tolerably high altitudes.
This race extends to Indo-China. Osgood described
specimens from Annam and Tong-king as representing a new
race, M. a. coolidgei, and diagnosed it as differing from
MACACA. 55
assamensis by its decidedly shorter tail, more greyish colora-
tion, and possibly heavier dentition. But he compared his
specimens with Sikkimese, not Assamese, skins ; and the
flesh-measurements he gave of two adult <J specimens show
the tail to be considerably less than half the length of the head
and body. Judging from the two specimens, including the
type, in the British Museum, I consider coolidgei to be a
synonym of typical assamensis.
6 6. Macaea assamensis pelops Hodgson.
Macacus (Pithex) pelops, Hodgson, Journ. As. Soc. Beng. xi,
p. 1213, 1840.
Macacus problematicus, Gray, Cat. Monkeys, etc., in Brit. Mus.
p. 128, 1870.
Macacus rheso-similis, Selater, Proc. Zool. Soc. 1872, p. 495.
Macacus assamensis, Blanford, Mamm. Brit. Ind. p. 15, 1888, and
of other authors (in part).
Vernacular. — Sahu (Lepcha) ; Pio (Bhotia) ; Bandar
(Pahari) ; and Panah Bandar (Darjeeling).
Locality of the type of pelops, the Nepal Kachar ; of problem-
aticus, Dhalimkot in Bhutan ; of rheso-similis, " Bast Indies."
Distribution. — The Himalayas from Mussooree through
Nepal, "the northern range of hills exclusively" (Hodgson),
and Sikkim, from 2,000 to about 6,000 ft., to Bhutan.
Distinguished from typical assamensis by the actually
and relatively longer tail, which typically is considerably more
than half the length of the head and body and about twice
the length of the foot ; also, possibly, by being a little smaller
The average lengths of the head and body and of the tail in
five adult <J examples from Sikkim are 22 § in and 13 in.,
whereas the same averages of the two adult $ examples of
typical assamensis from the Mishmi Hills are 25f- in. and just
under 8 in.
The only available specimens of this race measured in the
flesh, according to modern methods, and supplied with full
particulars of dates, altitude, etc., were collected for the
Survey by Crump and Baptista in Darjeeling and Sikkim ;
but there is no reason to doubt that these belong to the same
race as the Nepalese form named pelops and as the Bhutan
form named problematicus. At all events the types of those
two forms, as well as of rheso-similis, and all Hodgson's
Nepalese skins, have long tails like the skins from Sikkim.
The coat varies from about 55 to over 70 mm. in length
on the shoulder ; the colour is also individually variable in
accordance with the amount of buffy yellow in the pelage,
making it lighter or darker brown. An. immature specimen
56 CERCOPITHECnUE.
from Rongli, Sikkim, 2,700 ft., is brighter tinted than the
adults on the head, nape, shoulders, and arms, and is com-
parable to the specimen provisionally assigned to typical
assamensis from the Upper Chindwin Eiver in Burma, but is "not
so bright on the fore quarters, and lacks the yellow wash on
loins and legs.
Mesh-measurements (in English inches) of the two races of
this species are as follows : —
Head and
Name, locality and sex. body. Tail. Foot.
assamensis.
Mishmi Hills ; ad. <? 25f 7} 6|
Mishmi Hills ; ad. <J 25| 8 6|-
Adung Valley ; yg. t? Hf 4f 4
pelops.
Darjeeling ; ad. <J 23 12$ 6$
Darjeeling ; ad. <J 22$ 11$ 6f
Chuntang ; ad. cj 22f 13+ 6f
Dikchu ; ad. «J 22 13$ 6$
Darjeeling * ; ad. ? 20$ 15$ 6$
Rongli ; yg. <J 15$ 8$ 5J
The recorded weights do not coincide with the size. A <$
from the Mishmi Hills was 23 lb. ; the first $ on the list from
Darjeeling was 25 lb. and the £ from Chuntang 28 lb.
The measurements in the table (p. 57) suggest that <J skulls
of typical assamensis are a little larger than those of pelops.
The two skulls of assamensis from the Mishmi Hills differ
greatly in the shape of the facial profile. In the first the area
between the brow and the anterior nares is comparatively
lightly concave and long, the nasals measuring 47 mm. In
the second the area is deeply concave and short, the interorbital
septum being nearly vertical and the nasals measuring only
31 mm. But this skull was damaged during life in the
premaxillary region, and was shortened by about 5 mm.,
.and its total and condylobasal lengths, set in brackets, are
estimated.
In the case ofpehps a point to notice is the great difference
in size between the largest and smallest skulls from Darjeeling.
The former came from Sakiapokhu, the latter from Pashok.
This skull in its general form and size recalls in many respects
the skulls of M . mulatta.
Crump reports that this Macaque occurs throughout Sikkim
and Darjeeling at low elevations, its favourite zone in cold
* The head and body of this $ from the Rungbong Valley, Darjeeling,
5,200 ft., were given as 620 mm. (=24$ in.), but since the sktdl is,
considerably shorter than in the g examples from Sikkim, there is,
I think, no doubt that 620 was meant for 520, and I have assumed the
latter number to be correct.
MACACA.
57
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58 CEBCOPITHECID^!.
weather being from 2,000 to 4,000 ft. As a rule it frequents
heavy forest, where it occurs in large companies. Its ordinary
call-note is a loud " pio," rather a musical sound, and frequently
repeated. Its •warning cry is the same, hut uttered once
by the sentinel, which is generally on the look-out high up
in a tree. On hearing it the whole band descends to the
ground and moves away in absolute silence, concealed by the
dense undergrowth. It is hunted and eaten by the Lepchas,
who think its flesh has medicinal properties. Owing to
persecution on this account, and no doubt also to its fondness
for raiding maize-fields, as recorded on the label of one speci-
men, this Macaque is exceedingly shy and wary in certain
parts of Sikkim, but on the Nepal border, where it is pre-
sumably unmolested, it is much more fearless.
7. Maeaea nemestrina (Linnaeus). The Pig-tailed Macaque.
Simia nemestrina, Linn.., Syst. Nat. ed. 12, i, p. 35, 1766.
M acacus nemestrinus of most subsequent authors. (For bibliography
and synonymy of British Indian subspecies see below.)
Distribution. — From Uppee Burma and Abakan to Siam,
the Malay Peninsula, Sumatra, and Borneo.
A large or very large Macaque, the typical form the largest
of the genus, with the tail always less than half the length of
the head and body, on the average about one-third, mostly
covered with short hair throughout, sometimes with a small
terminal tuft and carried in an elevated arch above the anus.
Hairs on the crown short, radiating from a whorl and forming
a very definite " cap " differentiated from the hairs behind the
brow, on the temples, and sides of the crown by colour or
direction of growth or both combined. Hairs on fore part
of the cheek and temples growing backwards or obliquely
upwards to meet the forwardly directed hair in front of the
ear, and on the hinder part of the cheek to form a crest or
conspicuous thick fringe which curves over the top of the
ear on each side of the crown. The penis is long, with the
glans of normal form and irregularly subspherical. The
female has a conspicuous, monthly hypersemic swelling of the
external genitalia at pairing time.
The skull varies considerably in shape and size, the jaws
in the typical form, which does not enter the British Indian
area, being exceptionally long, almost " baboon-like " ; but
in the adult <J it is never so muscularly developed as in
M. assamensis and M. speciosa, having a comparatively
small occipital crest, and the temporal ridges, typically some
distance apart, seldom meeting to form a sagittal crest even
on the parietals.
MACACA. 59'
The typical form of this Macaque came from Sumatra.
It is also found, in Borneo and the Malay Peninsula, but does
not extend northwards into British Burma, although Blanford,
despite the evidence to the contrary supplied by Tickell
(see below, p. 62), ascribed the Tenasserim Pig-tail to typical
nemestrina, and gave a composite description of it derived
from two forms of the Pig-tail and from skulls of three species
of Macaca *.
Owing to the errors into which Blanford was led, partly
from want of properly localized material, I have included in
the following key to the races of British Indian Pig-tailed
Macaques the distinguishing characters of the typical form.
a. General colour much paler ; the back only
slightly darkened, much paler than the
blackish upper side of the tail ; jaws short . . leonina Blyth, p. 59.
a'. Much darker, the back deep blackish-brown,
the same tint as the top of the tail.
6. Hair, except on the back, profusely annu-
lated, general tint not so brown ; jaws [p. 62.
short, brows high blythii Pocock,
&'. Hair generally inconspicuously annulated ;
colour browner ; jaws long, brow-ridge [p. 58.
low, with sloping orbits nemestrina (Linn.),
Kloss in 1919 was the first author to refer all the described
Pig-tailed Macaques to a single species on the evidence of
a specimen from Trang in the Malay Peninsula intermediate
in colour between typical nemestrina and leonina. On this
and other evidence I entirely concur with his opinion.
7 a. Maeaea nemestrina leonina Blyth.
Macacus leoninus, Blyth, Cat. Mamm. Mus. As. Soc. p. 7, 1863 ;
id., Journ. As. Soc. Beng. xliv, p. 2, 1875 {Jnuus) ; Anderson ;.
Cat. Mamm. Ind. Mus. Calc. i, p. 71, 1881 (Blyth's type) ;
Blanford, Mamm. Brit. Ind. p. 18, 1888 (in part).
Macacus andamanensis, Baxtlett, Land and Water, viii,p. 57, 1869 i
Sclater, Proc. Zool. Soc. 1869, p. 467 ; Elliot, Rev. Primates, ii,
p. 208, 1912 (Pithecus).
Macacus nemestrinus, Blanford, Mamm. Brit. Ind. p. 20, 1888
(in part ; Tenasserim specimens which were not typical
nemestrina Linn.).
Macaca adusta and imulana, Miller, Proc. U.S. Nat. Mus. xxix,
pp. 559-60, 1906.
* The largest skull he measured is that of a menagerie-reared example
of typical M . nemestrina, the smallest, the skull of an adult <J of M. irus
aurea from Mergui (Oldham) and the skull of the alleged old $ from the
Toungyeen River (Bingham) is the skull of an adult $ of M. speciosa
(see p. 71). It is, perhaps, not surprising that on the strength of these
skulls he thought the (J and $ of M . nemestrina must be approximately
equal in size, and that there might be two species of it, a larger and
a smaller, living side by side.
60 CEECOPITHECIDJE.
Macaca nemestrina indochinensis, Kloss, Journ. Nat. Hist. Soo.
Siam, iii, p. 343, 1919.
Macaca nemestrina andamanensis, Poeock, Journ. Bomb. Nat. Hist.
Soc. xxxv, p. 302, 1931 *.
Vernacular. — Myouk-mai (Burm.) ; Myouk-la-haing (Ara-
kan) ; Myouk-padi (Burmese) ; Ta-o-ti (Burmese at Tavoy) ;
Bruh or Broh and Bruh-tana (Malay at Bankaehon).
Locality of type of leoninus, N. Arakan ; of andamanensis,
Port Blair, Andamans (introduced) ; of adusta, Champang,
Tenasserim ; of insulana, Chance Island, Mergui Archipelago ;
of indochinensis, Lat Bua Kao, E. Siam.
Distribution. — From Uppee Bxtbma to Tenassebim, the
Meegtti Archipelago, and Siam.
Coat without appreciable underwool, but longish, especially
on the shoulders, where at least in the adult $ it may form
a kind of mantle.
General colour comparatively pale, with an mdividual
tendency to erythrism, ranging from greyish-olive to buffy-
yellow, russet or golden-ferruginous, these tints being specially
noticeable on the long hairs of the shoulders, which are brighter
than the back, the lumbar and sacral regions being duskier,
* This synonymy is based on descriptions of the type of leoninus,
on the type of andamanensis, on several examples from Tenasserim
and the Mergui Archipelago, whence came Miller's examples of adusta
and insulana respectively, and on others from Siam referred by Kloss to
indochinensis. , The alleged differences between these forms seem
individual, not racial, and they all agree tolerably closely with Blyth's
description of the type of leoninus and with the independently published
descriptions of it by Anderson, Elliot, and Kloss, who saw it in Calcutta.
Elliot substituted the name andamanensis for leoninus under the mistaken
belief that leoninus was preoccupied in the genus Macaca by Simia
leonina Shaw (Gen. Zool. i, p. 34, 1800), which he relegated to the
synonymy of the species described below as Macaca silenus. In this
he was followed by Kloss ; and, unwisely, by myself, without verifying
the reference. On consulting Shaw's work I find there is no occasion
for the change. The name leonina was given by Shaw to a monkey,
described by Buffon, and believed to have come from Abyssinia, which
agrees with no species of Macaca, and differs essentially from M . silenus
in the tail being longer than the head and body, the former measuring
27 in., the latter 24 in., the precise dimension attesting their derivation
from an actual specimen. I am unable to identify Simia leonina,
but its general features, apart from the long beard, agree better with
John's Leaf-Monkey (Kasijohnii) than with M. silenus.
It is satisfactory to resuscitate the appropriate name leonina for the
Burmese Pig-tailed Macaque and drop the inappropriate name andaman-
ensis given to a specimen not indigenous in the Andaman Islands.
According to Hamilton (Proc. Zool. Soc. 1870, p. 220) it was one of
several imported from the adjoining mainland of Burma. Quite
possibly it came from Akyab in Arakan, the home of typical leonina.
At all events Tickell, in his MS., said that the Pig-tailed Macaque,
which he knew in Tenasserim, inhabited the hills to the interior of
Akyab, and was exported alive from that seaport and taken to Chitta-
gong. Equally likely some were taken thence to the Andamans.
MACACA. 61
especially in the middle lines ; hairs of the upper side, as a rule,
profusely annulated with buff, yellow or russet, and dark
brown or black bands. These vary in distinctness, being less
conspicuous on the hind back and legs than on the shoulders
and arms, and may be quite inconspicuous, especially in the
old faded coat before the moult sets in. Crown the same tint
as the nape or darker ; cheeks greyish or buffy, the hairs
forming a longish and thick apically annulated fringe. Under-
side greyish-white, with some darker annuJations on the
abdomen and the inner side of the limbs in some specimens.
Buttocks grey or buffy, sometimes indistiuctly annulated,
sometimes sharply contrasted with the back. Tail usually
with whitish fringe at the base on each side ; its upper side
dark and contrasted with the paler hue of the loins.
Flesh-measurements (in English inches) are as follows : —
Head and
Locality and sex. body. Tail. Foot.
Bankaehon, Tenasserim ; ad. <J 23 7£ 6f
Champang, Tenasserim ; ad. <J : 22§ 9£ 6i
Bed Point, Tenasserim ; ad. $ 20 7J- 6£
Sullivan Island, Mergui ; yg. ad. <J . . . 21 7 6
Chance Island, Mergui ; ad. $ 20| 7+ 5f
Chance Island, Mergui ; ad. <J 20£ 6J 6—
Bankaehon, Tenasserim ; ad. $ 19f 5f 5f
H'Kamti, Upper Burma ; ? 18$ 7J 5|
The measurements of the specimens from Champang and
Red Point, Tenasserim, and from Chance Island are taken
from Miller's records of adusta and insukcna respectively.
The specimens from Bankaehon and H'Kamti were collected
and measured by Gr. C. Shortridge, and the one from Sullivan
Island by C. Primrose. The tail varies from considerably
more to a little less than the length of the foot.
Miller's £ specimens from Champang and Red Point
weighed 20 lb. and 14 lb. respectively, his two from Chance
Island 13| lb. ; the <J from Bankaehon was 18 lb. and the
$ 10J lb. The <J specimens, although not very much shorter
in head and body, were much lighter than three adult males
of the typical form from Sumatra, which were 25, 27, and
30 lb. respectively, as recorded by Miller.
The skulls of this Macaque are about the same size as those
of M . mulatto, villosa True, and I am unable to find any constant
difference between the two, particularly in the $ skull*,
but the cj skull seems to have slightly more elevated brows
* By an error in registration two skulls carry the same number as
the skin of the 9 andamanensis collected by Shortridge at H'Kamti
on the Upper Cbindwin. One is the skull of M. mulatto,, the other of
M . nemestrina leonina, and I do not know which of the two belongs
to the skin.
62 CEBCOMTHECID-a:.
and stronger temporal ridges. The skull of the adult c? from
Bankachon is, unfortunately, not available ; but the skull
of an old $ from Tavoy figured by Tickell in his MS. has the
ridges strong and converging posteriorly, only about 7 mm.
apart on the parietals, -with a short median ridge between
them. In the skulls of typical nemestrina that I have seen
the ridges are farther apart, and this is accompanied by longer
jaws. But the difference in size between these two races
is not always very great. For instance, the skull of an adult $
>of nemestrina from Perak is only 143 mm. in total length,
but two from Pahang and Selangore are 151, a third from
Borneo is 161, and an adult $ from Sumatra is 134 mm.,
indicating considerably greater average length in nemestrina.
(For measurements of leonina, see p. 64.)
Tickell * in his MS. recorded this Macaque from Ye in Amherst
where it occurs in the thick jungles at the foot of the hills,
but is not nearly so common as the " fishing monkey "
(M. irus aurea), although it seems to increase in numbers
farther south in Tenasserim. The voice of the young is the
same, he says, as that of the Rhesus (M . mulatto). Shortridge,
who was acquainted with it at Bankachon, says it is plentiful
there, though not to the same degree as the Gibbon (H . lar
entelhides Geoffr.) and the Leaf-Monkey {Trachypithecus
obscwus Reid). It is gregarious, and when occurring round
villages is said to be very destructive to the rice-fields. He
found the cheek-pouches filled with pieces of this cereal.
With the specimen Shortridge collected at H'Kamti was
a note stating that it was secured on the west bank of the
Chindwin River and probably represented a hill-form, possibly
common enough round Sarameti and other hill regionsf .
7 5. Macaca nemestrina Wythii Pocock. Blyth's Pig-tailed
Macaque.
Macacus leonirvus, Sclater, Proc. Zool. Soo. 1870, p. 663, pi. xxxv
(ij) ; Anderson, Zool. Res. Yunnan, 1878, p. 52 (fig. skull) ;
Blanford, Mamm. Brit. Ind. 1891, p. 18 {$, fig.). Not M. leo-
nirms, Blyth.
* This able naturalist, to whom Blanford was much, indebted, published
an admirable description, illustrated by good coloured figures, of a £
and two young specimens of this Macaque. There is also an accurate
figure of the skull of a very old ^. The description and the figures
agree as precisely as can be with the description of leonina given above.
If Blanford, who saw this account, had paid heed to it, he might have
•seen that the Pig-tail of Tenasserim is not typical nemestrina, and that
its skull has the characters he regarded as distinctive of his leoninus.
f Anderson's accounts in 1878 and 1881 of the specimens of M. leoninus
and M . assamensis he said he collected near Bhamo are so confused that
I cannot make up my mind whether he secured a $ of each or of only
one of the species, and, if only one, which it was.
MACACA.
63
Pithecus andamanensis, Elliot, Bev. Primates, ii, p. 208, 1912 (in
part., fig. of <J).
Macaca nemestrina andamcmensis, Kloss, Journ. Nat. Hist. Soo.
Siam, iii, p. 344, 1919 (skull only).
Macaca nemestrina blythii, Pocoek, Journ. Bomb. Nat. Hist. Soe.
xxxv, p. 305, 1931.
? Macaca nemestrina, McCarm, Journ. Bomb. Nat. Hist. Soc. xxxvi,
p. 808, 1933.
Vernacular. — ? Kangh (Nagas).
Locality of the type, unknown.
Distribution. — Unknown, but probably some district of
British India east of the Ganges ; 1 Naga Hills in Assam.
Distinguished from n. leonina Blyth by the blackish-brown
tint of the crown and of the back behind the shoulders, which
%^-^T—rz
Fig. 15. — Blyth's Pig-tailed Macaque (Macaca nemestrina blythii).
(Prom a drawing by Wolf of the type when living in the Zoological
Gardens, London.)
resembles the tint of the upper side of the tail, resembling
in these respects typical nemestrina, but distinguished from it
by the generally profuse, conspicuous annulation of the
pale hairs, and by the shortness of the muzzle, the pronounced
brow-ridge, nearly vertical interorbital septum, and other
characters of the skull.
Coat full and longish, with well-developed facial ruff.
General colour dark, uniformly blackish-brown on the
crown, the back behind the shoulders, and the upper side of
the tail ; the nape, shoulders, and fore back paler, annulated
orange-red and black ; cheeks dusky grey, annulated, with the
tips of the hairs of the ruff black. Towards the root of the
64
CEECOPITHBOID M .
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MACACA. 65
tail, which is white below, with a white basal tuft on each
side, the bases of the hairs turn grey, the hairs beneath the
callosities on the buttocks white ; arms and hands paler
than shoulders ; legs dark olive ; underside greyish and
conspicuously annulated on the chest, browner, unannulated
on the belly.
This description is taken from the type and only available
specimen. The dimensions (in English inches) of the dried,
made-up skin are : — Head and body 24±, tail 8 J, foot 6.
The size is about the same as in the adult <J of n. leonina.
The skull, well figured by Anderson, is about the same length
as that of n. leonina and has similar short jaws ; but the
cranial portion is relatively short, owing to the unusual
uptilting of the occipital plane. It is exceptionally massive,
with high thick brow-ridges and strong temporal crests about
16 mm. apart on the frontals, but coalescing on the parietals
to form a low, thick, sagittal crest, a feature not observed
in any skull of n. leonina or typical nemestrina. These
features, however, may be a question of age, the skull being
that of an old animal.
This monkey has a curious history. It was imported to
the Zoological Gardens in London in 1870 as a full-grown
animal and was there seen by Blyth, who identified it as his
leoninus, overlooking the marked differences in colour between
it and his type of leoninus from Arakan. Sclater naturally
accepted this determination and was followed by Anderson
and Blanford. Sclater published a coloured plate of it, with
the type of andamanensis in the background, and considered
them to represent the sexes of the same species. Later
Anderson saw Blyth's type of leoninus in Calcutta and
described it, as well as this specimen, without commenting
on the differences between them. These were detected by
Kloss, who was naturally completely puzzled by the discrep-
ancies.
There is no proof that the type of blyfhh falls within the
limits of the fauna of British India. But Elliot says that he
saw a living specimen in the Calcutta Zoological Gardens
which resembled it. Not improbably this specimen came
from some nearby district, possibly northern Burma or Assam.
I have tentatively suggested that the specimens named
M . nemestrina which McCann collected at Changchang Pani,
500 ft., in the Naga Hills, may represent this race, because
he says they have much longer and darker coats than a speci-
men from Bankachon sent to Bombay as M. adusta, the
hairs being tipped with brown, especially on the hinder dorsal
region. He had three adult ?$ and one immature <J, which
in the flesh resembled Sclater's figure of M. leonina, the
. vol. i. *
66 CBECOPITHECIDJE.
reference presumably being to the (J in the foreground. His
description suggests that they are not as dark as the old $,
the type of blythii, but the difference may be a matter of sex
and age. His largest $ measured : head and body 22£ in.
tail 7f, foot 6. Two dimensions of its skull are entered in
the list below.
Assuming the correctness of this identification, M . n. blythii
inhabits, according to McCann, the same forests in the Naga
Hills as M . speciosa, but is far more arboreal and keeps strictly
to dense evergreen forests, 'which makes it difficult to observe
and secure. This may account for the comparative rarity
of specimens of this race in museums and for its having been
previously, so far as I am aware, unrecorded from. Assam.
8. Macaca silenus (Linnaeus). The Lion-tailed Macaque.
Simia silenus, Linn., Syst. Nat. ed. 10, p. 26. 1858 ; Schreber, Saug. i,
p. 87, 1775.
Macaous silenus of most subsequent authors, including Thomas,
Proc. Zool. Soo. 1911, p. 126.
Simia (Cereopithecus) veter albibarbatus and Silenus albibarbatus,
Kerr, Anim. Kingd. p. 64, 1792.
Simia ferox, Shaw, Mus. Leverian. ii, p. 69, 1793.
Vernacular. — Shia bandar (Hindi) ; Nil bandar (Beng.) ;
Nella manthi, Chingala (Mai.) ; Singalika (Can.) ; Karingode
(KLurg.) ; Kondamachu (Tel.) ; Kurankarangu (Tamil) ;
Carapu corongu (Nelliampathi).
Locality of the type of silenus, " Ceylon " * ; albibarbatus,
" Ceylon " ; ferox, " Ceylon."
Distribution. — S.W. India, the Westebn Ghats, principally
of Tkavastcore and Cochin, but alleged to occur from about
14° N. lat. to Cape Comorin.
Distinguished from all the other species of Macaca by the
combination of two characters : the growth of the hairs on
the temples and cheeks to form a long, thick, dark grey or
brownish-grey ruff or mane, which hangs down on each side
of the face like long whiskers and passes on to the throat as
a shorter, paler beard, and by the shining, black hue of the
* Nearly all the early records of this species give Ceylon as its locality,
and many of them, including the popular natural histories, cite it in
consequence as the " Ouanderu " or " Wanderoo," the vernacular
name for the so-called " Purple-faced Leaf -Monkey " or Langur of that
island. One or two comparatively recent authors have objected to the
use of silenus as its specific name because of the discrepancies between
Linnssus's description of silenus in the 10th and 12th editions of the
' Systema.' But I agree with Thomas that the description of 1758
is sufficiently exact to justify Sehreber's allocation of the name in 1775,
and the acceptance of it by nearly all recent zoologists, Elliot being an
exception, in calling it albibarbata. For the erroneous application of the
name leonina to this Macaque see p. 60, footnote.
MAMMALIA.
PLATE IV.
Photo D. Seth-Smith.
Lion-tailed Macaque (Macaco siknus).
MACACA.
67
rest of the head, body, limbs, and tail, although the under-
side, at least in the young, may be dark brown or greyish-
brown.
Despite the striking and distinctive appearance of this species ,
there is no doubt that it is not very remotely related to<
M. Tiemestrina leonina. The arrangement of the hairs on
the crown and cheeks is the same, and the huge mane or ruff
is merely an exaggeration of the shorter ruff seen in the Burmese
Pig-tailed Macaque. In both of them, also, the coat is in
general long and full, and the hairs of the greater part of the
tail are short, although in M. silenus the terminal tuft is
apparently always present and larger, and the tail itself is
longer typically, apparently about two-thirds the length of
Kg. 16. — Head of Lion-tailed Macaque (Macaca silenus).
the head and body and about twice as long as the foot. Addi-
tional resemblances to M . nemestrina are found in the structure
of the penis in the $ and the occurrence of the catamenial
swelling in the §, as recorded in my paper (Proe. Zool. Soc.
1926, p. 1571).
The following are some recorded measurements (in English
inches) : —
Locality arid sex.
Loc. unknown (Anderson) ; ad. $
Loc. unknown (B.M.) ; ad. $ . .
Loc. unknown (Elliot) ; ad. <? . .
Travancore (Bourdillon) ; ad. <J
Travaneore (Bourdillon) f ad. (J
Cochin (Riley O'Brien) ; yg. <J .
Travancore (Bourdillon) ; ad. $>
Travancore (Bourdillon) ; ad. $
1-2
Head and
body.
Tail.
Fo
24
10
—
23±
Hi
7-
21J
15£
n
21
13£
—
20
15
—
17$
10£
5
18
12£
—
18
10
—
68
CERCOPITHECID^B.
Of these the only one known to have been measured in the
flesh is the young specimen from Cochin collected by the
Survey. The records of the Travancore specimens were
quoted, by Blanford. They may be flesh-measurements,
but the dimensions given by Elliot were admittedly taken
from the made-up skin ; and there is very little doubt that
Anderson's were also. The tail of Anderson's specimen is
relatively very short, suggesting that the head and body
may be stretched. The records, for what they are worth,
show that M . silenus is about the same size as M . nemestrina
lamina.
The skull* of this Macaque is little known. Anderson
described one of an adult $ as characterized by its widely
bulging zygomatic arches, greatly developed orbital ridges, etc.
The only adult skull I have seen is that of a $ from Cochin
Pig. 17. — Skull of adult <J Lion-tailed Macaque (Maaaca silenits),
with occipital region restored, x £. From Cochin.
presented by Sir Frank Colyer, which, unfortunately, has its
occipital area broken away; hence the total and condylo-
basal lengths entered on the table (p. 64) were estimated;
but it agrees with Anderson's skull in the expanded zygomata,
prominent brow-ridges, and other points. The temporal
ridges are well defined and 13 mm. apart on the fronto-
parietal suture, but converge, without apparently meeting,
on the parietals. The muzzle is longer than in the type of
M . n. blythii ; but the facial profile is very similar to that of
the subadult <J of M . n. leonina from Sullivan's Island.
In the Nelliampathi Hills, according to B>. P. Eonloch,
this Macaque is rare and occurs in small troops up to about
* The photographs which Elliot published to illustrate the skull of
this species were taken from an adult <$ skull of the Bonnet Macaque
(M. radiata) from the Nilgiri Hills in the British Museum.
MACACA. 69
a dozen. It is sparingly distributed throughout the evergreen
forest and is a quiet monkey, its call-note being a deep " 0."
In 1859 this Macaque was recorded by Baker (Journ. As.
Soc. Beng. xxviii, p. 238) as occurring in the Western Ghats as
far north as Goa, but only in the lonely dense forests. In Tra-
vancore, according to Hill (Proc. Zool. Soc. 1937, Syst. p. 215),
it is not found below 2,000 ft., its altitude being usually from
2,500 to 3,000 ft. Like other Macaques it is gregarious,
living in parties of from twelve to twenty individuals or more.
In the rest of its habits it no doubt resembles its congeners.
Baker compared the call of the <J to the " cooyeh " of a native
astray in the forest and trying to keep in touch with his
companions. O'Brien secured for the Survey only one half-
grown specimen at Cotengady 3,500 ft.. in Cochin, and tersely
summed up his impression of the species by writing on the label
" rare, shy, and silent." These attributes help to explain
the comparative paucity of specimens in the museums of the
world.
The general likeness in colour between this Macaque and
John's Leaf-Monkey, both being alike mainly black with
brownish whiskers, is worth noting as an independently
acquired characteristic, probably in response to similarity
in habitat.
9. Maeaea speeiosa Cuvier. The Stump-tailed Macaque.
[Macactts] speciosus, F. Cuvier, Hist. Nat. Mamm. no. 46, 1825.
Inuus speciosus, Blyth, Cat. Mamm. Burma, p. 6, 1875, and of
most recent authors under Macaca or Lyssodes.
Macacus arctoides, Anderson, Zool. Bes. Yunnan, p. 45, 1878 ;
Blanford, Mamm. Brit. Ind. p. 17, 1888. (For further bibliography
and synonymy see under subspecific headings.)
Distribution. — Tibet, South China, Assam, Bttbma, Malay
Peninsula, Siam, and Indo-China.
Distinguished from all the British Indian species by the
shortness of the tail, which is reduced to a conical, sparsely
hairy stump, not more than one-third the length of the foot,
and by the excessive elongation of the glans penis. The hairs
on the crown, radiating from a central whorl, are elongated
laterally and posteriorly over the occiput, but in the adult are
very short anteriorly, leaving a nearly naked area behind
the brow-ridges ; hairs on the cheeks sweeping backwards, con-
cealing the ears, and downwards to the throat, where they
meet the hairs growing upwards from a whorl behind the chin,
and forwards along the side of the neck, forming with them
a tufted crest. Typically, at all events, there is an area of
naked skin round the callosities. General colour usually
dark brown, but varying from nearly black to reddish, with
the underside at most a little lighter than the upper. Face pink.
70 CEBCOMTHECIDiE.
la the shortness of the tail M. speciosa closely resembles
the Macaque from Celebes, M . maura. The tail is also tolerably
short in the Japanese Macaque, M . fusGata ; but it is still
more reduced than in these three species in the so-called
" Gibraltar Ape," M. sylvana, which is indigenous in Morocco.
But M . speciosa differs fundamentally from these species, as
it does from all the other species of Cynomorph Primates,
in the structure of the glans penis, which is remarkably long,
gradually tapers to a slightly upturned point, and is supported
by a correspondingly long bone, the baculum. This peculiarity
was first described by Anderson (Proc. Zool. Soc. 1872, p. 209)
and later by Murie in the same volume (p. 784), who, however,
wrongly identified his specimen as a Japanese Macaque.
The systematic importance of this structure was first insisted
Fig. 18. — Skull of adult <? Stump-tailed Macaque (Macaco, speciosa
speoiosa) from the Dikchu River, Naga Hills.
on by de Beaux (Giorn. Morf. dell' Uomo e del Primati, i,
pt. 1, 1917) and independently by myself (Ann. Mag. Nat.
Hist. (9) vii, p. 224, 1921, and later, Proc. Zool. Soc. 1925,
p. 1557). If M . speciosa be regarded as representing a distinct
genus or subgenus on account of the peculiarity of the penis,
the name available for it is Lyssodes Gistel, 1848.
Considering the great differences between this species and
M. assamensis McClell. in the length of the tail and the struc-
ture of the glans penis, it is perhaps surprising that the skulls
of the two are not easily distinguishable, that of speciosus
having similar massive brows and orbital margins, well-
developed occipital and sagittal crests, large upper canines, etc. ;
but it is on the whole more massive, and has the outer edge of
the orbits more vertical, straighter, and less concave where
they pass into the anterior root of the zygomatic arch.
MACACA. 71
Many specific or subspecific names have been given to
Macaques with the short tail and crimson face, described by
Cuvier as characteristic of the type of speciosus. There is
no doubt that all belong to that species, but the number of
subspecies is as yet unsettled. I provisionally assign the
British Indian representatives to two subspecies, a northern
and a southern.
9 a. Macaca speciosa speciosa Cuvier.
[Macacus'] speciosus, F. Cuvier, Hist. Nat. Mamm. no. 46, 1825.
Macaous brunneus, Anderson, Proc. Zool. Soc. 1871, p. 628, and
1872, p. 203 ; also Zool. Res. Tunnan, p. 45, 1878, where
brunneus is dropped as a synonym of arctoides.
Macacus (Magus) arctoides melli and esau, Matsohie, SB. Ges.
Nat. Fr. Berlin, 1912, p. 308.
Pithecus pullus, Howell, Proe. Biol. Soc. Wash, sli, p. 41, 1928.
Lyssodes speciosus melli, G. M. Allen, Amer. Mus. Novit. no. 429,
p. 3, 1930.
Macaca speciosa, Osgood, Field Mus. Nat. Hist. Zool. xviii, p. 202,
1932 ; McCann, Journ. Bomb. Nat. Hist. Soe. xxxvi, p. 797,
pis., 1933.
Vernacular. — Chantee (Nagas).
Locality of the type of speciosa, " E. Indies " ; of brunneus,
Kakhyen Hills, east of Bhamo ; of melli and esau, west of
Lo-chang, in western Kwantung ; of pullus, near Kuatun
in Fukien.
Distribution. — Assam, Upper. Bubma, S. China, Tong-king,
and Annam.
Coat in winter very long, on the shoulders up to about
112 mm. or more (4Jin.). Colour above varying from deep
olivaceous-grey to chocolate-brown, the hairs, at least in the
olivaceous-grey phase, speckled, but obscurely, with dull
buffish annulations ; the underside paler, greyish-brown to
brown.
Cuvier described speciosus from a coloured drawing that
was sent to him of a specimen in the Barrackpore Menagerie,
near Calcutta, and Blyth was the first to adopt this name for
specimens from Cachar and the Kakhyen Hills, east of Bhamo.
Anderson entirely agreed with the correctness of Blyth's
opinion, adding that the Macaque illustrated was probably
locally caught either in Assam or Cachar ; but for no intelligible
reason he chose for the species the later name arctoides, given
by Geoflroy to an example he received from Cochin China.
Blanford also adopted arctoides, because he followed Tem-
minck's erroneous and highly improbable view that the
type of speciosus was a Japanese Macaque (M. fuscata).
I unhesitatingly follow Blyth's verdict and Anderson's reasoning,
with the additional reason that specimens from the Naga Hills,
described below, agree better with Cuvier s description of the
colour of the upper side of speciosus as "grisvineux " than
72 OEECOPITHECID^!.
most examples of the species, although they are not so grey
below and on the inside of the limbs ; but a reproduced coloured
drawing cannot be implicitly relied on.
Anderson's type of brunneus from the hills on the western
border of Yunnan, a young specimen, had the coat long,
thick, and woolly, the colour dark brown above, washed with
yellowish below. Specimens he received later varied somewhat
in tint, and one that was kept alive acquired a speckled,
annulated coat during captivity. He also had an example
from Cachar, just south-west of the Naga Hills. I can find
no reliable characters, embodied in the descriptions, by which
the southern Chinese specimens named melli, esau, and pullus-
can be distinguished from brunneus.
The only available British Indian examples of this race
were collected by J. P. Mills in the Naga Hills, an adult $
on the Dikho River, 21 February, and an adult $ at Merang-
kong, 15 January. The <J has the hairs on the shoulder about
118 mm., and the general colour dark greyish-olivaceous above,
with the hairs obscurely annulated blackish and dull buff;
the crown has a grey hoary tinge, the limbs are a little browner
and paler than the back, there is some redder brown on the
tail and round the callosities, and the underside is also browner
and paler than the upper. The $ has the shoulder-hairs
"about 110 mm. ; the general colour is very similar, but the
flanks and underside are a trifle browner, and the crown
not so hoary.
In a good series of skins which he collected at Changchang
Pani in the Naga Hills, 500 ft., McCann traced the colour-
change from the newly-born young to extreme old age.
The general colour of the newly-born young is mainly creamy-
white, with a light brown tint on the rump and some brown
patches elsewhere. The brown increases rapidly in extent
and intensity. In the juvenile the tint is all brown. In the
adult the coat becomes speckled owing to a varying number
of pale annulations, usually about eight to ten, on the indi-
vidual hairs. In old age the .colour is more grizzled, the pale
annulations turning grey. In the skin of one aged <J there
were as many as sixteen annulations on the individual hairs
and some wholly grey hairs mixed with the rest. These
observations confirm Anderson's statement that his brown
example of brunneus became speckled during the time it was
in captivity.
McCann also referred to an adult specimen from the western
slope of the Divide between the Salween and the Irrawaddy,
5,000 ft., which was more ferruginous along the flanks than
in the Naga Hills skins.
Two adult $ skins from Chapa in Tong-king and Col de
Nuages, Annam (Delacour and Lowe), are very like my two
MACACA. 73
skins from, the Naga Hills, but the hair of the shoulders is
only about 75 mm. long, and the Annam skin has a lot of
long grey hairs on the crown and nape.
9 b. Maeaea speciosa melanotus (Ogilby).
Papio melanotus Ogilby, Proc. Zool. Soc. 1839, p. 31.
Macacus rufescens, Anderson, Proe. Zool. Soc. 1872, p. 204.
Macacus harmandi, Trouessart, Le Natural. 1897, p. 10.
Piihecus rufescens, Robinson & Kloss, Ann. Mag. Nat. Hist. (8)
xiii, p. 392, 1914.
Locality of the type of melanotus "said to be Madras" ; of
rufescens, " Singapore " ; of harmandi, Chantaboun, S. Siam..
Distribution. — Teitasseeim, the Malay Peninsula, and
Siam.
Distinguished from typical speciosa by its thinner, shorter
coat, up to about 70 mm. (less than 3 in.), in winter, by its
blacker or redder colour, and by the absence or more obscure
indications of pale speckling in the dorsal pelage.
The type of melanotus Ogilby, which was probably shipped
from the Malay Peninsula or Tenasserim to Madras, is mostly
glossy blackish above, browner on the flanks, and still browner
below, with the limbs a little lighter than the back, and with
scarcely a trace anywhere of pale speckling in the pelage.
It is a half -grown <J, with the hair on the forehead not con-
spicuously short as in the adult. The dry skin of the face is
quite pallid, showing that the specimen cannot be assigned
to the short-tailed black Celebean Macaque, M . maura, which
has a black face. The type of rufescens, bought alive in Singa-
pore, may have come from almost anywhere in the East,
but it so closely resembles some skins from the Malay Peninsula
that that country may be taken as its home. Like the type
of melanotus, it is a half -grown <J, with the hair on the forehead
longish, but differs in its generally brightish red colour above,
with some black-tipped hairs on the head and nape and
brighter brown hue below, the pale speckling in the hairs
being negligible. The type of harmandi does not seem to
differ in any way from that of melanotus. It was described
as very deep glossy brown, approaching black above, reddish-
brown below, with the face carmine, apart from the blackish
muzzle. The young was said to be tawny grey.
Although this Macaque has been recorded from districts
in Tenasserim as remote as the Toungyeen River, a tributary
of the Salween, in the north and Bankachon, Victoria Point,
in the south, the only available skin is that of a subadult <J
collected by H. Cecil Smith in Tavoy, December. The eoat
is about 70 mm. on the shoulder and everywhere thinner and
shorter than in the Assamese skins, with the hairs browner
at the base, a little darker, less olivaceous, and less annulated
74 CBEOOPITHBCIDiE.
distally ; there is no appreciable reddish-brown on the but-
tocks, but the crown is a trifle greyer. Bingham saw black
specimens at the foot of Mt. Mulai-yit, and a couple of obviously
young individuals, cream in colour, with a rusty tinge above,
were recorded by Davison from Bankachon (see p. 77).
These were probably not very different from the " tawny
grey " young of harmandi mentioned by Trouessart.
From the Malay Peninsula, in addition to the type of
rufescens, the British Museum has several skins collected at
Trang by Robinson and Kloss in December and January.
One adult $ is glossy black above, with hardly a trace of
annulations and no grey on the crown or brown on the rump.
A younger $ is like it, but browner, and a $ is still browner.
A young $ from Patalung (Skeat Expedition) is tolerably
uniformly rufous-brown, yellower on the crown and on the
limbs below the elbows and knees, closely approaching the
type of rufescens ; and a young $ from Nangkok Grabi,
Siam (Kloss), is very like the $ from Trang. The coat in these
skins is from about 55 to 70 mm. long. They were identified
as M . arctoides rufescens by Robinson and Kloss, who declared
that the reddish and brown colour was characteristic of $
and young ^ specimens. There is, however, the skin of an
adult (J killed in April at Hat Sanuh, near Koh Lak, Rajburi,
S.W. Siam, of special interest from the closeness of its likeness
to the skins from the Naga Hills assigned to typical speciosa.
The coat is a little thinner, in no sense woolly, but quite as
long, about 100 mm. on the shoulder ; the colour is only
a trifle browner and less olivaceous, but the hairs are a little
more conspicuously speckled with redder annulations. This
skin comes nearer typical M . speciosa arctoides, provisionally
regarded as a distinct race, than any I have seen (cf. infra,
p. 75).
The only flesh-measurements of British Indian specimens
of speciosa with which I am acquainted are those recorded
by McCann in his valuable paper. To these I have added the
dimensions (in English inches) of a few from other districts
belonging to both races : —
Head and
Name, locality and sex. body. Tail. Foot.
speciosa.
Changchang Pani, Naga Hills ; ad. <J . 25 J 2§- 7 —
Changehang Pani, Naga Hills ; ad. $ . 23£ If 7f
Changchang Pani, Naga Hills ; ad. (J . 22f 3 6f
KfiatBX^FT^deii (pullvs type); yg.ad. $. 24 2% 7 +
Changchang Pani, Naga Hills ; ad. $ . 21J- 2§ 6i
Col de Nuages, Aimam ; ad. $ 22$ £ 5|
Chapa, Tong-king ; ad. $ 21f if 5|
melanotus.
Koh Lak, Rajburi, S.W. Siam ; ad. <J . 21f If 7
Trang, Malay Peninsula ; ad. £ 19£ 2 6f
MAOACA. 75
These dimensions suggest that typical speciosa is larger
than melanotus, since the adult <J of the latter is about the
size of the $ specimens of the former, and smaller than the
young adult $ from Kuatan ; but that conclusion is not borne
out by the skulls (table, p. 76).
The female skull of speciosa from the Naga Hills, received
from Merangkong, without a skin, I identify as speciosa
because it has broad orbits, and a skin of this species was
collected at that locality, as recorded above. The skull of
melanotus from the Toungyeen River is the one Blanford
identified as an old $ of M. nemestrinus collected by Bingham
in the Meplay Valley. It has no skin, but is unquestionably
that of an old. $ M. speciosa, and the largest recorded.
The type of Macacus arctoides Geofiroy (Zool. Voy. de
Belanger, p. 61, 1831, and Mag. de Zool., Mammif. CI. i, pi. 2,
1833), from Cochin China, apparently represents a subspecies of
M . speciosa. It was said to be an adult <J, with the head and
body about 32 in. long, and to differ from the type of speciosa
in its conspicuously annulated coat. The annulation was
red and black, yielding a brownish-red general hue. On this
account I provisionally regard arctoides as a distinct subspecies
from melanotus, in which there is evidence that the adult <J
is black or nearly black above and unannulated, and the
annulations hardly appreciable even in the browner or redder
$ and young <J. It was the blackish, unspeckled colour of
harmandi that induced Trouessart to describe his Chantaboun
skin as representing a distinct race from arctoides. Gervais
substituted the name ursinus for arctoides (Hist. Nat. Mamm.
i, p. 93, 1854).
Geoffroy's description of atctoides suggests that it more
nearly resembles speciosus than melanotus. This is not
unlikely, since speciosus occurs in Annam just to the north
of Cochin. If the two are inseparable, arctoides comes into the
synonymy of speciosus.
The records* by Bingham and Davison of the occurrence
of this species both at the foot of Mt. Mulai-yit in Tenasserim
ajid at 7,000 ft., showing that it is found at comparatively
low and high altitudes above sea-level, were confirmed by
McCann's statements that in the Naga Hills it ranges from about
* Communicated in litt. by the observers to Blanford, who inserted
them under his account of the Lar Gibbon, being under the impression
at the time that they referred to an undeseribed species of Anthropoid
Ape. Later in his volume he surmised that they might apply to a species
related to his M . arctoides, but bigger. His estimate of the size was
derived from Davison's guess that one he saw standing erect was
ebout 4 ft. high and Bingham's guess that an old § sent to him would
have been 3J ft. high upright.
76
CEKOOPITHECID-aii .
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MACACA. 77
500 to 1,800 ft., and was taken at 5,000 ft. on the Salween-
Irrawady Divide. As might be expected from its abbreviated
tail, this Macaque is mainly terrestrial. On the label of one
of Mills's skins from the Naga Hills is written : " A ground
monkey which rarely climbs, and is found in bands which may
number 50 or more individuals." McCann says the parties
consist of twenty-five or thirty individuals of both sexes and
■all ages, apparently under a leader, but that occasionally
solitary old males, probably expelled from the troop, may be
seen. The party, however, seen by Bingham, as reported
by Blanford, at the foot of Mt. Mulai-yit, and described as
" large tailless black apes," numbered only four or five.
These, like the specimens recorded by Davison as inhabiting
a very dense part of the forest at 7,000 ft. on the mountain,
were on the ground, and McCann confirms these accounts,
stating that the species is found in dense forests and is essentially
terrestrial, feeding mainly on the ground and making its way
rapidly through the thick undergrowth when alarmed, and only
climbing trees when pushed or to feed on fruits or foliage.
The alarm-note is a short, harsh bark ; but, when feeding, the
bands keep up a continuous squealing and chattering and are
art all times apparently extremely noisy. They are also
exceptionally fearless and frequently show fight when driven
from potato fields and other cultivated crops, which they are
addicted to raiding. Their diet consists mainly of leaves,
fruits, and roots, but McCann thinks that they are probably
■omnivorous — and. no doubt rightly, because two young
specimens* kept alive at Bankachon by Davison were
" excessively insectivorous, preferring insects to fruit or
bread."
Davison made the interesting observation that they exhaled
a peculiarly foetid odour. This, too, was confirmed by McCann,
wno found that none of his party would tolerate nursing
a young captured specimen because of its unpleasant smell,
which adhered to clothing. These accounts suggest that
the odour, evidently much more potent than that of other
Macaques, which are frequently petted when young, may
be protective, rendering this species distasteful. It possesses,
at all events, two other attributes often associated with dis-
tastefulness, namely fearlessness and noisiness. The odour
no doubt emanates from the skin and is probably restricted
to it, since, according to McCann, some Nagas will eat the
flesh.
* Referred to by Blanford under his account of M. arctoides.
Accepting Davison's almost certainly erroneous statement that they had
adult dentition, although standing only about 15 in. high, Blanford
supposed they represented an undescribed species.
78 CEBCOMTHECIDJE.
10. Macaea Irus Cuvier. The Crab-eating Macaque.
Siima cynamolgus, Sehreber, Saug. i, p. 91, 1775 (not S. cynamolgus
Linn.).
Macacus irus, Cuvier, Mem. Mus. Hist. Nat. Paris, iv, p. 120, 1818
(substituted for cynamolgus, given by Linnaeus to an African
Baboon).
Locality of the type unknown — said to be W. Africa *.
Distribution. — From. Loweb Bukma eastwards to Borneo
and the Philippine Islands.
Superficially resembling M. radiata (Geoffi.) and M. sinica
(Linn.) in size, proportions, and the length of the tail, which is
at most only a little shorter than the head and body, short-
haired and tolerably cylindrical throughout, but distinguished
by the hair-growth of the head, the hairs of the crown typically
growing backwards from the brow, as in M. mulatto, (Zimm.),
Kg. 19. — Skull of adult $ Crab-eating Macaque (Macaea irus aurea)
from Tavoy.
and only very rarely showing signs of a parting or whorl, and
then never exhibiting the very definite radiating arrangement
constituting the "cap " or "toque" of the S. Indian and
Ceylonese species. The whiskers also are typically much
more bushy. But the greatest difference lies in the structure
of the penis, which is small, and has a small, subspherical
glans quite unlike the large, elongated glans of the other two ;
but, as in them, there is no periodic catamenial swelling in the
$ at pairing time.
* The type oicynamolgus Schreb. was a specimen, figured and described
by Buffon, said to nave come from Senegal or Guinea. Cuvier accepted
this locality when he substituted irus for cynamolgus in 1818 and when
he figured ctynamolQus in Hist. Nat. Hiram, pis. 30 and 31, 1819. But
in the same work (pi. 32, 1825), when he described and figured M. car-
bonaria (=M. irus carbonaria) from Sumatra, he corrected' the error
on the evidence of specimens resembling Buffon's, which he received
from the same island.
PLATE V.
MAMMA
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A. Head of Pig-tailed Macaque (Macaca nemestrina), typical form.
B. Head of Burmese Crab-eating Macaque (Macaca irus aurea).
C. Head of typical Macaca irus, showing hair-growth on the cheek.
MACACA. 79
The colour is very variable both individually and racially,
and also the size and the length of the tail ; but the British
Indian race is larger, both in its skull- and body-measurements
than M . radiata (Geoffr.), and has the tail a little shorter than
the head and body.
10 a. Macaca irus aurea Geoffroy. The Burmese Crab-eating
Macaque.
Macacus aureus, Geoffroy, Zool. Voy. de Belanger, pp. 58 and 76,
1831 ; Gervais, Hist. Nat. Mamm. p. 87, fig., 1854.
Macacus cynomolgus, Blyth, Cat. Mamm. Burma, p. 8, 1875 ;
Blanford, Mamm. Brit. Ind. p. 21, 1888 ; and of other writers
on the Burmese fauna (not cynamolgus Linn.).
Piiheous fascicularis, Wroughton, Journ. Bomb. Nat. Hist. Soe.
xxiii, p. 700, 1915 (not fascicularis Raffles, from Sumatra).
Pithecus vitiis, Elliot, Proc. U.S. Nat. Mus. xxxviii, p. 346, 1910.
Locality of the type of aurea, Pegu* ; of -vitiis, Domel Island,
Mergui Archipelago.
Distribution. — Lowee Btjema, Tenasseeim, Meegtti Aechi-
pelago, and S.W. Siam.
Distinguished from typical irus and other forms occurring
m Malaya, Sumatra, Java, etc., by the arrangement of the
hairs on the cheek. In the southern races the long hairs
below and in front of the ears sweep forwards and, meeting
the backwardly-directed hairs from the face, form with them
the characteristic fringe or crest on each side of the face,
rising sometimes to form a crest on the side of the head above
the ear. In aureus the hairs of the temple and upper part
of the cheek sweep backwards from the face, partly concealing
the ear, then turn downwards and forwards towards the corner
of the mouth and finally upwards, the general arrangement
being circular and resulting in a definite whorl and a small
crest low down on each side of the muzzle.
The colour and length of the coat vary considerably in
* Geoffroy cited Bengal and Pegu as the localities of this Macaque,
which he named aureus on account of the reddish-ochreous hue of the
specimens he saw. I have selected Pegu as the type-locality because
the species is not indigenous in Bengal. Belanger's specimen, which
Geoffroy was told came from Bengal, was no doubt bought in the
Calcutta Bazaar, where captive specimens, according to Belanger,
were offered for sale. Pegu, on the other hand, is a certain locality
for this Macaque.
The characteristic arrangement of the hairs of the cheek, which
I have used as the main diagnostic feature of this race, was illustrated,
though not commented on, by Gervais is 1854 when he figured, side by
side, the heads of aureus and carbonarius, which no doubt he saw in the
Paris Museum. Tiekell's unpublished figures, quoted so frequently
by Blanford, also show the same feature in specimens from northern
Tenasserim, one sketch indicating, though not emphatically, the hair-
whorl behind the mouth. I have added vitiis Elliot to the synonymy of
aurea entirely because of its locality, the description being valueless.
80
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MACACA. 81
specimens from the same locality in accordance with, the
moult and independently of it. A skin from Kathema Island,
Tavoy (H. C. Smith), 22 April, with the moult just finished,
has the hair of the new coat about 13 and 20 mm. on the hind
back and shoulders respectively, and the general hue of the
upper side at a little distance is dark greyish- or olivaceous-
brown, due to the blend of the blackish and ochreous annula-
tions of the hairs ; the limbs externally are a little paler,
but the upper side of the tail is black, becoming gradually
paler towards the tip ; the lower surface and insides of the
limbs are ashy grey, and there are some wholly black hairs
behind the brow-ridge.
Another, with the same locality and date, retains the old
coat everywhere, and its hairs are about twice the length of
those of the skin in fresh coat. It is also much paler, faded
to brightish tawny above, passing into dull ochreous-buff
on the flanks and hind quarters, its general hue recalling
that of the Rhesus Macaque (M. mulatto, (Zimm.)), the hairs
having scarcely a trace of ambulations, except on the head ;
the arms are duller and paler than the body, the legs below
the knee are grey, the tail is only slightly brownish above,
and the underside is paler grey. This skin is in the " red "
phase, like the type from Pegu.
All intermediates in coat-length and colour between these
two occur in skins from the Tavoy district (H. C. Smith),
two additional skins being " red " ; and a good series collected
by G. C. Shortridge in Southern Tenasserim between Tenas-
serim Town and Victoria Point agrees generally with the
Tavoy skins, although none of them exhibits the " red "
phase. But the coat may be longer and richer in colour, an
exceptionally handsome skin being that of a ^ from King
Island, 200 ft., Mergui Archipelago (Primrose), October,
which has a long glossy coat, 35 and 60 mm. long on the
hind back and shoulders, and the colour darker, with deeper
black and richer ochreous annulations. Another skin from
the Ye Forest near Ataran, in Amherst District, November, is
very like the King Island specimen, except that the coat is
shorter. An " orange-coloured " specimen, like the type from
Pegu, was recorded from Mergui by Blyth in 1875.
The size varies considerably, as shown by the following
flesh-measurements (in English inches) of specimens collected
for the Survey by Shortridge and Primrose : —
Head and
Locality and sex. body. Tail. Foot.
Tenasserim Town ; ad. $ 25£ 21| 5£
Thaget, Tenasserim ; ad. cJ 22£ 20$ 5|
King Island, Mergui ; ad. <J 21^ 19$ 5+
Pakohan River, Tenasserim ; ad. ? . . . 19§ 17f 5J
Tenasserim Town ; ad. $ 19$ 18$ 4$
VOL. I. G
82 CERCOMTHECID^.
The weights of the $ specimens from Tenasserim Town and
Thaget were 18 J and 13 lb. respectively, and of the two $
specimens 10J and 9 lb.
Habits. — The only peculiarity in the mode of life of this
monkey is its habit of frequenting tidal creeks and mangrove
swamps to hunt for crabs, upon which it feeds to a great extent.
iShortridge, who came across it in Tenasserim, said that
"although extremely plentiful where it occurs, this species is
here [in Tenasserim] confined to the neighbourhood of mangrove
swamps along the sea-shore and the banks of rivers. Its
chief habitat is along the edge of tidal creeks, where at low tide
it feeds on molluscs, crustaceans, and other marine animals.
It was so local round Victoria Point that, although swarming
along the banks of the Pakchan River, it was not once seen
near Bankaehon, only a few miles away from the river."
The monkey does not, however, appear to be absolutely
restricted to the coast and river-beds, since Primrose shot one
200 ft. above sea-level on King Island, Mergui. Nor does its
diet consist solely of crabs. No doubt, like other Macaques,
it eats insects as well ; and H. C. Smith, who examined the
stomachs of a number of specimens secured at Tavoy, found
that although a large percentage contained the remains of
crabs, these were generally mixed with vegetable food, and
in some cases the contents were entirely vegetable.
Tickell in his MS. states that since the tidal creeks in
Arakan, at the mouths of the Irrawaddy and in Tenasserim,
are the only high roads of the districts, the monkeys have
become familiarized to the sight of man and sufficiently
fearless to come near enough to pick up rice and fruits thrown
to them. This was confirmed by Blanford, who observed
them in Pegu follow a boat for some distance. Very naturally
also they are not only fearless of water but are adept swimmers.
Tickell tens of a wounded male jumping out of a boat, swimming
away, diving repeatedly and once swimming under water for
a distance of 50 yards.
10 b. Macaca irus umbrosa Miller. The Nicobar Crab-eating
Macaque.
Maoaaus cynomolgus, Blyth ,Journ. As. Soo. Beng. xv, p. 367,
1846 ; Blanford, Maura. Brit. Ind. p. 22, 1888.
Macacus umbrosus, Miller, Proc. U.S. Nat. Mus. xxiv, p. 789, 1902.
Locality of type, Little Nicobar Island.
Distribution. — Nicobar Archipelago, Great Nicobar, Little
Nicobar, and Katchal.
Distinguished from M . irus aurea and resembling typical
representatives of the species from the Malay Peninsula and
Sumatra in the mode of growth of the cheek-hairs, which
on the fore part of the cheek sweep backwards from the
OOLOBID.E. 83
face and on the hind part forwards from in front of the ear
And neck, the upturned points of the two hair-streams meeting
to form a conspicuous vertical crest which descends from the
■edge of the crown to the throat ; the eye-brow vibrissa are
.also less well developed, at least on the average*. The
general colour of the upper side and of the limbs is nearly-
uniform hair-brown, with a faint tinge of drab more pronounced
on the sides ; the hairs are marked with a cream ring which
is sharp on the crown and nape, less so on the buttocks and
-thighs ; the tail is dark drab above, almost black in its proxi-
mal half, pale drab below. The underside of the body is
pale drab. According to Miller the general colour differs
from that of specimens of irus from the Malay Peninsula
by being much darker and less yellowish.
Flesh-measurements (in English inches) : —
Head and
body. Tail. Foot.
Ad. <J 20f 21 i 5|
Ad. <J (type) 20£ 23£ S|
Ad. $ 18£ 20 5J
Skull-measurements of type : — <J. Total length 134 ; zygo-
matic width 90 ; orbital width 67 ; mandibular length 97 mm.
Although the flesh-measurements suggest that this Macaque
is on the average smaller, sex for sex, than M . irus aurea, the
skull of the $ appears to be about the same size.
Family COLOBIDiE.
The essential characters distinguishing this family from the
•Cercopitheeidse are given on p. 32; but in addition to
the sacculated stomach and absence of cheek -pouches, the
British Indian members of this family differ from the
Macaques in their longer, narrower hands and feet, with the
pollex and hallux shorter.
This family of monkeys is represented in Africa by the so-
called Guerezas or Coldbus, whence its name is derived ; but
the Oriental Region, where it ranges from India, Tibet, and
South China to Borneo, may be regarded as its headquarters.
* I am indebted to Mr. Miller for telling me (in litt., 1937) about
the hair-growth on the cheeks and the eyebrows in his skins of umbrosa.
In these particulars, he says, they differ markedly from the specimens
from the Mergui Archipelago described by Elliot as vitiis, thus justifying
the conclusion I had already reached that vitiis is a synonym of aurea.
From a comparison of Elliot's Mergui specimens with Geofiroy's account
of aurea, Miller suggested to me that the former should perhaps rank as
a local insular race distinguished by its duller colour ; but a majority
•of mainland skins is dull-tinted. The rest of the characters of umbrosa
1 have entered above are derived from Miller's original description.
G2
84 COLOBIDJE.
In this region there are several well-defined genera, the
Proboscis Monkey (Nasalis) and three Snub-nosed Monkeys
{Rhinofifhecus, Presbytiscus, and Simias), which are un-
represented in the British Indian fauna. The genera occuring
within Indian limits are not so well defined from each other.
They have been popularly called Langurs, a name open to the
objection that, strictly speaking, it belongs to a particular
kind found only in parts of India. The title Leaf-Monkey,
recently adopted by Hill, is perhaps the most appropriate.
Although some of the different kinds are fond of frequenting
rocks, or even old buildings, the Leaf-Monkeys are essentially
arboreal, inhabiting for the most part high forest trees and
feeding upon foliage, flowers, fruits and seeds of various
kinds, sometimes perhaps occasionally taking animal food and
Fig. 20. — Sacculated stomaoh of Oeylonese Leaf-Monkey
(Kasi senex).
but seldom coming to the ground except for water or a garden
raid. On the ground they can gallop, and bound along at
a good pace on all fours ; but it is in the trees that their
remarkable agility is chiefly manifested. They run with
speed and surety along the big branches, spring lightly from
one to another or take prodigious leaps from tree to tree,
often landing, spread-eagle fashion, on a mass of foliage.
Their structure is essentially adapted to this form of activity.
The loins are powerfully developed for the " take off," the
tail is exceptionally long and used, as a balance, and the hands
are long and narrow to play the part of grasping hooks.
The palm is about twice as long as wide, the thumb is greatly
reduced in size and nearly functionless, but the four remaining
digits are long and strong, the two middle ones being subequal
COLOBIDiE.
85
and a good deal longer than the others. The hand differs greatly
from that of the Macaques, in which it is comparatively broad
and stumpy, with the palm much wider, the thumb longer,
and the remaining digits about as long or shorter. The feet
also differ tolerably similarly.
There have been many confusing changes in the nomen-
clature of the Oriental monkeys of this family. So long as they
were all considered to represent a single genus they were cited
as Presbytia or iSemnopithecus, the latter mostly being preferred,
e. g., by Anderson and Blanford, • until Miller pointed out
that Presbytia is the older name. Then Thomas recommended
Kg. 21. — A. Right hand of Indian Langur (Semnopithecus entellus).
B. The same of Lion-tailed Macaque (Macaco, silenus).
the still older name Pithecus, and in this he was followed by
Wroughton, Fry, Hinton, myself, Phillips, and Hill. But
Pithecus was later authoritatively ruled out of use because its
type-species, veter Linn., cannot be identified from the descrip-
tion, and its type-specimen has disappeared.
In captivity at least the Leaf-Monkeys are sedate and
dignified in appearance and behaviour as compared with the
Macaques. The latter may be regarded as the "plebeians,
the former as the "patricians" of the Indian monkey-
population.
86
COLOBID.ffi.
That the species fall into a number of natural groups was:
realized by practically all modern systematists ; but the
overlap in external characters and the individual variability
in skulls made very difficult any satisfactory definition of
the groups. I made such an attempt in 1928 and, adopting
the namePiiSAecws, divided the genus into three sections, based
upon the colour of the newly-born young, namely, the Entelhos-
group, containing the species found in India and Ceylon, in
Fig.:
-A. Right foot of Indian Langur (Semnopithecus entellus).
B. The same of Lion-tailed Macaque (Macaca silenus).
which the young is uniformly black ; the Pyrrhus-gvowp
found to the east of the Bay of Bengal, in which it is golden-
red ; and the Aygula-gioup, also found to the east of the Bay
of Bengal, in which it is typically of the " cruciger "-type,
white with a black cross-mark on the back, the group-names
being taken from the oldest or most familiar specific names
in each section.
COLOBIDiE. 87
But in 1934, in an attempted revision of the species of the
Pyrrhus- and Aygida-gFOUips found to the east of British
Indian territory, I took for these groups the generic names
Trachypithecus and Presbytia. The Indian and Ceylonese
species I left as in my earlier paper, with the remark that, if
Pithecus be upheld, Semnopithecus Desm. (entellus) and
Kasi Reichenbach (jdhnii) would be synonyms of it. Pithecus,
however, was condemned ; and Hill in 1934 showed that my
association of johnii and its Ceylonese allies wth the typical
Indian Langurs on the evidence of the colour of the young was
wrong. It is true they are black in johnii and in the races of
entellus, in which they have been recorded, the only ones known
to me, but not in the Ceylon species, to which johnii is very
closely akin. Hill therefore in 1936, accepting the burial
of Pitheous, divided my unnatural Entellus-gcovcp into two
genera, . Semnopithecus and Kasi, which are respectively
precisely equivalent to the two species, entellus and senex,
which I adopted in 1928.
The four genera mentioned above occur within the British
Indian area ; but their distinguishing characters are not very
convincing, and in some respects are average rather than
absolute, owing to deviations from the typical form which
may crop up.
The characters given in the analytical key and in the
definitions which follow are taken from the British Indian
species.
Key to the Genera based on British Indian Species.
a. Newly -born, young never exhibiting "eruciger"
pattern ; no conspicuous white band on inside
of thigh and continued as streak down leg ;
skull with nasal profile straight or concave,
and brow-ridges usually more or less promi-
nent above interorbital septum, at least in.
adult <J.
6. Hairs of croup not appreciably shorter than
of area of back in front,
c. Newly -born young black, where known ;
head, always with hair radiating from
frontal whorl and with long bristly
eyebrows ; coat coarser ; skull, at least
in adult <J, with very prominent brows. [Desm., p. 88.
(India and Ceylon.) Semnopithectjs
c'. Newly-born young typically golden-red,
never black ; hair of crown smooth or
shaggy (a frontal whorl only in one race),
and eyebrows less well developed ; coat
finer; skull with weaker brow-ridges. [Reiehenb., p. 120.
(E. of Brahmaputra.) Tkaohtstithecds
&'. Hairs of croup noticeably shorter than of area
of back in front ; newly -born young typically
grey with white cheeks, black only in one
species ; other characters as in Trachy- [p. 146.
pithecus. (S. India and Ceylon.) Kasi Reiehenb ,
88 COLOBIDiE.
■a'. Newly-born young of " oruciger "-type, white
with cross-shaped blackish pattern on back ;
a conspicuous broad white stripe on inside of
thigh and continued as streak down leg below
knee ; skull with convex nasal profile and brow-
ridges obsolete mesially above interorbital sep- [scholtz, p. 158.
turn. (Tenasserim.) Pkesbytis Eseh-
Genus SEMNOPITHECUS Desmarest.
Semnopithecus, Desmarest, Mamm. ii, p. 533, 1822 (latinized form
of Semnopitheque, F. Cuvier, Hist. Nat. Mamm. iii, pt. 30, 1821,
which has often been cited as the source and date of the scientific
name); ¥. Cuvier, Dents Mamm. p. 247, 1825; and of many later
authors, including Anderson and Blanford (in part) ; Poeock,
Proc. Zool. Soc. 1934, p. 895, and Hill, Ceyl. J. Sci. (B) xx, p. 117,
etc., 1936 (sensu stricto).
Type of the genus, entelhis Dufresne.
Distribution. — Practically the whole of India, except the
western deserts, from the Himalayas southwards, and Ceylon.
Newly-horn young, where known, black throughout.
Adult always with the hair behind the brow radiating from
a frontal whorl and the eyebrows well developed ; hair on the
crown sometimes rising into a crest or tuft ; on the sacral
area (croup) not shorter than on the mid-back ; individual
hairs of the back comparatively thick, with coarse surface
imbrication and somewhat scanty pigmentation in the medulla
and cortex (Hill). Male and female alike in colour of the
pubic region. Clitoris sometimes elongated and pendulous,
with its tip projecting below the labia of the vulva *. End of
the penis with median notch on the corona and tumid margins
to the orifice. The skull is powerfully developed, especially
in the adult <J, the brow-ridges being prominent and the area
behind them forming a wide depressed platform.
In the days gone by, before the admission of " subspecies,"
the Langurs of this genus, only known from the geographically
isolated and extreme modifications of the type, were of course
assigned to several distinct species, notably by Blyth and,
following him, by Blanford, who accepted four, namely,
schistaceus from the Himalayas, entellus from the northern
plains, hypoleucos from the Malabar coast, and priamus from
the Coromandel coast and Ceylon. But Blanford's acquaintance
with Himalayan Langurs was apparently almost limited
to a few Nepalese skins procured by Hodgson, which are not
typical schistaceus of that author, and his knowledge of the
Langurs of Peninsular India was apparently equally imperfect
owing to want of specimens.
* According to Hill this condition of the clitoris indicates recent
parturition. That was not the case in the specimen in which I recorded
it (Proc. Zool. Soc. 1926, p. 1554). She was imported alive from
India and exhibited in the London Zoological Gardens, and had not
bred for some time.
SEMNOPITHBCD'S. 89
The Mammal Survey of British India, however, obtained
a large number of skins and skulls from many localities in
India and Ceylon, providing far more material than was
available to Blanford and his predecessors. The examination
of this material in 1928 revealed the existence of a number
of previously undescribed kinds and confirmed the distinctness
of those described by Blyth, but convinced me that the best
•way of dealing with them was to regard them as subspecies
of a single species because they are definable by characters
which actually or inferentially intergrade, the differences
being mainly differences in size, in the length of the tail, in
the length and fulness of the coat, and modifications of the
same style of coloration, all of which fall within the limits
of subspecific variation, whether the distribution be con-
tinuous or not.
That some of the differences in colour are associated with
wetter or drier environment is illustrated by the various
forms that inhabit Peninsular India. Typical entellus, occurring
in the plains of Northern India, south and south-west of the
Ganges, is mainly characterized by the strong contrast between
its black hands and feet and the rest of the limbs, which is
pale. When this monkey is traced southwards on the eastern
■and western sides of Peninsular India it gradually changes.
In parts of the Deccan and towards the Coromandel coast,
and as far south at least as Kurnool and the Nallamalai Hills,
it is represented by a race, anchises, in which the black of the
hands and feet is to a considerable extent replaced by white.
Farther south in the hills near Salem occurs a race, priam,
in which the hands and feet are wholly whitish, matching
the arms and legs, and the pelage is generally paler. But in
this kind another feature sets in, namely, the uplifting of the
hair on the crown behind the frontal whorl to form a tuft
or longitudinal crest. This is not invariably present, at least
in made-up skins, and its systematic value is doubtful. Two
races with similar crests, but darker in general hue, especially
■on the limbs, occur farther to the west in S. India, namely,
■elissa in S.E. Coorg, which has black hands and feet, and a form
in Travancore closely related to the Ceylonese race thersites.
There is also a form in Cochin, priamellus, known from a single
specimen, which, although tuftless, seems to be akin to those
with that ornament. These three tufted Langurs from
Salem, S.E. Coorg, and Travancore show that in the drier
■eastern zone the colour is paler than in the wetter western
.zone.
A similar general darkening of hue is revealed when entellus
is traced from the plains of the north southward into the wetter
.zone of the Western Ghats. The most northern of these
darker forms, achates, found in Bellary, Dharwar, and Kanara,
intergrades with entellus, but is on the average darker, especially
90 COLOBIDiE.
on the arms and legs, and has a blackish tinge between the
eye and ear. A little farther south on the Kanara-Mysore
boundary is another form, iulus, which has still darker arms
and legs. Farther south, in S. Coorg, is the darkest form of
all, seneas, darker even than hypoleucos, alleged to have come
from Travancore, which is nearly intermediate between
seneas and iulus.
Proof of the intergradation of all these forms is not yet
forthcoming, there is much yet to be learnt about them, and
future collecting may increase or diminish the number of
local races.
The distribution of the Himalayan forms is discontinuous
from that of entellus, which is apparently not indigenous
north of the Ganges and Chambal River. Nevertheless the
differences between the former and Peninsular Indian forms
are practically only definable as average differences, justifying
Anderson's view that " schistaceus " is only a " variety ' r
of entellus ; and Blanford himself thought typical schistaceus
from the Nepal Tarai must be the same as entellus. But
neither of these authors was personally acquainted with the
Langur of Kangra and Chamba, ajax, which has black hands
like entellus, and it is possible that in the past the distribution
of these two forms was continuous west of the Jumna.
In this volume I have adhered to my original treatment
of these Langurs as subspecies of entellus ; but Hill (Proc.
Zool. Soc. 1937, pp. 209-11, and Ceyl. J. Sci. (B), xx, p. 213,
1937) has recently reverted to Blanford's opinion that
schistaceus, entellus, hypoleucos, and priam represent distinct
species, each probably represented by several subspecies,
a conclusion derived, partly at all events, from embryological
differences observed between priam and entellus. There
will be time to consider the systematic value of this evidence
and its possible application to the problem when it has been
tested in several specimens of each of the different kinds of
Indian Langurs. The matter, however, is of no great moment,
since authors seldomagree about " species " and "subspecies " ;
and some I have no doubt would, give full specific status
to all or most of the forms to which I have accorded the lower
rank. There is also the seeming paradox to be borne in
mind that two forms may intergrade and interbreed in one
district, where they are " subspecies," and keep apart in
another, where they are " species."
Owing to the intergradation between most of the races of
this Leaf-Monkey and to some individual variation within
the limits of each, the construction of a satisfactory analytical
key for their ready determination is not an easy matter.
In the following key an attempt has been made to enumerate
the principal features upon which they have been separated.
SEMNOPITHECTTS. 91
But of two of the races described nearly a century ago, namely,
hypoleucos and dvssumieri, I have been unable to examine
specimens and have been compelled to judge their characters
from what the authors said. I have been unable to match
any of the skins at my disposal with either of them, as
described, but both were based upon a single individual.
Key to the described forms of Semnopithecus.
a. Whiskers long and thick, mostly or com-
pletely hiding the ears ; head and whiskers
uniformly whitish or pale buff and contrasted
with the tint of the back.
b. Arms below the elbows black and strongly
contrasted with the paler hue of the body ;
coat long, shaggy and mane-like ajax (Poc), p. 96.
&'. Arms and hands not black and not sharply
contrasted with the dark hue of the body ;
coat shorter, not shaggy and mane-like,
c. General colour dark earthy brown ; coat
thick and woolly achiUes (Poc), p. 95-
c'. General colour paler, slaty or greyish- [p. 92.
buff ; coat shorter and less woolly .... schistaceus Hodgs.,
a'. Whiskers shorter, not or only partly covering
ears ; head and whiskers variable, but never
so white and strongly contrasted with the
body as in section a.
d. Hands and feet black; generally contrasted
with arms and legs,
e. Crown of head at most a little paler than
nape and shoulders ; hands and feet
black or brown and typically strongly
contrasted with arms and legs entellus(Dufr .), p. 98.
e'. Crown of head clearly defined from
shoulders and back by its paler tint.
/. No tuft on crown ; upper portion of
whiskers typically clouded with
blackish.
g. Paler ; crown of head uniformly
huffish in tint ; tail and outside and
inside of legs below knees not black.
h. Arms below elbows not black,
contrasted with black hands . . . achates (Poc.),p. 103-
h'. Arms below elbows black, blending
with hands. [P- 107.
i. Paler; cheeks pale like crown . . dussumier i GeoSr.,
i'. Darker ; cheeks infnseate above, vulus (Poc), p. 104.
g'. Darker ; crown suffused with dusky
hairs ; tail and leg below the knee
black.
j. Inner side of thigh clothed
with white hairs ; appar-
ently smaller and not so [p- 108.
dark above and below hypoleucos Blyth,
j' Inner side of thigh mostly
clothed with black hairs ;
apparently larger and darker
above and below senate (Poc), p. 106.
92 COLOBIIXffl.
/'. A tuft on the crown ; upper portion of
whiskers pale like the crown elissa (Poo.).p- 113.
d'. Hands and feet not black, either white
mottled with black or brown or whole
coloured, but matching the arms and legs.
k. Hands and feet white mottled with dark [p. 101.
patches anchises (Blyth),
h'. Hands and feet not mottled, the feet at
least typically whitish, and the hands
darker, but both matching respectively
the legs and arms.
I. Cheeks, crown of head, and nape
practically the same pale tint, and
much paler than the back; no tuft [p. 112.
on crown priamellus (Poc),
I'. Cheeks lighter than crown, which is
tufted and usually nearly the same
tint as the back.
m. General hue paler priam Blyth, p. 109.
m'. General hue on the average darker ; fp- 115.
smaller thersites (Blyth),
If in this key the presence and absence of the head-tuft,
to which Blanford attached importance, had been taken as
an alternative character to the black feet under the headings
d and d', elissa would have been transferred to d' and
distinguished from priam and thersites by the pronounced
contrast between its black hands and feet and pale arms
and legs, and anchises and 'priamellus would have come under
d and be distinguished from the rest under that heading,
the former by its mottled hands and feet, the latter by its
white feet.
11a. Semnopitheeus entellus sehistaceus Hodgson.
Semnopithecus sehistaceus and nipalensis, Hodgson, Journ. As.
Soc. Beng. is., pt. 2, p. 1212, 1840 (not sehistaceus Blanf.).
Pithecus sehistaceus, Hinton & Fry, Journ.. Bomb. Nat. Hist. Soc.
xxix, p. 404, 1923.
Pithecus entellus sehistaceus and hector, Pocock, Journ. Bomb.
Nat. Hist. Soc. xxxii, pp. 477 and 481, 1928.
Vernacular. — Langur (Mallaha) ; Derdca (Pahari) ; Oooni
(Hindi in Kumaun).
Locality of type of sehistaceus, the Nepal Terai ; of hector,
Sitabani, Ramnagar in Kumaon, 2,000 ft.
Distribution. — The Nepal Terai, Ottdh, Kumatjn and
Gabhwal in the drainage of the Ganges and its northern
tributaries.
" Dark slate-grey above with the hands and feet somewhat
darkened ; the head and underside pale yellow ; the coat
more or less wavy."
This epitome of Hodgson's description does not apply
exactly to any of the skins, here assigned to sehistaceus,
SEMNOPITHECTJS.
93
available for examination. The only example from the
Nepalese area is a subadult $ collected at Hazaria Pathergatti,
300 ft. It evidently differs from Hodgson's type in its paler
colour, being uniformly pale greyish buffy-brown, not dark
slaty-grey, above and on the outside of the arms, with the
legs and tail a little paler, the general effect being concolorous,
except for the creamy- white head and nape ; the chest is
yellowish. The winter coat, 15 February, in accordance with
the low altitude, is comparatively short, with the hairs sub-
equal and about 50 mm. (2 in.) on the shoulder.
The examples I described as hector from Kumaun may,
I now think, be assigned to this race, as Hinton and Fry-
supposed . They tolerably closely resemble the skin from
Kg. 23. — Skull of subadult £ of Semno^ithecus entellus schistaaeus
from Hazaria Pathergatti in the Nepal Terai. X f .
Hazaria in colour, the general hue of the upper side being
slaty-grey tinged with buff or pale brown, with the arms
slightly darker, more fuscous -grey than the back. But the
coat in November, January, and February is longer than in
the Hazaria skin, being from about 65 to 80 mm. (3 in.) in
length on the shoulder.
The flesh-measurements (in English inches) are as follows : —
Locality and sex.
Nepal Terai (Hodgson's type) ; ad. S ?
Nepal, Hazaria Pathergatti ; subad, $
Kumaun, Almora (hector) ; ad. <J
Kumaun, Eamnagar (hector) ; ad. <J . . .
Kumaun, Ramnagar (hector) ; ad. $ . . .
Head and
body.
Tail.
Foot
30
36
8i
23
36
7*
27^
39
9
26
37
Si
24
33$
7J
m
COLOBIDtE.
The weight of the $ from Ramnagar was 38 lb., of the $
39 1b.
The skulls of the Kumaun specimens assigned originally
to hector have the nose projecting, the plane of the nasals
being sloped at an angle of approximately 45°, a very familiar
but not invariable feature in the other Himalayan races.
But in the skull from Hazaria Pathergatti, which from its
locality I regard as typical schistaceus, the nose is not nearly
so prominent, having more of a concave dip at the base of
the nasals.
Of the specimens I have seen and assigned to this race the
only one that came from the Nepal Terai, the type-locality
Kg. 24. — Skull of adult $ of Semnopithectis entellus sahistaoeus from.'Naiai
Tal, one of the specimens originally assigned to hector. This skull
differs considerably in shape from the skull from Hazaria Pathergatti,
but the differences are probably individual, not racial, x J.
of schistaceus, is the subadult £ from Hazaria Pathergatti.
This is approximately full-sized, yet neither in its flesh- nor
skull-measurements does it exceed large $ specimens of
entellus and priam from Peninsular India. Blanford, indeed,
thought the Langurs from the foothills of the Himalayas
would prove to be entellus, not schistaceus. But not realizing
that Hodgson described schistaceus from the Nepal Tarai,
he applied that name to the larger, darker form, achilles,
of the highlands of Nepal. So far as size is concerned schistaceus
is intermediate between the highland Himalayan races and
those of Peninsular India. Evidence of the existence of this
SBMKOKTHECUS. 95
.race in Oudh and Garhwal is supplied by an adult <J skull
from Bahraich in the former province and of three adult $
skulls from Garhwal (B. B. Osmaston), all received without
skins. The condylobasal length of the first is 112 mm., and
its average in the others is 111£. These dimensions are
practically the same as in the skull of an adult $ from Almora
in Kumaun, entered in the table (p. 117), one of the specimens
I described as hector. These skulls are only 6 mm., about
a quarter of an inch, longer in that dimension than the largest
skull of typical entellus from Hazaribagh.
Habits. — In Kumaun, according to Crump, this Langur
ranges from 1,100 ft. at Ranmagar to about 7,650 ft. at Khati,
.but may extend up to about 9,000 ft. Apparently it seasonally
migrates from higher to lower levels and vice versa. It is not,
for example, found at Lohaghat, 5,600 ft., in February,
but arrives later, in warmer weather. It is common in all
the heavy forests, sometimes associating during the day
with the Rhesus Macaque (Macaca mukttta viUosa True),
although the two species separate in the evening before
settling down for the night. Baptista found it fairly plentiful
in Hazaria near the Soonson River, where it was not
very shy. The call is " hoop ! hoop ! " generally uttered
as a warning cry by one of the troop.
Regarding the breeding habits the evidence is conflicting.
Blanford quoted a MS. remark by Hodgson that in schistaceus
pairing takes place in February and young are born in April
and May, the period of gestation being only two months.
This is clearly an erroneous inference. Young born in April
and May must be the product of pairing some time in the
latter half of the previous year. McCann, on the contrary,
found that females in the Terai were pregnant, or with
newly-born young, in February. It is not improbable that
Hodgson's statement referred to achilhs, which occurs at
high altitudes in Nepal and Sikkim, since his specimens,
originally named schistaceus in the British Museum, belong
to that race. If there is a definite breeding season, as McCann
thinks, it is likely enough that the young would be bom
earlier in the warmer Terai than in the colder mountains of
Nepal.
11 b. Semnopitheeus entellus achilles (Pocoek).
Semnopithevus schistaceus, Blanford, Mamm. Brit. Ind. p. 30,
1888 (not of Hodgson).
Pithecus entellus achittes, Pocoek, Jonin. Bomb. Nat. Hist. Soo.
xxxii, p. 478, pi. ii, fig. 2, 1928.
Vernacular. — Sahu Kdboo (Lepcha) ; Propyaka (Bhotia).
Locality of the type, Sathar Hill, Gorkha, 12,000 ft., 50 miles
north-west of Katmandu.
96 COLOBID^!.
Distribution. — Sikkim and Nepal at high altitudes ; ? Kash-
mir.
Distinguished from schistaceus by the nearly uniform
chocolate or dusky sepia-brown colour of the body, tail
and outside of the limbs sometimes with a paler brown or
buffy sheen, a pale tail-tip and patches of black on the hands
and feet ; nape, whiskers, and crown, except for the black
hairs on the brow, cream or dirty white and sharply contrasted
with the shoulders ; lower side and inside of limbs whitish
or tinged with buff. Coat full and moderately long, the hairs
subequal in length, on the shoulder from 50 mm. in $ (December
19th) to 90 mm. in <J (January 18th) (type).
Flesh-measurements (in English inches) : —
Head and
Locality and sex. body. Tail. Foot.
Nepal, Gorkha (type) ; ad. <J 30 36& 8£
Sikkim, Chuntang ; ad. $ 26J 33 8
Sikkim, Chuntang ; ad. $ 24 30£ 8
An adult $ from Lachen, Sikkim, weighed 35 lb. The skull
of achilles is a little larger than that of schistaceus, exceeding
it in eondylobasal length by about the same amount as the
skull of schistaceus exceeds that of entellus.
The recorded altitudes for this race are : Chuntang in Sikkim,
5,350 ft. ; Lachen, Sikkim, 8,800 ft. ; and Gorkha, Nepal,
12,000 ft.
Habits. — In Sikkim Crump recorded this Langur as fairly
plentiful in the Lachen Valley from about 5,000 to 10,000 ft.
Apparently it does not descend below Chuntang. He found it
always shy and wary and confined entirely to the heavy forests,
where it goes about in troops composed of a small number of
individuals. It is a very silent animal, even when fired at,
and he never heard its call-note.
lie. Semnopithecus entellus ajax (Pocock).
Pithecus entellus, Lindsay, Journ. Bomb. Nat. Hist. Soc. xxxi,
p. 599, 1926.
PUhecus entellus ajax, Poooek, Journ. Bomb. Nat. Hist. Soe.
xxxii, p. 480, pi. ii, fig. 1, 1928.
Locality of the type, Deolah in Chamba, 6,000 ft.
Distribution. — Chamba, Kangra, and Kttlu, at high
altitudes, districts drained by tributaries of the Indus.
? Kashmir.
Distinguished from achilles by its much longer, shaggier
winter coat and colour. General colour above greyish-brown,
greyer and paler than in achilles, with the pale hue of the crown
less sharply denned as a rule by the greyish tinge on the nape
blending with that of the shoulders ; also in adults the arms
below the elbows and the hands are black or blackish-brown,
PLATE VI.
MAMMALIA.
r<
4 ( tt r at (>
*~ •** ~*±
1. The Kangra Langur (Semnopithecus entellus ajax).
2. The Nepal Langur (Semnopithecus entellus achilles).
3. The Tarai Langur (Semnopithecus entellus schistaceus).
SEMNOPITHBCXJS.
97
and darker than the legs, which are paler greyish-brown,
with some sooty-grey on the thigh and knee, the feet a little
darker than the shin, bordered with white, and with white
hairs on the toes ; the tail like the back, but with the terminal
three or four inches white ; underside and inside of limbs white.
The coat is loose and shaggy owing to the inequality in the
length of its hairs, whichform a kind of mane over the shoulders,
upper arms, and flanks, especially in the <J, where the longest
hairs may be from 150 mm. (6 in.) to about 240 mm. (about
9| in in the type) ; whereas in the $ they are from about
100 mm. (4 in.) to 145 mm. (nearly 6 in.).
Fig. 25. — Skull of adult o* of Semnopithecus entettus ajax from
Chamba. x f .
Flesh-measurements (in English inches) about the same as
in achilles : —
Head and
Locality and sex. body. Tail. Foot.
Kangra Fort ; ad. <J 31 34£ 8$
Chamba (type) ; ad. <J 30 38 9
Chamba ; ad. $ 30 32 8
Kangra ; ad. $ 27 28£ 8$
Kangra ; ad. $ ; 23 34 8J
The weight of the $ from Kangra was 46 lb., of a (J from
Chamba 43 lb., and of the larger $ from Kangra 28 lb.
The skull is about the same as in achilhs.
VOL. I. h
y© colobidjj:.
The recorded localities and altitudes of this race are as
follows :— Chamba, from 6,000 to 7,500 ft. ; Kangra, 2,450 to
9,500 ft. ; Kulu, 9,800 ft. The specimens from Chamba and
Kangra were collected in December, January, March, and
April, those from Kulu in June, and the last were still carrying
their winter coats.
Habits. — The Langurs seen by Hutton at Simla amongst the
nr-trees laden with snow were probably representatives of
ajax. At Kangra Wells made the interesting observation
that the troops of this Langur consisted of a single adult <J,
a number of females with young, and also young males.
MeCann later reported the same habit in typical entellus
during the breeding season (see below). Wells was greatly
impressed by the muscular development of the largest <J,
its biceps being quite as large as an ordinary man's.
11 d. Semnopitheeus entellus entellus (Dufresne).
Simla entellus, Dufresne, Bull. Soc. Phil. Paris, i, p. 49, 1797 ;
and of most subsequent authors, at least in part.
Pithecus entellus entellus, Pooook, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 477, pi. ii, fig. 3, 1928.
Vernacular. — Langur and Hanum&n (Hindi) ; Vandra
(Kathiawar).
Locality of type, Bengal.
Distribution. — Noethekn Peninsular, India, south of the
Ganges from Bengal to Kathiawar.
Distinguished from the races, above described, occurring
north of the Ganges and in the western Himalayas by its
smaller size, thinner, typically shorter coat, the whisker-
hairs concealing the ears to a much less extent, by the darker,
less contrasted tint of the head, and by the sharp contrast
between the black hands and feet and the rest of the arms
and legs.
Colour of the upper side usually tolerably uniform, darker or
lighter shining sepia-brown, rather paler on the shoulders
and head ; the crown sometimes noticeably paler than the
shoulders, but blending through the intermediate tint of the
nape ; limbs, apart from the hands and feet, and the tail
nearly the same as the back, but the tail-tip often pale, and
the leg below the knee often paler than the thigh.
Flesh-measurements (in English inches) of some specimens : —
Head and
Locality and sex. body. Tail. Foot.
Hazaribagh, Bengal ; ad. <J 25$ 42£ 8£
Junagadh, Kathiawar ; ad. $ 23 37f 7£
Midnapore, Bengal ; ad. §> 22£ 38 7
Palanpur, Gujerat ; ad. ? 2l| 35 7£
A c? and $ from Midnapore weighed respectively 35 and 25 lb.
SBMNOHTHECTTS.
99
The range in altitude is approximately from 200 ft. in
Midnapore and Palanpur to 1,000 ft. in Hazaribagh.
The skull of entellus is smaller on the average than that of
schistaceus ; it is on the whole less prognathous and has the
plane of the nasals more vertical, the nose not projecting to
the same extent, so that the facial profile is differently shaped.
But there is a skull of entellus from Midnapore with the nose
projecting in the way characteristic of the northern races,
and in one or two skulls of the latter the nose is approximately
as vertical as in the majority of entellus skulls.
Habits. — The habits of this race are better known than those
of any other. Blanford described them at great length, and
Fig. 26. — Skull of adult <J of Semnopithecus entellus entellus from
Hazaribagh. x f .
some interesting information, corroborating, extending, and in
a few particulars emending his account, have recently been
supplied, by McCann (Journ. Bomb. Nat. Hist. Soc. xxxvi,
p. 622, 1934) . Probably all the more southern races, enumerated
below, on which few observations have been made, resemble
entellus in all essentials connected with mode of life.
By many Hindus they are regarded as sacred, and by being
unmolested they have no fear of man, and frequently invade
villages and houses and make themselves a great nuisance
by pilfering food and pillaging fruit-gardens and fields of
crops. Away from the villages they mostly frequent high
trees and rocky hills, never far from water. They are ex-
ceedingly active, and can leap from one tree to another
h2
100 coLOBnxaE.
20 or 30 ft. away, with a drop of perhaps 40 or 50 ft. in. descent.
They bound from rock to rock with similar ease, and run on
all fours with a succession of long leaps, covering the ground
with considerable speed. They go about in troops composed
of females and young of both sexes under the leadership
and guardianship of a single large male. According to
McCann, who has studied their daily life, the troop sets off
in the early morning and spends the forenoon feeding and sitting
about, their food consisting of leaves, buds, flowers, shoots,
and a great variety of fruits. Towards mid- day they usually
return to their resting place and sit about on branches and
rocks, dozing or sleeping for a couple of hours or so. The
troop then sets off again to its feeding ground, and towards
evening returns homeward for the night, the male being
usually the last to retire to sleep when the others have settled
down after much quarrelling for places near the ends of the
branches — not on the thicker boughs. Every member of the
troop is at all times on the watch for danger, and if one utters
a warning cry and bolts, the rest follow in the same direction
without waiting to investigate the cause. Alarm or anger is
expressed by a loud, harsh, guttural cry, uttered, as sportsmen
well know, at the sight of a tiger or leopard, and repeated as
the beast is followed through the jungle. But, as Blanford
says, the cry may be provoked by the sight of a running deer.
Crump supplied the following note on this Langur in the
parts of Bengal where he collected and observed them. In
the small State of Chainpur they are plentiful and protected.
In parts of the Hazaribagh district they appear to be remarkably
scarce, only four specimens being seen. A few living on
Parasnath Hill are rigidly protected. In the Santara Range,
Singhbhum, they are well distributed,though not very abundant.
Owing to persecution by the Jumsare tribe, who kill them
for food, and by the Kols, who shoot them with arrows when
threatening the crops, they have taken to the thick forests,
seldom venturing into the open, and have become so shy and
wary that it was impossible to get near them through the
undergrowth.
To the west in Kathiawar Crump found them fairly well
distributed in the Gir Forest, although he did not meet with
many. In the Girna Mountains, where they are held sacred
and are fed by the natives, they were reported to occur in
great numbers. At Danta they were tolerably common,
and some were seen at Uria ; but in and near the town of
Palanpur, where the Hindoos hold them sacred, they were
plentiful, and might be seen on the house-tops and verandahs.
Crump also made some interesting observations on their habits
at Hewra in Nimar. He saw some come down to the ground
from the top of a very lofty tree by a quick succession of about
SEMNOPITHECUS. 101
iour vertical drops in an upright position, not as a rule from
one thick branch to another, but amongst the foliage. When
alarmed and running through long grass they move with great
bounds, frequently stopping and raising themselves to their
full height to take a look round. He saw one female, playing
with her young one, throw it into the air and catch it by its
two hands as it fell. She was no doubt instinctively giving
it practice in the use of its hands for grasping branches.
These Langurs from Hewra in Nimar were not typical entellus,
having paler hands and feet. One of them that was shot is
referred to below as an example provisionally assigned to
anchises.
As evidence of what he considered to be a difference in the
courage and prowess of these Langurs, Mr. Alison Minchin
recently published the following accounts of the behaviour
of two specimens (' Field,' Oct. 31, 1936). In Ganjam District,
■on the Coromandel coast, a large male, one of a big party
raiding the crops, was attacked by pariah dogs and, being
unable to escape, buried his face in his paws on the ground
and resigned himself to his fate, without apparently making
any attempt at defence. In the second instance another
male was the tyrant of a little village near Attikan, robbing
the huts as he pleased, and was a terror to the pariah dogs.
One evening he visited a rest-shed, where a man was staying
with a fine bull terrier, and was taking fruit off the table
when the dog intervened. The first intimation the occupier
had of the contest that ensued was the yelling of the dog,
and on rushing into the room he found the Langur had seized
his assailant by the fore legs and was pulling them apart.
On the man's appearance the monkey released the dog and
retreated to the other end of the room, but when the man
picked up the dog the Langur returned furiously to the
attack and made him release the dog, and the desperate
fight that followed ended ultimately in the Langur being
killed by the bull terrier, which, however, was badly bitten.
The Langurs concerned in these two episodes were identified
by the writer as " entellus." They were probably referable
to the next race, anchises, or to priam (p. 109).
lie. Semnopitheeus entellus anchises (Blyth).
Presbytia anchises, Blyth, Journ. As. Soe. Beng. xiii, p. 470, 1844 ;
and xvi, p. 733, 1847.
Pithecus enteRus anchises, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 488, 1928, and xxxv, p. 53, 1931.
Locality of the type, " the Deccan."
Distribution. — South of the area occupied by typical entellus ;
the Central Provinces, and Kurnool in the Eastern Ghats.
102
coLOBiD.a:.
Distinguished, according to Blyth, from typical entellus by-
its paler hands and feet which, instead of being all black and
sharply contrasted with the areas of the arms and legs above,
as in that race, are mottled, the hands being mixed white
and blackish and the feet whitish with dusky black above
Kg. 27. — A, hand of Senmopithecus entellus entellus from Hazaribagh ;
B and C, of two specimens identified as 8. entellus anchises from
Kurnool ; D, of <S. embeUus priam from Salem. The figures
illustrate the gradual -whitening of the hairs when the Langurs are
traced from Bengal to the Eastern Ghats.
the base of the toes and on the terminal phalanges, the leg
from the knee downwards being whitish, and the coat very
long.
SEMNOPITHECTTS. 103
The interest of this monkey, of which Blyth had only one
specimen, lies in its being intermediate in the colouring of
the hands and feet between typical enteLhis and the more
southern race described below as priam. Blanford dismissed
it as questionably the same as entellus.
In 1928 I provisionally assigned to anchises some specimens
collected at Hewra in Nimar, 1,000 ft. (Crump), and at Seoni-
Malwa (S. H. Prater), and in 1931 specimens were received
from Diguvametta and the ISTallamalai Range, 2,000 ft., in
the Kurnool district (A. S. Vernay's collection). These
specimens are not all alike in the tint of the head and dorsal
surface, one from Seoni-Malwa approaching the race described
below as achates in the tint of the crown ; but all agree with
Blyth's description in the reduction of the black on the hands
and feet, although this varies in extent, and it is to be noted
that the skins from Diguvametta, Kurnool, the most southern
locality known for the race, have the least black on the
extremities, thus coming nearer the white-handed, white-
footed priam, which occurs in the Palkonda and Shevaroy
Hills farther south in the Eastern Ghats.
The flesh-measurements (in English inches) of two examples
are as follows : —
Head and
Locality and sex. body. Tail. Foot.
Diguvametta, Kurnool ; old § 25j- 39 —
Diguvametta, Kurnool ; ad. $ 23 38 —
The weights of the two specimens were 24 and 23 lb.
respectively.
In five adult £ skulls the eondylobasal length is 85 mm. or
a trifle less, about the same as in entellus and priam.
11/. Semnopitheeus entellus achates (Pocock).
Semnopithecus anchises var., Blyth, Joura. As. Soo. Beng. siii,
pp. 471, 844.
Pithecus entellus achates, Poeook, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 488, 1928.
Vernacular. — MusJiya, Kari-Koti, Adari-Koti (Kanarese) ;
Yerpa-Moti-Koti Karrimikka, Mungya (Waddars) ; Kari-
Mangya (Haran Shikaris) ; Wanga, Wanar, Maha (Maratbi) ;
Langur (Dekhan) ; Kumdamuchu (Telegu).
Locality of the type, Haunsbhavi in Dharwar, 2,000 ft.
Distribution. — Dharwab, Bellary, and Kastara.
Distinguished from typical enteUus, with which it inter-
grades, principally by its slightly darker dorsal colour, by
its paler crown, which is buff or greyish-buff, and noticeably
contrasted with the shoulders, although not to the same extent
as in schistaceus and other northern races, by the blackish-
grey tint of the area between the eye and the ear, and by
104 COLOBID.SE.
the darker hue of the limbs below the elbow and knee, so that
the black hands and feet are not nearly so emphasized.
Flesh-measurements (in English inches), apparently slightly
exceeding those of typical entellus, are as follows : —
Head and
Locality and sex. body. Tail. Foot.
Havmsbhavi, S. Dharwar; ad. 3 27 43 8
Vijayanagar, Bellary ; ad. (J 26£ 40 8£
Samasgi, Dharwar-Kanara boundary ;
ad. 5 24 34 7
Vijayanagar ; ad. $ 26 35 7i
Haunsbhavi ; ad. $ 24| 33 7
Samasgi ; ad. ? 224 31 J 6J
The weights of the three g specimens, in order, were 30, 35,
and 22| lb., and of the three $ specimens 27, 22J, and 17 lb.
The skull is like that of entellus, but is on the average
decidedly smaller (see tables, pp. 117-19).
The recorded ranges in altitude of this race are : Bellary,
1,500 to 1,600 ft. ; Dharwar and the Kanara-Dharwar
boundary, 2,000 ft.
Habits. — According to Shortridge this race is extremely
plentiful in Dharwar, and is generally fearless and inquisitive,
easily distinguishing between Europeans and natives and much
more suspicious in the presence of the former. He confirmed
the observation made in the case of other kinds of Leaf-
Monkeys that, when frightened, they can crouch amongst
the tops of thick trees and by drawing the branches together
become completely hidden. It is equally abundant apparently
throughout Kanara, where it extends to the coast. Among
the hills round Vijayanagar at Bellary it was fairly plentiful
and unusually large. Its habitat is varied, and Shortridge
noticed that where it occurs in the open country it is much
less shy and suspicious than in the forests.
Newly-born young, probably not more than a few weeks old,
from Kanara, dated January 18 and 30 and March 22, suggest
the same breeding habits for achates as for typical entellus ;
but the data are obviously insufficient to warrant the con-
clusion that the young are born only at that time of year
approximately.
11 g. Semnopithecus entellus iulus (Pocock).
Piihecus entellus iulus, Pocock, Journ. Bomb. Nat. Hist. Soo.
xxxii, p. 490, 1928.
Locality of the type, Jog, Gersoppa Falls, on the Kanara-
Mysore boundary, 1,300 ft.
Distribution. — Unknown, except from the type-locality.
Distinguished from achates by its closer, softer fur, smaller
size, and generally darker colour, especially of the arms and
legs below the elbow and knee.
SEMNOPITHBCUS.
105
General colour of the back deep brown, sharply contrasted
with the head, which is buff and has, as in. achates, a good
deal of blackish hair between the eye and ear ; arms in $
black, a little paler in $ ; legs deep greyish-brown outside ;
tail blackish-brown, with its terminal third whitish and strongly
contrasted, as in many examples of achates. Underside and
inside of limbs pale.
Flesh-measurements (in English inches) of the only two
specimens known : —
Head and
Locality and sex. body. Tail. Foot.
Gersoppa Falls ; ad. <J 20£ 34 6J
Gersoppa Falls ; ad. $ 20 36 6j
The weights of the two specimens were : <£ 21 lb., $ 18J lb.
Judging from these two specimens, which, like those of
■achates, were obtained and measured by the same collector,
'^ <rf*U
Fig. 28. — Head of Semnopiihecus entellus iulus ftom the Gersoppa Falls,
showing the black hair on the cheeks and the contrast in tint
between the light-tinted head and the dark nape, which is like the
back, two characters also exhibited by S. entellzts achates.
G. C. Shortridge, the tail of iulus is as long on the average as
that of achates, whereas the head and body and the foot are
shorter. The shortness of the head and body is borne out
by the smaller skull, although that of the $ is about the same
length as the skull of the small $ from Samasgi. No doubt
the two forms completely intergrade.
In his note on the Langurs of Kanara (achates) Shortridge
has the following pertinent remark : — "Adults vary con-
siderably in size and weight and also in the amount of black
on the hands and arms, specimens from Jog and Gersoppa
(below Ghats) being unusually dark." The detection by the
106
coLOBma:.
collector in the field of the characters upon which this
subspecies was established supports the view that it is a
distinguishable form.
11 A. Semnopithecus entellus seneas (Pocock).
Presbytia hypoleucos, Dollman, Journ. Bomb. Nat. Hist. Soc.
xxii, p. 493, 1913.
Pithecus entellus seneas, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 492, 1928.
Locality of the type, Makut, S. Coorg, 250 ft.
Distribution. — Makttt, 250 ft., and Wottekolle, 2,000 ft.
Distinguished from achates and iulus by its generally
darker colour everywhere, noticeably on the inside of the
legs.
Back dark brown ; crown of head much darker than in
achates and iulus, and blending with the fore back ; whiskers
Fig. 29. — Head of Semnopithecus entellus seneas, from Wottekolle, S-
Coorg, showing the tolerably uniform dark hue of the cheeks, crown,
and nape, features in which this race differs from achates and
iulus, but resembles S. entellus entellus, which, however, is much
lighter on those areas and elsewhere.
reddish-white, blackish above ; limbs dark brown close to-
the body, but black elsewhere externally below to the hands
and feet ; tail black to the tip ; throat and chest reddish-
white, like whiskers, but inside of arms and legs black, as are-
the thighs to the pubic region, where the colour is sharply
contrasted with the whitish hue of the belly.
Flesh-measurements (in English inches) are about the.
same as in achates : —
Head and
Locality and sex. body. Tail. Foot.
Makut, S. Coorg ; ad. cj 28 37 7
Makut, 8. Coorg ; ad. ? 25 36 6J
SBMITOPITHBCFS. 107
The weights of the <J and <j> were respectively 25£ and 22 lb.
The measured skulls are very slightly larger than those of
iulus. The data are insufficient to justify a definite conclusion
on that point, except that they are in keeping with the
superiority in the flesh-measurements.
Shortridge recorded this Langur as fairly plentiful on the-
slopes and at the foot of the Ghats around Wottekolle and
Makut. It is replaced by elissa to the east of the Brahmagiri
Hills, which appear to separate the two at this point. He
added that he could get no information about the occurrence
of Langurs in North Coorg.
11 *". Semnopithecus entellus dussumieri Geoffroy.
Semnopithecus dussumieri, I. Geofiroy, C. R. Acad. Sei. Paris, xv,
p. 719, 1842 ; Arch Mus. ii, p. 538, pi. 1843.
Semnopithecus entellus dussumieri, Poeock, Journ. Bomb. Nat.
Hist. Soo. xxxii, p. 491, 1928.
Locality of the type,, Malabar coast. (Probably exported
from Mahe.)
Distribution. — Unknown apart from type-locality.
Distinguished apparently from hypoleucos, to which it is
nearly allied, by the paler colour of the back, which is greyish-
brown, not " deep dusky brown " ; by the inner surface of
the limbs being brown throughout, like the back, not black
below the elbows and knees and white above them ; and by
the terminal third of the tail being decidedly paler than the
rest instead of blackish throughout.
This Langur, of which only the type, an adult £ with its
young, has been recorded, is known to me only from the two-
descriptions, which disagree in one respect, and from the
plate, which does not agree precisely with either. In its
pale yellowish (fauve) nape and head, contrasted with the
brownish back, it evidently resembles achates, iulus, and
hypoleucos ; but the figure shows no trace of the dusky band
between the ear and eye present in" achates and iulus. It
further differs from those two races in the brown hue of the
inner surface of the limbs, which are like the back, darker
than the yellowish under surface of the body and contrasted
with it. The arms below the elbow outside and the hands
and feet are black ; but the leg outside is darkish brown
to the foot, the tail in its basal two-thirds is darker than
the back, and its terminal third is slightly paler than the back.
The race differs from eeneas in being much paler everywhere,
except on the hands and feet.
The measurements of the type, taken no doubt from the
skin, converted into English inches, are : head and body
24*, tail 34, which, if correct, are about the same as in $
achates and seneas and considerably larger than in the <J of
hypoleucos.
108 COLOBIDJE.
Blanford cited dussumieri, without question or comment,
as a synonym of hypoleucos. But unquestionably he had not
the material to justify the affiliation, and the descriptions of
the types of the two enforce their separation, at all events
until more is known of these Langurs of the Western Ghats.
11 j. Semnopitheeus entellus hypoleucos Blyth.
Semnopithecus hypoleucos, Blyth, Journ. As. Soc. Bengal, x,
p. 839, 1841 ; Anderson, Zool. Res. Yunnan, p. 20, 1878.
Pithecus entellus hypoleucos, Pocock, Journ. Bomb. Nat. Hist.
Soc. xxxii, p. 492, 1928.
Locality of the type, Travancore.
Distribution. — Tbavancoke, so far as at present ascertained.
A race known apparently only from the type-specimen in
Calcutta, which, according to Blyth's description, resembles
seneas in its wholly black tail and in the blackness of the
arms and legs below the elbows and knees, but differs in having
the inside of these limbs white above the elbows and knees
and in its much paler head, which, with the whiskers, was
described as " brownish- white, a little darker on the crown,"
and was evidently much paler than the back, said to be " deep
dusky brown." In the contrast between the paler head and
the darker back and in the whiteness of the upper half of the
limbs inside it is similar to achates and iulus, but differs from
them in its wholly black tail and in the dark hue of the lower
half of the limbs inside. It is thus in many respects inter-
mediate between seneas on the one hand and achates and
iulus on the other, although its stated locality is remote
from those of these three races.
The measurements of the type, an old $, taken no doubt
from the dried skin, are, according to Anderson : head and
body 21 in., tail 36 in., suggesting a small monkey about the
size of iulus. This conclusion is borne out by the length
of the skull, 106 mm., as also recorded by Anderson.
Kinloch identified as hypoleucos a Leaf-Monkey in the
Nelliampathy Hills which he described as comparatively
silent and occurring here and there in troops of about half a
dozen on the northern cliffs, but seldom, if ever, in the ever-
green forest. The Mulcers eat it, but do not consider its
flesh has the medicinal value of that of John's Leaf-Monkey.
But since the name hypoleucos has been used as a general
term for the dark forms of entellus found in the Western
Ghats, Kinloch's identification cannot be accepted without
reserve.
During his recent visit to Travancore Hill failed, in spite
of many inquiries, to get satisfactory evidence of the existence
of such a Languf there.
SEMNOPITHECUS.
109
312,
Ilk. Semnopithecus entellus priam Blyth.
Semnopithecus pallipes, Blyth, Ann. Mag. Nat. Hist. p.
1844 (April).
Semnopithecus priam, Blyth, Journ. As. Soe. Bene, xiii, p. 470,
1844 (Oct.).
Semnopithecus priamus, Blyth, Journ. As. Soe. Beng. xvi, pp. 732
and 1271, pi. 54, 1847 (in part) ; Anderson, Zool. Res. Yunnan,
p. 19, 1878, andBlanford,
Pithecus entellus pallipes, Poooek, Journ. Bomb. Nat. Hist. Soe.
xxxii, p. 395, 1928 (in part, excluding Ceylonese specimens),
and xxxv, p. 55, 1931 *.
Locality of the type, the Coromandel coast.
Distribution. — The Dhabmapixbi, Shevaroy, and Palkonda
Hills. Range elsewhere not certainly ascertained, but
recorded from the Nilgiei Hills.
Kg. 30. — Head of Semnopithecus entellus priam, the Tufted Langur, with
white hands and feet, from Salem, showing the contrast in tint
between the crown and the brow and cheeks.
Distinguished at once from the foregoing races by the
* As shown by the synonymy, I formerly called this Langur pallipes,
the earliest name given to it, and cited by Anderson, Forbes, and Elliot
as valid, but apparently not adopted by either of them because they
were unaware that it antedates priam. But although it is perfectly
clear that pallipes was given to the pale-footed Entellus of Southern
India, Blyth's remark about it does not, in the strict sense of the word,
constitute a technical description whereon a name can be based. He
merely recorded it as " the reputed Entellus Monkey of Southern India,
which is quite distinct from that of Bengal, and will bear the appellation
S. pallipes." Since the name is not sufficiently important to be
established by "fiat," which would be the sensible course, and is not
likely, I think, to be accepted in the future, I think it better, for the
sake of stability, to adopt here the name priam, which will probably
ultimately prevail.
.110
COLOBIDJE.
-presence of an upstanding tuft or longitudinal crest of hair
•on the crown behind the frontal whorl. This tuft may perhaps
.sometimes be lost in prepared skins and possibly absent at
times in the living animal when moulting ; but it is a normal
feature of the race.
Colour darker or lighter greyish-brown on the dorsal surface
.and the middle of the crown, with the nape paler, greyish-
tawny ; arms greyer than back, the hands whiter than the
fore arm ; thighs like the loins and back, but the leg becomes
gradually paler towards the foot, which is white ; the tail
.Fig. 31. — Skull of adult g, provisionally identified as Semnopithecua
entelkis priwn, from the Nilgiri Hills (Davison), x §.
usually a little darker than the back, sometimes blackish-
grey, with the tip whitish ; sides of the crown, whiskers, throat,
-and underside whitish.
Flesh- measurements (in English inches) : —
Head and
Locality and sex. body. Tail. Foot.
Shevaroy Hills ; ad. <J 25| 39£ 8f
Dharmapuri Range ; ad. <J 25£ 37$ 7£
Dharmapuri Range ; ad. ? 25 40 7£
Palkonda Hills ; ad. $ 23£ 37£ 7 J
The weight of the <J from the Shevaroy Hills was 37 lb.,
■of the $ from the Palkonda Hills 19| lb.
Special localities and recorded ranges in altitude for this
race are : Hogainakal and Murther in the Dharmapuri Range,
850 and 950 ft. ; Tirtamalai in the Shevaroy Hills, 1,000 ft. ;
SEMNOPITHECtrS.
Ill
and Dasarladoddi in the Palkonda Hills, 1,600 ft. Davison
told Blanford that this monkey ascends the western slopes of
the Nilgiri Hills to a height of 6,000 ft. ; but since only skulls
without skins are available, the identification is doubtful.
One of the skulls, that of an adult <J, taken by Thomas from
a, head-skin certified to have been tufted, agrees very closely
in size with the skulls from the Eastern Ghats and also with
the skulls of a couple of specimens from Travancore referred
to below. The skull of priam does not differ appreciably from
that of typical entellvg.
In the British Museum there are two adult c? unmeasured
skins, with tufted heads, from Travancore which, although
differing from each other a good deal in colour, are darker
JFig. 32. — Skull of adult S, referred to under Semnopithecm entellus
priam, from the Mahendragiri Range, xf. (For comparison with
fig. 31.)
than skins of priam from the Eastern Ghats. One came
from S. Travancore (Col. Dawson), the other from Aramboly
in the Mahendragiri Range (A. S. Pillay). I formerly tenta-
tively identified these as thersites on account of their hue.
But their skulls, measuring respectively 98 and 97 mm. in
•condylobasal length, are about the same size as those of
priam and larger than those of fhersites I have measured.
They may represent a race intermediate between the two,
but more specimens are needed. On geographical grounds
Hill suggested that they might belong to the next race,
priamelius, from Cochin ; but the only known specimen of
the latter, an adult $, has a paler head, without the tuft,
■and a considerably shorter skull, with a condylobasal length
of 83 mm. and no crest.
112
COLOBIDiE.
11 1. Semnopithecus entellus priamellus (Poeock).
Pithecus entellus priamellus, Poeock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 494, 1928.
Locality of the type, Cochin.
Distribution. — Cochin. Not recorded elsewhere.
Colour of back pale sepia-brown and sharply contrasted with
the creamy or slightly buffy- white nape ; crown a little
paler, and brow and whiskers a little yellower than nape,
the general hue of the head and its marked contrast with the
nape being very much as in achates, but there is no dusky
tract between the eye and ear ; tail greyish-brown, becoming
gradually greyer distally, with a small white tip ; arms similar
to the back outside, but becoming darker distally and blending
Kg. 33.-
-Skull of adult <J of Semnopithecus entellus priamellus
from Cochin.
with the hand, which is deep brown with black fingers ; legs
externally paler and greyer than the arms, the feet mostly
covered with dirty white hairs ; lower side and inner side
of the limbs all white or creamy- white.
There are no flesh-measurements of the only known specimen,
an adult $, but the skull is small, about the same size as the
<J skulls of iulus and hypoleucos.
Of the above-described western Indian forms of entellus,
which have no tuft on the head, priamellus comes nearest to
achates in coloration, and not to its geographically nearer
allies hypoleucos and seneas ; but it differs from achates in
its pallid feet, in the hands blending with the fore arm, and
in its uniformly tinted whiskers, which have no dusky band
between the eye and ear. Its affinities are doubtful. In
my analytical table of the races of Indian Langurs, in which,
SEMNOPITHECUS.
113
as stated, they were arranged in accordance with my views
of their kinship, I placed it under the same heading as
priam and alongside it, attaching subsidiary importance to
the absence of the crest. These opinions were confirmed
by Hill, who placed priamellus in priam, which he regarded
as a distinct species, adding that the absence of the crest in
the type of priamellus was not surprising because part of the
skin of the brow had slipped. The condition of the crown
does not suggest this to me, but, since its hair is thin and short,
the absence of the tuft may be due to moulting. On the
assumption that the tuft is potentially present, priamellus
is distinguished from the other forms of the Langur which
possess that ornament — from elissa by the blending of the
hue of the hands and feet with the area of the limbs above,
Fig. 34. — Front view of skull of adult <? of Semnopiihecus entellus
priamellus from Cochin.
instead of the sharp contrast ; from priam and thersites by
the pale hue of the nape and crown and their strong contrast
with the back, and by the noticeably smaller skull of the
adult $, and from priam at least, in addition, by its much
darker arms and hands.
11m. Semnopithecus entellus elissa (Pocock).
Presbytia entellus amchises, Ryley, Jburn. Bomb. Nat. Hist. Soe.
xxii, p. 494 1913 (not of Blyth).
Pithecus entellus elissa, Pocock, Journ. Bomb. Nat. Hist. Soe.
xxxii, p. 493
Locality of the type, Nagarhole, S.E. Coorg.
Distribution. — S.E. Coobg. Hitherto only known from the
type-locality.
vol. i. t
114
colobim:.
Resembling priam in the presence of an upstanding crest
on the crown, but distinguished by its black hands and feet
and the sharp contrast between them and the rest of the
limbs. The general hue is nearly as in achates, but there is
no fuscous tint on the cheek, and the end of the tail is less
extensively pale.
The upper side is brown ; but the head and nape are buSy-
or greyish-white and sharply contrasted with the back, the
crown being a trifle darker ; the tail above is like the back,
with the extreme tip whitish ; outside of arms and legs as
dark as the back, or darker, greyer, not so brown, and the hands-
Fig. 35. — Head of Semmopitheous entellus eUssa, the Tufted Langur,.
with black hands and feet and pale, uniformly coloured crown, brow,
and cheeks.
and the feet black ; lower side and inside of limbs greyish,,
whitish or creamy.
Flesh-measurements (in English inches) : —
Head and
Locality and sex. body.
Nagarhole ; yg. ad. <J 24-
Nagarhole ; ad. $ 25
Nagarhole ; ad. $> 24}
Curiously enough the flesh-measurements are about the
same as in priam, but the skulls are very noticeably smaller,
closely approaching those of iuhts.
According to Shortridge the single troop of this Langur
seen north of Srimangala was conspicuous from the crested
heads of the individuals. It was plentiful round Nagarhole,
and occurs in Coorg only to the east of the Brahmagiri Balls,
its place to the west of that range being taken by aeneas,^
which has no crest.
Tail.
Foot
36
36}
31}
7
6}
SEMHOPITHECTTS. 115
11 w. Semnopitheeus entellus thersites (Blyth).
Presbytis thersites Blyth, Journ As. Soc Bang, xvi, p. 1271, 1847.
Presbytia priamus and thersites, Kelaart, Prodr. Faun. ZeyL
pp. 3-5, 1852.
Semnopitheeus priamus, Blanford, Mamm. Brit. Ind. p. 31, 1888-
(in part).
Pithecus entellus pallipes, Pocock, Journ. Bomb. Nat. Hist. Soc
xxxii, p. 495, 1928 (in part).
Pithecus entellus thersites, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxv, p. 56, 1931.
Pithecus (Semnopitheeus) entellus thersites, Phillips, Man. Mamm.
Ceylon, p. 25, 1935.
Vernacular. — Grey Wanderoo ; Vandhura, Konde vandhura
or Elli vandhura (Sinhalese) ; Mundi (Tamil) ; Mundu
Kurangu (Jaffna Tamil) ; according to Phillips.
Locality of type, Trincomalee.
Distribution. — Ceylon and apparently Travastcoke.
Distinguished from priam, on the average at least, by its
smaller size and generally darker hue *.
Head typically with, a tuft on the crown, but this is occasion-
ally absent f. The general colour is individually variable;
the upper side is from greyish to darker brown, the tips of
the long hairs having a paler sheen ; the limbs are like the
back near the body, but become gradually paler distally,
the hands and feet being the same tint as the forearm and
shin, and not contrasted with them, the feet being whitish;
the crown is approximately the same hue as the back and
darker than the whitish cheeks and chin ; the underside
and inside of the limbs are mostly greyish-white.
The following approximate dimensions (in English inches)
and weights (in lb.) are taken from Phillips's volume : —
Head and
body. Tail. Foot. Weight.
Largest <J 28 35f 7$ 29$
Average of 6 £<? 24$ 34$ 7 27$
Largest $ 26$ 30$ 6| 19
Average of 7$$ 21$ 31 6$ 15$
Of the 4 <J(J and 4 $£ from Cheddikulam, Mankeni, and
Arucam Bay, whose measurements I recorded in 1931, none,
* Commenting on this race, Hill (Proc. Zool. Soc. 1937, Syst. p. 210)
said he was prepared to admit that Ceylonese specimens might be on
the average larger and darker above than specimens from the Coromandel
coast, justifying their subspecific distinction. In the material at my
disposal Coromandel specimens (priam) are larger on the average in
flesh-dimensions and weights, and also in the skxdl, than Ceylonese
specimens (thersites).
f Blyth originally regarded this Ceylon Leaf-Monkey as identical
with the Coromandel coast form described as priam. He considered the
crest an invariable character, and on the receipt of a specimen from Trin-
comalee -without the crest thought it represented a distinct species and
described it as thersites. In this view he was not followed by later writers.
l2
116
COLOBOUB.
either <J or £, was so large as Phillips's largest ; but their average
length of head and body was almost exactly the same.
Except that the skull is on. the average smaller than in
priam, it does not appear to differ from it. The average
eondylobasal length of" 7 adult <J skulls is 87 mm., about half
an inch shorter than in the 2 adult <J skulls of jyriam from the
Eastern Ghats.
Habits. — According to Phillips this Langur is commonly
found in the whole of the low- country dry zone, but does
not ascend the foothills to any great altitude, and is absent
from the wet zone. In its general habits it does not apparently
differ from its Indian allies. The troops or family parties,
Pig. 36. — Head of Semnopithecus entellus thersites, the Ceylonese Tufted
Langur, drawn by Blanford's artist from a skin from Ceylon
referred to by Blanford as S. priamus.
numbering from half a dozen to twenty or thirty, are composed
of mdividuals of all ages and both sexes. Near the villages
they are fearless of man because he leaves them unmolested ;
but in wilder districts they are more timid, especially in the
Vedda country, where the jungle men hunt and eat them.
They are as often seen on the ground as in trees, especially
about rocks and old ruins and near tanks and rivers. They
feed upon fruits, leaves, and grain of various kinds, and raid
cultivated ground. They are said, indeed, to eat without
ill effects the plant from which strychnine is prepared. Practi-
cally their only enemy in Ceylon is the leopard, which captures
them either by lying in wait or by so frightening them by
rushing at the tree in which they have taken refuge that they
fall to the ground from panic . This confirms Dunbar Brander 's
observations on the Entellus of Central India. Breeding
apparently takes place at all seasons of the year.
SEMNOPITHBCtrS.
117
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120 COLOBID-3E.
Genus TRACHYPITHECUS Reichenbach.
Trachypithecus, Reichenbach, Vollst. Nat. Affen, p. 88, 1862
(in part.) ; Trouessart, Bev. Mag. Zool. 1879, p. 57 (emend.).
The " Pyrrhus-grotxp " of Pithecus, Poeock, Journ. Bomb. Nat.
Hist. Soc. xxxii, p. 475, 1928.
Trachypithecus, Poeock, Proc. Zool. Soc. 1934 (1935), p. 928;
Hill, Ceyl. J. Sci. (B) xx, pp. 118 et seq. 1936.
Type of the genus (selected 1935), pyrrhus.
Distribution (as at present understood). — From Assam
through Bubma into Indo-China, the Malay Peninsula, Suma-
tra, Java, and Borneo.
Distinguished from Semnopithecus by the colour of the
newly-born young, which has the coat tolerably uniformly
golden-red throughout. In the adult the eyebrows are less
well developed, and only in one race is there a distinct
Pig. 37. — Head of Trachypithecus pileatus subsp. ?, drawn by Blanford's
artist from a skin of which no particulars were given.
frontal whorl with radiating hair. The hairs of the back
also, according to Hill, are thinner, more silky, and have
finer surface imbrications, and are pigmented only in the
cortex, not in the medulla as well*. The clitoris of the $ is
small, and contained within the labia of the vulva, not
elongated and pendulous as it is in at least some adult $
Semnopithecus. In the typical species, pyrrhus and some
others, the sexes differ, the $ having a patch of white hair
beneath the callosities ; but in other species the sexes are
alike in the colouring of the pubic region as in Semnopithecus.
* It -would be interesting to know what species of Trachypithecus
were examined for this character. It hardly seems probable that the
hair-pigmentation is the same in the black Javan pyrrhus as in the grey
Assamese pileatus.
TBACHYPITHECUS. 121
The skull has the brow-ridges less well developed, at least
on the average, than, in Semnopithems.
I have seen skins of the newly-born young of several different
kinds of Trachypithecus, ranging from Assam to Java, all of
the colour stated. The only exception to the rule apparently
was supplied by an example of T. pileatus, recorded by
McCann as white. But the young of this species known
to me are typically golden.
In the following key to the species here admitted the
characters of obscurus and pyrrhus apply to the races found
within British Indian territory, i. e., Tenasserim, not
necessarily to all the races found elsewhere.
Key to the British Indian Species of Trachypithecus *.
o. Sexes alike in the colour of the pubic region, the
tint of the hind abdomen continued back to the
callosities, to a certain extent down the thighs,
and sometimes on to the root of the tail ; lower
side paler, whitish-grey or ochreous.
6. The face black, hair on the crown forming a
mat typically differentiated from the hairs of
the temples and fore-nape by its greater
length and darker hue; size on the average [p. 121.
larger pileatus (BIyth),
6'. Face with a pale patch in front on the lips, and
a similar pale ring or half -ring round the eyes ;
no such mat on the head ; size on the average
smaller,
c. Hind legs, tail, and crown not sharply dis-
tinguished from the back by their much paler [p. 129.
tint phayrei (BIyth),
c'. Hind legs, tail, and crown sharply contrasted [p. 138.
with the back by their whitish hue obscurus (Reid),
a'. Sexes differing in colour of pubie region, the £ with
a pale patch of hair beneath the callosities ;
darker, browner above and below ; no pale [p. 142.
patch on month pyrrkus (Korsf.).
12. Trachypithecus pileatus (BIyth).
Semnopiihecus pileatus, BIyth, Journ. As. Soe. Beng. xii, p. 174,
1834 ; and xiii, p. 467, 1844. (For other bibliography, see under
the subspeeiflc headings.)
Locality of type unrecorded, but no doubt Assam.
Distribution. — Assam and the adjoining districts of Chitta-
gong and Uphbb Buhma.
Size large, face black, the crown from the brow, where
there is no whorl, to the occiput typically covered with a thick
* The species described by Blanford (op. cit. p. 38) as Semnopiihecus
chrysogaster, on the evidence of two specimens alleged to have come
from Tenasserim, is not represented in the fauna of British India.
That name is a synonym of Trachypithecus potenziani, which is found, ■
so far as is known, only in the Mentawei Islands, west of Sumatra.
122 COLOBIM3.
mat of soft, mostly erect hairs, which posteriorly overlap
the short hairs of the fore-nape and laterally those above
the ear and on the temple ; hairs of lower cheek long, whisker-
like, covering the lower part of the ear, and blending with
long hairs low down behind the ear. General colour of the
upper side, outside of the limbs, and the basal half of the tail
grey to blackish-grey with some long glistening hairs ; the
incrassate tail blackish in its distal half ; the hands and feet
also sometimes darker than the fore- arm and lower leg ;
the underside of the body, the inner surface of the limbs,
and the cheeks typically paler than the rest and sharply
contrasted, often suffused with red.
Key to the Races of T. pileatus.
a. Whiskers long, whitish or reddish, and
sharply contrasted with the darker tint of
the crown.
b. A sharp contrast in colour between the
upper and undersides of the body and
between the outer and inner surfaces of
the limbs ; throat and breast not paler
than belly.
c. Breast, throat, and whiskers pale, at most
tinged with bufi ; abdomen greyish- [p. 122.
white, at least in adult pileatus (Blyth),
c'. Breast, throat, and whiskers bright rusty
red.
d. Upper side paler, head and nape nearly
uniformly slaty-grey ; red suffusing [p. 125.
underside at least to groins durga (Wrought.),
d'. Upper side darker, head blackish,
a grey patch on nape ; red on under- [p. 126.
side fading away on abdomen tenebriais (Hint.),
6'. No sharp contrast in colour between upper
and undersides of body and between
outer and inner surfaces of limbs ; throat [p. 128.
and breast whiter than belly brahma (Wrought.),
a'. Whiskers shorter, grey, and not noticeably [(Wrought.), p. 128.
lighter in tint than the crown shortridgei
12 a. Trachypithecus pileatus pileatus (Blyth).
Semnipithecus ptteaVus, Blyth, Joum. As. Soe. Bene, xii, p. 174,
1843.
Semnopiiheous argentatus, Horsfield, Cat. Mamm. E.I. Co. p. 7,
1851.
Pithecus pileatus pileatus, Hinton, Journ. Bomb. Nat. Bust. Soc.
xxix, p. 79, 1923 ; Pocoek, Journ. Bomb. Nat. Hist. Soc. xxxii,
p. 661, 1928.
Locality of the type of pileatus, unknown ; of argentatus,
Sylhet.
Distribution.' — The Garo, Khasi, Jaintia, and Naga Hills
, in Assam.
Distinguished by the pale hue of the whiskers, throat,
TEACHYPITHECTTS. 123
underside of the body, and inner side of the limbs, which
are white or whitish, and at most faintly tinged with buff
or pale red and are sharply contrasted with the grey hue of
the upper side.
The colour varies a good deal individually. A half-grown $
from Mokokchung in the Naga Hills, 5,000 ft. (Wells), is
pale slate- or smoky-grey above and on the outside of the
limbs and on the tail, with the head and hands a little darker ;
the underside and the whiskers are white, and there is some
white on the hands and feet, but the feet are not darker than
the legs. An adult <J from the same locality differs in having
the whiskers and underside tinged with buff or very pale red,
the hands and feet darker, and no white on the fingers or toes .
An adult <J from Tura in the Garo Hills, 1,400 ft. (Wells),
differs from the last in having the arms and legs paler, the
outside of the leg below the knee and the top of the foot
being much greyer. An immature <J from Lait Kynsao in the
Khasi Hills, 2,000 ft., is very like the adult <J from Tura,
and an immature <J from Konshong in the Jaintia Hills,
3,000 ft., has the outer side of the leg below the knee greyer
than in the older specimen from the Khasi Hills, and the end
of the tail buff instead of blackish.
Flesh-measurements (in English inches)* and weights
(in lb.) :—
Head and
Locality and sex. body. Tail. Foot. Weight.
Tura, Garo Hills ; ad. $ . . . 28 41 7£ —
Changchang Pani, Naga
Hills ; ad. <J 26$ 35| 8 —
Changchang Pani, Naga
Hills ; ad. <J 25f 35| 7$ —
Konshong, Jaintia Hills ;
ad. $ 23 35 7£ 21$
■Changchang Pani, Naga
Hills ,- ad. $ 23 30| 6f —
As recorded by Hinton, on information supplied by
Mr. Mills, this race in the Naga Hills is only found in the high
cool jungle above 4,000 ft., and never mixes with the next
race, which occurs at lower levels on the slopes and in the
valleys.
Habits. — According to MoCann (Journ. Bomb. Nat. Hist.
Soc. xxxvi, p. 626, 1933), who observed this monkey at
Changchang Pani in the Naga Hills, it inhabits dense forests
intersected with streams, but was not seen to come to the
ground to drink, the plentiful dew supplying all the water it
requires, and it feeds on buds, leaves, and fruits. It is polyga-
mous, and at least during the breeding season in the winter
* The measurements of the specimens from Changchang Pani are
taken from McCann, -who identified them as pUeatus, without nominating
the subspecies.
124
COLOBIDJ3.
months goes about in small parties of eight or ten, consisting
of a master male, three or four females and young ; but there
are also parties of immature, non-breeding individuals. It
Kg. 38. — Skull of adult <J of Trachypitheaus pileatus pUeatus from Tura
in the Garo Hills. A. Frontal view. B. Lateral view.
makes a squealing noise at times, but its -warning cry is a harsh
bark, and, being exceedingly shy, it either, when alarmed,,
hides and remains quiet in the topmost foliage or takes to-
TRACHYPITHECTJS. 125
flight, making a tremendous noise by breaking and bending
the branches as it traverses the forest. This account refers
to typical pileatus or durga ; but no doubt the habits of
all the races are similar.
12 b. Trachypithecus pileatus durga (Wroughton).
Presbytia pileatus, Blyth, Journ. As. Soc. Beng. xvi, p. 735,
pi. xxvi, fig. 3, 1847.
Presbytia durga, Wroughton, Journ. Bomb. Nat. Hist. Soo.
xxiv, p. 655, 1916.
Pithecus durga, Hinton, Journ. Bomb. Nat. Hist. Soc. xxix, p. 79,
1923.
Pithecus pileatus saturatus, Hinton, Journ. Bomb. Nat. Hist. Soc.
xxix, p. 81, 1923.
Pithecus pileatus durga, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 663, 1928.
Locality of the type of durga, Cachar ? ; of saturatus, Bara
Hapjan in Lakhimpur.
Distribution. — From Lakhimpur in Uppeb Assam south-
wards to the Naga Hills, Cachar, Tipperah, Chittagong,
and Upper Chind-win to the west of the river.
Distinguished from the foregoing typical race by the
intensity of the redness of the whiskers, throat, underside of
the body, and inner surface of the limbs.
When Blyth received examples of this race from Tipperah
and Chittagong he regarded them as representing the males
of his pileatus, based on a pale-bellied female from an unknown
locality. But the difference in coloration is not sexual,
nor can durga be regarded as an erythristie mutant of pileatus,
because the two forms do not appear to live together, and there
is evidence of their occurrence at different altitudes. On the
label of an example of pileatus shot at Mokokchung, 5,000 ft.,
in the Naga Hills, Mills, the collector, wrote : " Grey-bellied
variety much less common than red-bellied, and found at
higher altitudes." He secured durga at Lakhuni in the
Naga Hills at 2,000 ft., but most of the specimens obtained
by the Survey were from much lower levels, 200 ft. at Lak-
himpur, 300 and 400 ft. at Golaghat, 400 ft. at Lanka in
North Cachar, and 800 ft. at Lamsakhang in the Cachar Hills.
Shortridge secured a specimen with " ginger yellow under-
parts and whiskers " at Nansun Chaung in Upper Chindwin,
and believed that this race occurs on the west side of the river
and the related form, shmtridgei, on the east side. A skull
picked up by Mackenzie near the Kabaw Valley is probably
referable to durga.
The locality of the type oi durga, collected over half a century
ago by 3>r. Reid, is doubtful. It was first labelled " Cachar "
and later " Upper Assam." But the point is of no great
moment, since this red-bellied Langur is found in northern
126 colobid^;.
Assam, at Golaghat and Lakhimpur, and also in Cachar.
Although Hinton regarded the types of durga and saturatus
as representing distinct species, I consider the differences
between them to be due to the age and the " make-up " of
the skins. The type of durga is darker above and duller
below than that of saturatus, but the latter is a fresh skin,
whereas the former has been many years in the Museum,
and is probably soiled and faded. The teeth of durga were
supposed to be larger than of saturatus. This is true of the
two types, as the table of skull-measurements shows, but
the cheek-teeth of an example of saturatus from Golaghat
are only 1 mm. shorter than in the type of durga.
Flesh-measurements (in English inches) and weights (in lb.)
are as follows : —
Head and
Locality and sex. body. Tail. Foot. Weight.
Barahapjan (saturatus type);
ad. 3 27f 39 7f- —
Golaghat ; ad. <? 23f 35f 7§ —
Lanka, N. Cachar ; ad. <? 23J 40* 9£ 27
Golaghat; ad. $ 20 33J l\ —
Lamsakhang, Cachar Hills . . 19f 33f 6f 25
ad. £
12 c. Trachypithecus pileatus tenebrieus (Hinton.)
Pithecus pileatus tenebricus, Hinton, Journ. Bomb. Nat. Hist. Soc.
sxix, p. 81, 1923 ; Poeock, Journ. Bomb. Nat. Hist. Soc. xxxii,
p. 664, 1928.
Locality of the type, Matunga River, N. Kamrup.
Distribution. — Assam, north of the Brahmaputra, Noeth
Kamkttp.
Distinguished from the two preceding races by the darker
tint of the upper side, the head, back, outer surface of the
arm, the hand, and the base of the tail being ashy black, but
the cap is denned behind by a half-collar of grey-tipped hairs
and the leg is somewhat greyer than the arm ; the throat,
breast, and shoulders, and the base of the whiskers are almost
as deep a red as in durga, but the whiskers are apically grey
as in the next race, brahma, and the redness of the underside
is much less pronounced on the belly and inguinal area,
which in the <J are whitish, in the $ pale red. A young
specimen is darkish grey above and on the outside of the
limbs, but whitish below, with a yellow tinge on the chest
and cheeks. It is darker than a young <J of the typical
race of about the same age from Konshong in the Jaintia
Hills.
Examples of this race were collected by Wells at Menaka
Nadi, 500 ft., Matunga River, 1,200 ft., and at Bogra Nadi,
2,000-3,000 ft.
TRAOHYPITHECtTS.
127
1st
WW-*
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occooco
CO W CO CO C^l
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CO CO (PI
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-IT*
00 t-
OOQCSO
If
%
£
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128 COLOBID-gl.
The flesh- measurements (in English inches) of two specimens
and the weight (in lb.) of one are as follows : —
Head and
Locality and sex. body. Tail. Foot. Weight.
JVIatunga Biver (type) ; yg.
ad. (J 22* 36f 7| 19£
Bogra Nadi ; ad. $ 22 j 34 7 —
The evidence of these measurements would hardly in them"
selves justify the opinion that this race is smaller on the
-average than the preceding two, but the skulls point to the
.same conclusion, the adult $ having the same eondylobasal
length as the smallest known adult £ of durga and being a little
less in that dimension, as in the length of the mandible,
•,than the young adult $ of pileatus from the Garo Hills.
12 d. Trachypithecus pileatus brahma (Wroughton).
Presbytia brahma, Wroughton, Journ. Bomb. Nat. Hist. Soc.
xxiv, p. 654, 1916, and xxv, p. 559, 1918.
Pithecus brahma, Hinton, Journ. Bomb. Nat. Hist. Soo. xxix,
p. 79, 1923.
Piihecus pUeatus brahma, Pooook, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 604, 1928.
Locality of the type, Seajuli in the Dafla Hills, North
Xakhimpur.
Distribution. — North Lakhimpue, so far as known.
Distinguished principally from the typical race, pileatus,
by the absence of sharp contrast in tint between the outer
.and inner surfaces of the limbs and upper and undersides
of the body, and by the slight suffusion of the flanks and abdo-
men with red, making them brighter and darker than the
•chest, throat, and whiskers which are white, with only the
tips of the whisker-hairs grey.
General colour above and on the outside of the limbs
uniformly slate-grey, but the limbs and tail darkening towards
their extremities ; the inside of the limbs pale grey, blending
with the tint of the outside, and there is no pale line invading
the hallux on the inner side of the foot ; whiskers white,
with ashy tips as in tenebricus ; throat and chest whitish,
•but the belly tinged with reddish-buff, becoming more intense
laterally where it passes into the flanks.
No measurements were recorded of the only known example
• of this race, a barely adult <J.
.12 e. Trachypithecus pileatus shortridgei (Wroughton).
Presbytia shortridgei, Wroughton, Journ. Bomb. Nat. Hist. Soc.
xxiv, p. 56, 1915.
Preebyiis shortridgei belHger, Journ. Bomb. Nat. Hist. Soc. xxiv,
p. 57, 1915.
TEACHYPITHEOXJS. 129
Pithecus shortridgei, Hinton, Journ. Bomb. Nat. Hist. Soo. xxix
p. 79, 1923.
Pithecus pileatus shortridgei, Pocock, Journ. Bomb. Nat. Hist.
Soo. xxxii, p. 665, 1928.
Locality of the type of shortridgei, Homalin in Upper
Chiadwin ; of beiliger, H'Kamti.
Distribution. — The Upper Chiadwin district of Uppeh Burma
to the east of the river.
Distinguished from all the preceding races by its nearly
uniform greyish hue, except on the hands and feet and the
greater part of the tail, which are blackish, the whiskers,
the underside of the body, the throat, and the inside of the
limbs being grey, with no trace of erythrism or whiteness and
only slightly paler than the upper side, which is slate or greyish-
brown. Also the whiskers are noticeably shorter and the
tail is distally short-haired ; but since all the known skins
were collected in July and August these may be seasonal
characters.
Flesh-measurements (in English inches) and weights (in
lb.) are as follows : —
Head and
Locality and sex. body. Tail. Foot. Weight.
Homalin (shortridgei type) ;
ad. $ 2S£ 40*- 7£ 30
Miasm ; ad. <J 27$ 39§ 7f —
H'Kamti (bettiger type); ad. <J 26£ 37£ 8£ 28
Homalin ; ad. $ 26$ 38i 7£ 21
This race inhabits the valley of the Upper Ghindwin River
apparently at comparatively low levels. Shortridge collected
it at H'Kamti, 500 ft., and at Homalin, 400 ft., and according
to him it is plentiful between those two localities and is found
only on the east side of the river. He also secured it at
Minsin * ; and H. C. Smith shot specimens in the Pidaung
Reserves, Myitkyina.
13. Trachypithecus phayrei (Blyth).
Presbytia phayrei, Blyth, Journ. As. Soc. Beng. xvi, p. 733, 1847.
(For other references see under the subspecies.)
Locality of the type, Arakan.
Distribution. — The whole of Btjbma, including Tipperah,
from at least as far north as Bhamo to northern Tenasserim,
and thence through Siam to Indo-China.
Distinguished from T. pileatus by the absence of the " cap "
* As Shortridge pointed out, Wroughton was misled into describing
the skins from H'Kamti as representing a distinct race, belliger. The
character on which he relied was due to discoloration of the skins by
smoke during preparation.
VOL. I. K
130 COLOBIDJi!.
on the summit of the head, which in that species is differen-
tiated from the hair of the temple by being longer, fuller,
and in all the races except one by its darker and contrasted
hue, the hairs in T. phayrei on the top and sides of the head
being essentially similar in colour, texture, and length ;
also in T. phayrei the dark hue of the face is relieved by
a pallid patch on the mouth and more or less round the eyes,
the end of the tail is not blacker than the back, there is no
tendency to erythrism in the coloration, and the size is, on
the average, considerably smaller.
Key to the British Indian races of T. phayrei here admitted.
•a. No whorl or parting in the hair of the forehead ;
eyes encircled by pale ring.
b. Darker on the average above and with sharper [p. 130.
contrast between dorsal and ventral surfaces, phayrei (Blyth),
£>'. Paler on the average above and with less
contrast between dorsal and ventral sur- [(Elliot), p. 134.
faces crepusculus
a'. A whorl or parting in the hairs just behind the [(Wrought), p. 136.
brow; a pale half-ring on innei side of eye. .. . shtmiem
13 a. Traehypithecus phayrei phayrei (Blyth).
Presbytia obseurus, Blyth, Journ. As. Soc. Beng. xiii, p. 466, 1844
(not obseurus Reid).
Presbytia phayrei, Blyth, Journ. As. Soe. Beng. xvi, p. 733,
1847, and of Anderson and Blanford under Semnopitheeus ;
Wroughton, Journ. Bomb. Nat. Hist. Soc. xxiii, p. 464, 1915,
and xxiv, p. 297, 1916 (in part)
Presbytia barbei, Blyth, Journ. As. Soc. Beng. xvi, p. 734, 1847
(not barbei Blyth, 1863 and 1875).
Presbytis melamera, Elliot, Ann. Mag. Nat. Hist. (8) iv, p. 267,
1909.
Pitheeus pyrrhus phayrei and barbei, Pocock, Journ. Bomb. Nat.
Hist. Soc. xxxii, pp. 668 and 669, 1928.
Vernacular. — Myauk-Ugenyo, Myauk-myet-kwin-byu (Bur-
mese) ; Myauk-Nyo (Toungoo).
Locality of the type of phayrei, Arakan ; of barbei, Tipperah
Hills*, of melamera Caud Ciaung, near Bhamo.
Distribution. — The whole of Btoma from at least as far
north as Bhamo to Pegu.
The explanation of the above-given synonymy, especially
the relegation of barbei, is as follows : — In his original descrip-
tion Blyth gave the name barbei to specimens said to have
been collected in the Ye district of Tenasserim, and stated
* Originally recorded by Blyth from the Ye district of Tenasserim ;
but this was corrected in 1863 to the interior of the Tipperah Hills.
Presumably the alteration applied to the original specimen of barbei
■and not to the one cited under that name at the later date (see p. 131.)
TRACHYPITHECrjS. 131
most positively that they resembled Malayan specimens of
T. obscurus, with which he was evidently familiar, in the
light markings on the face, i. e., the pale hue round the eyes
and on the lips. But in 1863 he not only changed the locality,
•on information received from Barbe, but contradicted his
first description by saying that the faces were black *. More-
over, in 1875 he stated that the skins were received without
skulls. Anderson, however (Cat. Mamm. Ind. Mus. i, pp. 48-9,
1881), when referring to specimens in the Calcutta Museum
as Blyth's types of barbei, described two skulls, citing their
catalogue numbers. Further, the Calcutta skins regarded
by Anderson as the types of barbei, and no doubt the same
as those called barbei by Blyth in 1863, have black faces.
This is shown by the reproduced photograph of the head of
one of them sent by Annandale to Wroughton (Journ. Bomb.
Nat. Hist. Soe. xxv, p. 47, 1917) and by the account of the
two given by Osman Hill (Proc. Zool. Soc. 1936, pp. 105-8),
who came to the conclusion that they represent a distinct
black-faced species of Leaf-Monkey from Tipperah. But
the evidence cited above shows they are not the original
specimens of barbei described in 1847, and there is no evidence
that they came from Tipperah if the alteration in the locality
applied, as may be supposed, to the original pale-lipped,
pale-eyed skins. There is one more point. Blyth was
largely actuated in separating barbei from phayrei in his
description of 1847 by the absence in the former of the crest
•on the crown, but he subjoined a footnote stating that the
taxidermist lad who prepared the skins was positive they
had crests when fresh. That, in my opinion, is not unlikely.
If so, it is another item of evidence favouring the view of
close likeness between the types of phayrei and barbei f.
Another name to be considered is mektmerus. In 1928
I considered this to be a synonym of barbei, which was regarded
as a race distinguished from phayrei by its generally darker
hue. But I had only one skin, the type of melamerus, to
* The view that the eyelids and lips changed from pale to black
in sixteen years is untenable. Skins 100 years old retain the pallor of
those areas.
t The source of the confusion about barbei will probably never be
traced. But it seems clear that Blyth must have had skins from the
Ye district in Tenasserim, whence came the type of the race described
below as atrior, which is black-faced, like his examples of barbei described
in 1863. But G. M. Allen (Amer. Mus. Nbvit. no. 429, p. 4, 1930)
has recorded as barbei a good series of fresh skins from S.W. Yunnan
which have blackish faces and are silvery above and below ; and these
seem to agree better with Hill's description of the supposed types of
barbei, allowing for soiling and deterioration of the latter, than do my
•specimens of atrior. It is quite likely that specimens resembling
Allen's will turn up in Upper Burma awl perhaps prove to be T.piUatus
shortridgei.
k2
132
COLOBID^.
judge from. Since then the examination of fresh material,
notably of a skin collected by Garthwaite near Bhamo,
whence the type of melamerus came, convinced me that the
supposed differences are individual, not racial.
The eyes in the hving animal are encircled by a white ring
which turns livid and reddish in dried skins. The hair on
the crown may or may not be raised into a crest, but there
is no parting or whorl behind the brow. The general colour
of the upper side may be deep, dull lustreless drabby, not
reddish, brown or buffy or greyish-brown, the pale area of
the hairs being more or less lustrous ; the forehead and cheeks
are blacker ; the arm below the elbow gradually darkens
to the black hands, and the feet are black ; the tail above is
usually paler than the back ; the chin, throat, chest and belly
are whitish or grey, and the pale colour extends in both
Fig. 39. — Head of Trachypithecus phayrei phayrei, without trace of
tuft, from 35 miles north-west of Toungoo.
sexes to the callosities and down the inner side of the thigh,
fading away inferiorly below the knee, but sharply defined
against the dark hue of the outer side of the thigh behind ;
frequently too it spreads on to the base of the tail below*.
One or two cases may be cited to illustrate individual
variation in colour in the same district. The type of mela-
merus from near Bhamo is mostly deep brown above and darkish
grey below. Garthwaite's specimen, also from near Bhamo,
500 ft., January 12, which, like the other, is crestless, differs in
being much paler and greyer, both above and below, and
closely resembles a skin from Kin in Lower Chindwin, whereas
another from Kin is decidedly darker and more like, although
* When the monkeys are in flight, with the tail uplifted, these whitish
areas may perhaps act as guide-marks.
TEACHYPITHECXTS.
133
not so dark as, the dark Bliamo skin. In a series of eight
skins from about thirty miles north-west of Toungoo (Mack-
enzie), one $, January 12, is very dark, lustreless drab-brown
above, very like the dark Bhamo skin, whereas in two ?$,
November and January, the long hairs from the nape back-
wards have extensively buff-grey, somewhat lustrous ends.
The rest of the Toungoo series are intermediate between these
extremes. A skin from the southern Zamayi Reserve, 6 miles
north of Pegu (Mackenzie), March 10, is even darker above
than the dark Bhamo skin, but has the belly dirty white.
This skin is like two from 30 miles south-east of Prome,
November 24 (Mackenzie), and it seems to resemble the type
of 'phayrei from Arakan, except that it has no crest.
Fig. 40. — Head of Trachypithecus phayrei phayrei, with, well-developed
tuft, from 35 miles north-west of Toungoo.
This crest, to which some importance has been attached,
is too unreliable in its incidence, at least in made-up skins,
to be used as a systematic character. In the series of Toungoo
skins it is present or absent. It is present as a thin upright
tuft in one from Kamri Island, Arakan, in the two from Kin
in Lower Chindwin, and in two from Mt. Popa (Shortridge) ;
but in two others from Mt. Popa and in those from Prome
and the Zamayi Reserve there is no definite crest, although
the hair on the crown, as in many of the Toungoo skins, is
long and irregularly straggling. In the dark Bhamo "skin
the hair on the crown is as long as in some of the southern
skins, but it is less untidy and has a generally backward
direction. In Garthwaite's skin from Bhamo it has the same
direction, but is shorter and smoother.
134 COLOBIDJE.
Habits. — The habits of this monkey are probably the same-
throughout Burma. According to Shortridge it is less noisy
and less seldom seen than the Indian Langurs (Semnopithecus),
being almost exclusively forest dwellers, and rarely frequenting
cultivated ground. He found it fairly plentiful on the higher
slopes of Mt. Popa, which he considered to be the northern
limit of the range of the race to the east of the Irrawaddy,
its place farther north and in the Shan States being taken
by shanicus. Near Toungoo Mackenzie reported that it is
usually found in dense high forest in parties numbering
about twenty. On being alarmed the females made off at
once with their young, but the males sometimes stayed
behind, barking at the intruders. More of them were con-
sequently shot then of the other sex. In flight the whole
party follows the same route, running along the same branches,
jumping from and to the same spots and now and again
stopping at the same place for a long look back. Their
leaps appear tremendous. Usually they seem to jump into'
a group o^ branches or the top of a lower tree, rather than
on to a particular branch, " spread-eagling " themselves
so as to distribute the weight, the tail apparently being
used as a balancer. All the specimens he secured were from
the west bank of the Irrawaddy ; but he saw a troop on the
east side of the river. The Burmans, he added, eat the monkey.
At Kin in Lower Chindwin the monkey was fairly plentiful
on the west bank of the river, but it was not seen on the east
bank, nor on the Upper Chindwin.
At Letsigan, 3,000 ft., in Lower Chindwin, in a party of
about twenty, Mackenzie shot on February 10 two $?,
one in milk, the other with a three-quarter-grown naked
foetus. This shows that this monkey may breed early in
the year. It is likely enough, however, that young are born
at all seasons.
13 6. Traehypitheeus phayrei crepusculus (Elliot).
Presbytia phayrei, Tiekell, Journ. As. Soc. Beng. xxviii, p. 428,.
1850.
Presbytia crepuscula and P. crepuscula wroughtoni, Elliot, Ann.
Mag. Nat. Hist. (8) iv, pp. 271, 272, 1909.
Presbytia ctrgenteus, Kloss, Journ. Nat. Hist. Soo. Siam, iii, p. 388,
1919.
Pithecus pyrrhua crepusculus, Pooook, Journ. Bomb. Nat. Hist.
Soc. xxxii, p. 672, 1928 ; Osgood, Field Mus. Nat. Hist., Zool.
xviii, p. 202, 1932.
Traehypitheeus phayrei crepusculus, Pocoek, Proc. Zool. Soo. 1934,
p. 949.
Locality of the type of crepusculus, Mt. Mulai-yit, Tenas-
serim ; of wroughtoni, Pachebon, Central Siam ; of argenteus-
Lat Bua Kao, west of Korat, S.W. Siam.
TRACHYPITHEOTTS. 135
Distribution. — Northern Tenasserim, the adjacent areas
of Siam, and thence apparently to Laos and Annam.
Closely resembling typical phayrei, with similar pale eyelids
and lips, but distinguished by being on the average paler
and greyer above and not so white below, the ventral surface
being grey and sometimes hardly paler than the dorsal.
Crest on the crown present or absent.
This race differs considerably in coat and colour according
to the season. This is illustrated by two skins from.
Mt. Mulai-yit, 5,000 ft. (Davison) which Blanford identified
as phayrei, rightly considering them to be the same as the
Langur from near Moulmein referred by Tiekell to phayrei.
One is tolerably uniformly brown above, with some rusty-
brown hairs on the arms, hands, and feet, so that the hands
and feet have lost their typical black hue. The other in
fresh coat is greyer, with black hands and feet, and scarcely
differs from examples of typical phayrei from Mt. Popa,
except that the belly is greyer and the base of the tail the same
tint as the loins. Very similar to the last is a skin from
Lampha, Tenasserim, 1,000 ft. (A. S. Vernay). An instructive
series showing considerable individual variation in tint, in
the same locality and the same month, was secured by A. S.
Vernay east of Urn Pang on the Mewong River, Siam, between
February 3 and 27, probably at the time of coat-change.
Some are paler grey, some darker and browner than the
pale skin from Mt. Mulai-yit, but they are obviously the same
monkey. An unusually brilliantly -tinted skin, with a golden
sheen, was collected by Vernay south-west of Kempenpet,
600 ft., on the Klong Klung River on February 27. A topo-
type of argenteus Kloss (September) is paler than the average
of Siamese and Tenasserim skins, but very like the palest
Um Pang skin.
Tickell's MS. account of phayrei, and his figure of a specimen
shot on the ground while drinking at a stream in the hills
between Kawkareik and Midiawaddee in Tenasserim, refer
no doubt to crepus&dus. In the living animal the face is
livid purple with white rings round the eyes and white lips ;
the general colour is smoke- or ash-grey aE over except the
brow and hands and feet, which are black, and the hind
abdomen and inner side of the thighs, which are white.
Habits. — It was not at all common near Amherst, keeping to
the depths of the forest and frequenting in small parties only
the hugest trees. It was seldom seen owing to its fear of man,
but might be heard crashing through the trees in full flight.
When by chance observed in flight, the party was seen to
keep in line, headed by the biggest, galloping along the
large boughs of the trees and occasionally flinging themselves,
with prodigious leaps, from one to another. The main body
136
COLOBIDJB.
continues its flight for some time, but an old male 'will
sometimes stop behind, uttering, with the mouth wide open,
a short deep bark, not unlike that of the Indian Entellus.
At other times they are usually silent. In flight they were
never seen to carry the tail upright, like the Entellus in India.
Soo.
13 c. Trachypithecus phayrei shanieus (Wroughton).
Presbytia barbei, Wroughton, Joum. Bomb. Nat. Hist.
xxiii, p. 465, 1915 (not of Blyth).
Presbytia shanieus, Wroughton, Joum. Bomb. Nat. Hist. Soo.
xxv, p. 47, 1917.
Piikecus melamerus, Wroughton, Journ. Bomb. Nat. Hist. Soo.
xxvii, p. 552, 1921 (not of Elliot).
Pithecus pyrrhus shanieus, Poeook, Joum. Bomb. Nat. Hist. Soo.
",p. 670, 1928.
Vernacular. — Lingmun, Ling Kang ? (Shan) ; Myauh-
Ugengo, Myauk-myet-kwin-byu (Burm.).
Locality of the type, Seen in the Hsipaw State.
Distribution. — The North Shan States and their neighbour-
hood to the east of the Irrawaddy, in the dry zone of Upper
Btjema.
Distinguished from the specimens assigned to typical
phayrei by the presence of a whorl, rarely represented by
*■*
"-Vrt^S
Fig. 41. — Head of Traehypishecus phayrei shanieus, from Hsipaw State in
the N. Shan States, showing the whorl on the brow.
a short longitudinal parting, in the hairs of the forehead
just behind the brow ; and, according to Shortridge, the
pale area round the eyes is limited to the inner portion of the
orbit.
Shortridge collected this monkey on the east bank of the
Irrawaddy, 560 ft., opposite Kyaukmyaung ; at Se-en, 1,411 ft.,
in the Hsipaw State ; at Gokteik, 2,133 ft. ; at Mansam Falls
and at Pyaungyaung in the North Shan State. H. C. Smith
TBACHYPITHECUS.
137
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subsequently got it at Nattaung, 700 ft., in Maymyo, Mandalay
District. In this specimen and in one from Gokteik the
parting on the brow is a short streak with the hairs diverging
right and left. In the others it is a whorl with radiating
hair, and this whorl is well defined in the immature specimen
from Pyaungyaung, which is just changing its baby coat.
Shortridge found this monkey plentiful in the Hsipaw
District and fairly abundant at Ngapyimin opposite Kyouk-
myoung, where it probably occurs as a wanderer from the
Shan Plateau, which at this point is not very far from the
river ; but he surmised that probably it is not found on the
Irrawaddy below Mandalay, the country being too open.
It lives in troops, is not apparently very noisy, but is shy of
man. It was on living specimens that he noticed the difference
between this race, then identified by Wroughton as barbel,
and. typical phayrei from Mt. Popa in the coloration of the
orbits, adding that the difference is not observable on dried
skins. That is true. In most of his skins of shanicus the
orbits appear to be coloured as in phayrei.
Flesh-measurements (in English inches) and weights (in lb.)
of the three races phayrei, crepusculus, and shanicus : —
Head and
Name, locality and sex. body. Tail. Foot. Weight.
phayrei.
Mt. Popa ; ad. (J 24 32 7 17}
Kin, Lower Ohindwin ; ad. <j . 22| 26£ 6$ 17}
Toungoo ; ad. $ 22} 31 7 —
Toungoo ; ad. <J 214 34 7 —
Mt. Popa ; ad. ? 21 J 31 6 15}
W. of Kindat ; ad. $ 184 25} 6 10}
W. of Kindat ; ad. $ 17} 28$ 6& 15
crepusculus.
Lampha, Tenasserim ; ad. $ . 22 31} 6 16}
sIumiAcus.
Gokteik, N. Shan St. ; ad. $ . 24 30 6} 19
Gokteik, N. Shan St. ; ad. $ . 23 31 ■ 6} 15
14. Traehypithecus obseurus (Reid).
Semnopithecus obseurus, Reid, Proo. Zool. Soe. 1837, p. 14 ; Martin,
Mag. of Nat. Hist, vi, p. 440, 1838 ; and of many subsequent
authors under the same or other generic names.
Trachypithecue obseurus, Pocock, Proc. Zool. Soc. 1935, p. 940.
Locality of the type, unknown *.
Distribution. — The Malay Peninsula from Johore northwards,
into Tenasseeim and S.W. Siam.
Resembling the typical race of T. phayrei in the pale hue
on the mouth and round the eye, bat t he eye not entirely
* Martin's good description of Reid's type, a specimen exhibited
in the Zoological Gardens, Loudon, shows tolerably conclusively that it
came from some place in the southern part of the Malay Peninsula. It
was probably shipped from Singapore.
TEACHYPITHEOTTS.
139>
encircled, the pale area being restricted to a semicircle above
and on its outer side. Further distinguished by the presence
of a cap of pale hair on the crown usually sharply contrasted
with the dark hue of the rest of the head and of the body,
which is brown or blackish, without silvery sheen, but has the
dorsal area usually paler than the flanks, often bronze-brown,
and this tint spreads over the shoulders in front ; the legs-
and tail are usually paler than the loins, sometimes a little,
sometimes very conspicuously, and the underside is dark
brownish or greyish-brown to nearly black.
There is no doubt about the close kinship between this
species and phayrei, and in my paper on the Langurs of British
India in 1928 I regarded them as subspecies of the same
species on the supposition that intermediate forms would
be found ; and it is true that skins of typical obscurus from
Fig. 42. — Head of Trachypithecus obscurus flavieauda from the mouth of
the Tavoy River, showing the pale crown and nape.
the Malay Peninsula are often very like examples of phayrei
from Upper Burma ; but in Tenasserim the characteristic
colouring of obscurus, i. e., the white cap and the pale con-
trasted tint of the hind legs and tail, become emphasized,
and there is no indication whatever that in this district
obscurus blends with the southern form of phayrei, i. e.,
crepusculus, which also occurs there.
The two races here admitted as members of the British
Indian fauna may be provisionally * diagnosed as follows : —
a. Tail much paler than the legs, which are only [p. 140.
a little paler than the loins sanctorum (Elliot),
a'. Tail approximately the same tint as the legs, [p. 140.
both being much paler than the loins flavicmtda (Elliot),
* Provisionally because Elliot did not describe the colour of the legs
in sanctorum. He merely compared the type-specimen with a more
southern, darkish-legged form, without mentioning any difference in
the hue of these limbs.
140 COLOBID^!.
14 a. Trachypithecus obseurus sanctorum (Elliot).
Presbytis sanctorum, Elliot, Proc. U.S. Nat. Mus. xxxviii, p. 351,
1910.
Pithecus pyrrhus sanctorum,, Pooook, Journ. Bomb. Nat. Hist.
Soo. xxxii, p. 673, 1928.
Trachypithecus obseurus sanctorum, Poeock, Proc. Zool. Soo. 1934,
p. 944.
Locality of the type, St. Matthew Island, Mergui Archipelago.
Distribution. — St. Matthew Island, so far as known.
Distinguished apparently from the other British Indian
races by the comparatively slight contrast between the tint
of the hind legs and of the loins, a feature in which it comes
nearer to the typical race of obseurus from Selangor, Johore,
etc., but differing from the latter in having the tail much
paler than the legs.
The characters of this race are merely inferred from Elliot's
untrustworthy description of it. He compared it to a race
named carbo by Thomas in 1909, which inhabits Terutau
Island, off the western coast of the Malay Peninsula, and
is a blackish form with the legs and tail approximately
matching and not very sharply contrasted with the loins.
From this he distinguished the type of sanctorum by its
creamy-white cap and buff-grey tail. If his description is
correct, sanctorum is distinguished from the next race by the
sharp contrast in colour between the tail and the hind legs.
The uncertainty about sanctorum is particularly unfortunate
because the name has page priority over the name of the next
race, which is well known.
14 6. Traehypithecus obseurus flavicauda (Elliot).
Presbytis flavicauda, Elliot, Proc. U.S. Nat. Mus. xxxviii, p. 352,
1910.
Presbytis obscura smithi, Kloss, Journ. Nat. Hist. Soo. Siam, ii,
p. 5, 1916.
Trachypithecus pyrrhus flavicauda, Poeock, Journ. Bomb. Nat.
Hist. Soc. xxxii, p. 672, 1928.
Trach/i/pithecus obseurus flavicauda, Poeock, Proc. Zool. Soe. 1934
(1935), p. 942.
Trachypithecus obseurus corax, Pocoek, Proo. ZeoL Soc. 1934
(1935), p. 944*.
* I have seen the type of smithi, not the type of flavicauda ; but the
examination of a large number of skins from the northern portion of
the Malay Peninsula convinces me that the alleged differences between
them are merely individual, not racial. The supposed race corax was
based on several specimens darker in hue both above and below than
typical flavicauda, and regarded as representing the most northern
race of T. obseurus. But since describing it I have received from
Mr. H. 0. Smith a skin from the mouth of the Tavoy River, a locality
to the north of the type-locality of corax, and this skin is of the flavi-
cauda type. Since a distinct subspecies is not likely to cut into the
area of flavicauda, I abandon, for the present at all events, the idea that
corax is a distinguishable form.
TKACHYPITHECTTS. 141
Vernacular. — Lutong, Lutong-laboo (Malay at Bankachon).
Locality of the type of flavicauda, Trang, Peninsular Siam ;
of smithi, Patiuyu in Peninsular Siam ; of corax, Tenasserim
Town.
Distribution. — From the northern part of the Malay
Peninsula northward through Tenasserim to Tavoy and to
the Pechburi district of S.W. Siam.
Distinguished from typical obscurus from the southern
portion of the Malay Peninsula and from others found in
the islands off the coast by the strong contrast between the
pale whitish or greyish tail and hind legs from the hips to
the ankles and the dark brownish or blackish hue of the rest of
the dorsal surface up to the head, which has the typical pale,
often silvery, conspicuous cap. From Elliot's description of
sanctorum it differs from that form by the legs being pale,
like the tail, not brownish, and contrasted with it.
This Leaf-Monkey is represented in the British Museum by
skins from the following British Indian localities : Bankachon
near Victoria Point, Tenasserim Town, Banlaw on the Great
Tenasserim River (Shortridge), King Island, Mergui Archi-
pelago (Pilgrim), and Tavoy (H. C. Smith).
A series of nine skins from Bankachon, December and
January, shows considerable individual variations in details.
The cap is usually silvery and conspicuous, sometimes soiled
yellowish, sometimes quite inconspicuous ; the back is mostly
black or blackish, with a varying amount of bronze-brown or
grey-brown on the shoulders ; the arms vary from decidedly
brown, only a little lighter than the shoulders, to pale,
glistening, buffy-whitish or grey almost to the wrist, and
nearly as pale as the legs ; the legs, mostly clear whitish,
may be ashy or pale smoke-grey ; the tail may match or be
contrasted with the legs, and is whitish or ash-grey or decidedly
yellowish throughout, or ashy-grey in its proximal, smoke-
grey in its distal half. Individual skins of this series closely
match skins from localities in the northern part of the Malay
Peninsula.
The proximity of St. Matthew Island, Mergui, to Victoria
Point suggests the likelihood of identity between the Leaf-
Monkeys of the two areas ; but the description of sanctorum
does not admit that determination, especially as Elliot referred
a specimen from James Island, a few miles to the north,
to flavicauda. This skin Kloss assigned to smithi, as also
a skin from Kissaraing Island. The skin in the British
Museum from King Island (September), as well as those from
Tenasserim Town (March) and Banlaw (April), have the hairs
of the belly darker, blackish when massed, than in the skins
from Bankachon, in which they are paler, palish brown when
massed ; but the skin from Tavoy is like the Bankachon set.
142 COLOBIDJB.
A flat > native imperfect skin from near Pechburi, S.W. Siam.
-(K. G. Gairdner), seems to agree best with. Tenasserim Town
and Banlaw skins.
Flesh-measurements (in English inches) and weights (in lb.)
•of J 1 , obscurus flavicauda:—
Head and
' Locality and sex. Body. Tail. Foot. Weight.
Tenasserim Town (corax type); 23£ 28 6£ 16J
ad. 6*.
Banlaw, Tenasserim ; ad. $ . . 22£ 29£ 6$ 15£
King Island, Mergui ; ad. $ . 25£ 30} 6} —
Bankachon ; ad. ? 25£ 31 6J —
Bankachon ; ad. ? 23 31£ 6| 19
Bankachon ; ad. $ 21£ 31£ 6* 18J
According to Shortridge this Leaf-Monkey is almost as
plentiful in the localities where hie collected it as the White-
handed Gibbon {Hylobates lar), but is much less noisy. The
alarm-note is almost like that of a Macaque, and quite unlike
the curious " hoot " of the Indian Langurs. It keeps to the
thickest jungles and is rather shy of man.
15. Traehypithecus pyrrhus (Horsfield).
Sernnopithecus pyrrhus, Horsfield, Zool. Res. Java (unpaged),
1820.
Presbytia pyrrha, Thomas & Wroughton, Proe. Zool. Soo. 1909,
p. 372.
Traehypithecus pyrrhus, Pocock, Proc. Zool. Soc. 1934, p. 929.
Locality of the type, Java.
Distribution. — From Borneo through the Sunda Islands
to the Malay Peninsula, Indo-China, Siam, and Tenasserim.
Distinguished from T. obscurus and T. pkayrei by the
absence of the pale patch on the mouth, although the eyelids
may be pallid, and by a sexual difference in the colour about
the callosities. In the $ the hairs round the callosities are
•of the same tint approximately as those of the underside,
but in the $ there is a patch of pale, usually white, hair beneath
the callosities.
The general colour of this widely ranging species is very
variable both individually and racially, ranging on the upper
side from black, deep brown or dark grey, with the tips of the
hairs of certain areas, occasionally restricted to the head and
hind legs, but generally of other parts as well, pale huffish
or grey, giving the pelage a spangled or frosted appearance ;
the hands, feet, and forehead are black, and typically at
least the end of the tail as well, but there is never an isolated
-pallid cap on the occiput and crown as in obscwrus.
TEAOHYPITHECTTS. 143
15 a. Traehypitheeus pyrrhus atrior (Pocock).
? Presbytia barbei, Blyth, 1863, and of Anderson, Blanford
& Hill under other generic names (for references see above,
p. 131) (not P. barbei Blyth, 1847).
Pithecus pyrrhus atrior, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 673, 1928.
Trachypithecus phayrei atrior, Pocock, Proc. Zool. Soc. 1934
(1935), p. 952.
Locality of the type, Ye Forest, 500 ft., inland of Moulmein
in the Ataran district of Tenasserim *.
Distribution. — The northern part of Tbnasseeim and
adjoining areas of Siam.
Distinguished from T. phayrei and T. obscurus, the other
British Indian Langurs of this genus found in Tenasserim,
by the characters mentioned in the diagnosis of pyrrhus
and also by the generally darker hue of the upper and under-
sides. From cristatus, the Malayan race of pyrrhus, and
germani, the Indo-Chinese and Siamese race, it differs by
having the tail dark greyish, slightly paler than the back,
instead of black distally, and darker than the back.
In the type, from the Ye Forest (Thurling), the coat is short,
rather coarse and lustreless, with a few silvery spangles on the
nape, the whiskers black, and the tail dark olive-grey. A
second specimen $> from the foot of Mt. Nwalabo in Tavoy
(Davison), April, has the long hairs over the ears, on the nape,
upper arm, and on the leg to a less extent, with silvery sheen
at the tip ; there is also some silvery spangling on the shoulders
and back and a good deal of ashy-grey on the chest. In this
there is only the merest trace of yellow hair detectable
about the callosities in the badly made up skin ; but in a $
fromMenam Hoi, S.W. Siam (K. G.Gairdner), there is a visible
patch of scanty hair below the callosities. In the type the
eyelids are cut away, but in the skins from Nawlaboo, Menam
Hoi, and two from Sai Yoke, S.W. Siam (K. G. Gairdner), the
eyelids have a livid, yellowish hue.
No flesh-measurements are available.
I suspect Blyth may have had in his hands a specimen or
specimens of this monkey from the Ye District when, in 1863,
he redescribed barbei and said it came from Tipperah. Also
1 think there is little doubt that the specimen from
Mt. Mulai-yit assigned by Anderson to barbei represented it.
Its affinities always have been a puzzle. I now think it is
more nearly related to T. pyrrhus cristatus, which extends
into the Malay Peninsula, than to the more northern species
T . phayrei ; that it is, in fact, the most northern representative,
* This locality is the same as that originally assigned to barbei by
Blyth, and the monkey agrees tolerably well with the one he described
as barbei in 1863, when he corrected the locality to the Tipperah Hills,
but it does not agree with the one described in 1847.
144
COLOBID^B.
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TEACHYPITHECtTS.
145
.apart perhaps from T. pyrrfms germani of Siam and Cochin
■China, of T. pyrrhus rather than one of the most southern
representatives of T. pliayrei. But although it is an isolated
form, not, so far as is yet known, intergrading either with
■Kg. 43. — A. Facial view of skull of adult $ of Trachypitheeus phayrei
shanicus from Gokteik, N. Shan States. B. The same of adult <J
of T. pyrrhus atrior from the Ye Forest, Tenasserim, differing in
its larger orbits and complete brow-ridge. Both x £.
VOL. L
146
COLOBID^!.
cristatus or germani, I provisionally regard it as a distinct
race of pyrrhus. One circumstance connected with it stands
out quite clearly. Living almost alongside each other in
Tenasserim and keeping absolutely apart, and entirely distinct
from each other in colour, are the three Leaf-Monkeys.
T. phayrei crepusculus, T. obscurms flavicauda, and T. pyrrhus
atrior. A fourth species coming into the same area is Presbytia
femoralis keatii (see p. 161), which, according to the present
conception of atrior, is, like it, a migrant from the south.
Genus KASI Reichenbach.
Kasi, Reichenbach, Vollst. Nat. Affen, p. 101, 1862 ; Hill, CeyL
Journ. Sei. (B), xx, pp. 118 et seq. 1936.
Type of the genus, johnii (Fischer).
Distribution. — S. India and Ceylon.
Distinguished from Trachypithecus by the newly-born young
being black or very light - silvery-grey with white cheeks.
Fig. 44. — Skull of adult $ of Kasi senex nestor from Panadura, showing
the sloped brow and more prominent angular portion of the
mandible, by which the skull of this species may be distinguished
usually from that of Semnopttheeus mtelkis thersites. x f .
and by the hairs of the sacral area (croup) being shorter than
those of the fore-back. As in the type of Trachypithecus,
namely, pyrrhus, the sexes differ a little in colour, the $ having
a distinct patch of white hair below the callosities and some-
times spreading to a varying extent on to the base of the
thigh.
KASI. 147
The principal differences between the two species here
admitted may be summarized as follows : —
os. Hairs of the cheeks typically tolerably uniformly
brown and nearly matching those of the crown,
never white at the base on each side of the face ;
throat and chin darker than the crown ; end of [p. 147.
the tail black johnii (Fisch.),
a'. Hairs of the cheeks generally whitish throughout
and contrasted with the crown, always white at
the base close to the face ; throat and chin
whitish or white and paler than the crown ; [p. 150.
end of the tail lighter than the rest senex (Erxl.),
16. Kasi johnii (Fischer).
Cercopithecus johnii, Fischer, Syn. Mamm. p. 25, 1829, and of most
later writers on the Indian fauna, including Anderson and
Blanford, but quoted as johni and cited under Semnopithecus,
Pithecus, etc.
Semnopithecus cucullatus, I. Geoffrey, Zool. Vog. Belanger, p. 38,
p.. 1, 1834, and of Blyth, 1859 (Presbytia).
Semnopithecus jubatus, Wagner, Sehreb. Saug. Suppl. i, p. 305,
1840, and of Jerdon, Mamm. Ind. p. 7, 1867 (Presbytia).
Pithecus senex johnii, Pocock, Journ. Bomb. Nat. Hist. Soe. xxxii,
p. 503, 1928.
Pithecus johni, McCann, Journ. Bomb. Nat. Hist. Soc. xxxvi,
p. 624, 1932.
Pithecus vetulus johni, Hill, Ceyl. J. Sci. (B), xix, p. 79, 1934.
Kasi johni, Hill, Ceyl. J. Sci. (B), sx, p. 124, 1936 ; Proc. Zool.
Soe. (Syst.), 1937, p. 206 (johnii)*.
Vernacular. — Turuni Kodan, Pershk (Toda) ; Korangu
(Baduga andKurumba), Karing Korangu (Mai.) ; Kari-Mushya
(Coorg) ; Mandi (Tamil).
Locality of the type of johnii, Tellicherry ; of cucullatus,
the Ghats (Bombay) ; of jubatus, South India.
Distribution. — The hill-tracts of Southern India, the
Western Ghats from Coorg southwards, the Mlgiri, Anamalai,
Brahmagiri, Tinnevelly, and Palni Hills, usually not below
3,000 ft.
General colour of the body, limbs, and tail, to the apex,
jet black above and below, but the dorsal surface typically
shows a faint pale speckle under reflected light and frequently
* In 1928, when I associated this monkey and its Ceylonese allies.
in the same group as the enteUus-like Langurs, although giving them
specific rank as P. senex and P. entellus respectively, the obvious course
to follow was to regard the comparatively trivial differences between
johnii and senex as of subspeeific value to avoid suggesting their
equivalence to the much greater differences between either of these
forms and entellus. But now that entellus and senex+johnii are
regarded as distinct genera, I agree with Hill that specific rank may
be given to johnii, which, although very obviously closely allied to
senex, does not actually intergrade with it.
L2
148 colobim;.
there is a conspicuous grizzled patch on the loins, whence it
spreads on to the extreme root of the tail and the upper part
■of the thighs ; this patch varies greatly in distinctness and
may be undetectable ; head with a black brow-band, but the
rest of the crown brown or buffy-brown, this tint spreading
■on to the nape ; the long cheek-hairs also brown, generally
a little darker than the crown, but the hairs black close to the
face and on the chin ; the throat with dark blackish- brown
hair, darker than the crown.
Flesh-measurements (in English inches) and weights (in lb.)
■of some specimens in the British Museum and of others
recorded by Kinloeh and McCann and entered in Hill's Mono-
graph : —
Head and
Locality and sex. body. Tail. Foot. Weight.
Nelliampathy Hills (Kinloeh) ;
6* 28| 28f 7f —
Nelliampathy Hills (Kinloeh) ;
6* 25* 30f 7* —
Tinnevelly (McCann) ; <J 26 36 — 20
Kodaikanal, Palni Hills (Mc-
Cann) ; £ 25 27 — 29
Kodaikanal, Palni Hills (Mc-
Cann) ; S 23 32 — 24
Hrahmagiri Hills, S. Coorg
(Shortridge) ; ad. $ 25$ 30f 7 —
Aiiainaad,S.Malabar(0'Brien) ;
just. ad. <J %2\ 37 7f —
Kukkal Shola, Palni Hills
(MeCann) ; ad. 2 24= 32 7 25
Nelliampathy Hills (Kinloeh) ;
2 22$ 28$ 6 —
McCann recorded several wild-caught specimens in which the
end of the tail was broken off, his larger <J from Kodaikanal
being an instance. No doubt Kinloch's large $ had the same
■defect.
The skulls differ profoundly individually both in the
degree of prognathism, which affects the slope of the facial
plane, and in the length of the cranial. This is shown by the
differences between the two skulls from KLotengady and
Anamaad. The latter, an older skull, has a shorter, more
vertical face and a very short back behind the auditory orifice.
The length and width of the cranial portion of the two is as
follows : — Kotengady, 83 X 60 ; Anamaad, 74 X 60 mm. Others
are intermediate. The mandibles also differ. In the Anamaad
skull the postdental portion has the hind border strongly
convex in its lower half and no inferior emargination, whereas
in the Brahmagiri Hills skull the postdental portion has the
hind border much less convex and the lower border emarginate
helow. (For the dimensions of these skulls see p. 157.)
KASI. 14&
Habits. — In Coorg, according to Shortridge, this monkey is.
apparently confined to the southern slopes of the Brahmagiri
Hills on the Wynaad border, where it occurs in the "sholas "
or patches of jungle surrounded by grass country on the
mountain sides. In this district, which he thinks is probably
its northern limit, Shortridge recorded the species from
2,700 to 5,135 ft. It is eaten by the jungle tribes, who use its
skin to make drums.
On the label of a specimen collected on the Colengody
Estate, Cochin, 3,500 ft., Ryley O'Brien wrote: "Common,
but very wideawake and difficult to get a shot at." At
Anamaad in S. Malabar he secured the species at 3,200 ft.
Kinloeh in 1923 described this species as formerly extremely
common in the evergreen forest on the Nelliampathy Hills,
where its noisy cry, " hoo-ha, hoo-ha, hoo," could be heard in
every direction all day long ; but its numbers are now greatly
reduced owing to persecution by the jungle Mulcers and
Kadars and coffee-estate coolies, who are inordinately fond
of its flesh. He recorded its altitude in these hills as 3,500 ft.
Confirming and extending these observations, McCann
(Journ. Bomb. Nat. Hist. Soc. xxxvi, p. 624, 1934) states
that the species occurs at the foot of the Tinnevelly Hills,
but in the Palnis is not found below 3,000 ft. Here it frequents
dense " sholas " and may often be noticed passing from
one to another over the intervening open green stretches.
Occasionally a solitary male, much scarred from fighting with
a probably successful rival, may be seen, but more usually
pairs or small companies. In the morning they are fond
of sunning themselves in the tallest eucalyptus trees near the
" sholas," but the afternoon is spent in sound sleep. When
cornered in a tree, the monkey frequently drops to the ground
and attempts to escape by running on all fours at a good
pace. The natives, who highly prize the flesh, blood, and
gall as cures for all diseases, use dogs to locate them, and are
attracted to the spot by the barking of the monkey at its
pursuers. At Kukkal Shola, in the Palni Hills, McCann
secured a 2 at 6,100 ft.
According to Hill the roar of the $ is like that of the Ceylonese
Bear-Monkey (K. senex monticola). Other sounds are a short,
sharp growl and a whining cry, the latter used more by the
2 and young.
McCann's discovery at Kodaikanal of a 2 with a full-time
fcetus on May 8 and of very young specimens on December 20
and 29 suggests that there is no definite breeding season.
150 COLOBIDJE.
17. Kasi senex (Erxleben).
Cercopitheeus senex, Erxleben, Regn. Anim. p. 24, 1777 *. (For
other references see under the subspecifie headings.)
Locality of the type and distribution, Ceylon.
General colour of the body and limbs varying from black
to brown, the crown of the head and nape darker or lighter
brown, usually paler than the shoulders and back, but the
long whiskers, directed back over the ears, typically white
and contrasted with the crown, sometimes pale brown at the
ends, but always white at the base close to the black face ;
the throat and chin white or whitish, like the whiskers, and
paler than the crown ; the distal end of the tail becoming
gradually paler to a varying extent.
Typical specimens of the four intergrading races of this
species may be distinguished as follows : —
•a. Crown and croup sharply contrasted by their
paler hue from the darker shoulders and back ;
distal portion of tail more extensively pale.
6. General colour of body and limbs black ;
crown deeper brown ; whiskers sometimes
brownish distally ; croup-patch often sil- [p. 151.
very ; size rather larger vetulus (Erxl.),
b'. General colour of body and limbs drabby or
greyish-brown ; crown paler brown ; whis-
kers white ; croup-patch not silvery ; size [p. 153.
smaller nestor (Bennett),
■a'. Crown and croup less contrasted by their paler
hue, sometimes scarcely at all, from the
generally brown hue of the shoulders and back ;
distal end of the tail less extensively pale. [p. 154.
o. Coat shorter and thinner ; tail longer . . . senex (Erxl.),
o'. Coat thick, shaggy, and long ; tail [(Kelaart), p. 156.
shorter monticola
* Eeasons for the adoption of this name are given on p. 154 (footnote).
Another name that has been used for these Ceylon monkeys, even
as lately as 191 8 by Wroughton.istfeie) - T-inn., 1766, based upon a monkey
described by Kay in 1793 as "white, with black beard " (? whiskers),
and alleged to have come from Ceylon. No doubt Ray was told it
was from Ceylon ; but from the evidence of the description, by which
one must abide, it did not, since no Ceylonese monkey has a black
beard or whiskers. That is the reason why Blanford (Proc. Zool. Soc.
1877, p. 622), quite rightly, rejected it, and. why it was not considered
either by Hinton or myself. But in 1934 Hill said veter was the earliest
name given to the " Purple-faced Monkey " of Ceylon. If so, it would
have to come in for one of the races, as Wroughton, who, like HilL
preferred the alleged locality to the description, perceived. But
Hill rejected it, not because of the "black whiskers," but because of
the white body of the type, being unable to determine to which of
the three races addicted to albinism the name applied. Now veter,
which is not a nomen nudum, as Hill declared, is the type of Pitheeus ;
and if assignable to the Purple-faced Monkey, as claimed, Hill should
have taken JPithecus, not Kasi, as the generic name. Another name for
this species, cited by Hill, and no doubt copied from Elliot or Forbes,
Macacus silenus var. alba, Fischer (Syn. Maura, p. 28, 1829). I can
KASI.
151
17 a. Kasi senex vetulus (Erxleben).
Cercopithecus vetulus, Erxleben, Regn. Anim. p. 25, 1777.
Cercopitheeus kephalopterus, Zimmermann, Geogr. Ges. ii, p. 185,
1780, and of many subsequent authors mostly as cephalo-
pterus.
Pithecus vetulus vetulus, Hinton, Ann Mag. Nat. Hist. (9)
p. 509, 1923 (with full synonymy) ; Phillips, Ceyl. J. Soi. (B) ;
xiii, p. 273, 1926.
Pithecus senex vetulus, Pocock, Journ. Bomb. Nat. Hist. Soo.
xxxii, p. 501, 1928.
Pithecus vetulus vetulus, Hill, Ceyl. J. Sci. (B), xix, p. 58, 1934
Phillips, Man. Mamm. Ceyl. p. 14, 1935.
Kasi vetulus vetulus, Hill, Ceyl. J. Soi. (B), xx, p. 127, 1936.
Vernacular. — Kala vandhura or Vandhura (Sinhalese) ;
Mundi (Tamil).
Locality of types of vetulus and h&phalopterus, Ceylon.
Distribution. — The wettest parts of the lowlands of the
west and south- west.
Face black, bordered at the sides by the white hairs of the
cheeks, which are produced into long whiskers towards the
•ears, where they are frequently pale brown ; chin and throat
Tig. 45.-
-Head of John's Leaf-Monkey {Kasi johnii) from the Brahma-
giri Hills, S. Coorg.
white or whitish. Crown and nape generally dark brown ;
shoulders, arms, flanks, and greater part of the back black
or brownish-black, finely frosted with grey or buff in reflected
light ; hind-back to the base of the tail covered with a large
find no such entry in this work. There is, however, a paragraph
-reading " Simia silenus y alba ; barba nigra," followed by a reference
-to Simia veter linn. But from the context it is perfectly clear that
■" alba " is the first word of the description, not a systematic name.
26
29*
6*
17
24
28
6*
—
21*
29
6*
12
17-|
27*
6
21
28*
6
14*
23
274
6
22
28"
6
11*
81*
28
6
21
30
6
11*
17*
27f
6
19*
27
6
—
152 OOLOBID^E.
patch of greyish or silvery short hairs, sharply defined against
the black in front, and spreading on to the outside and back
of the thigh, but dying away at the knee, the rest of the leg,
like the whole of the underside, except the throat and chin,
black or blackish ; tail like rump-patch at base, then darker
grey, but with about the terminal third conspicuously paler,,
dirty white*.
The following are the approximate flesh-measurements
(in English inches) and weights (in lb.) of some specimens
in the British Museum, and of others recorded by Phillips,
and Hill :—
Head and
Locality and sex. body. Tail. Foot. Weight.
Anasigalla (Hill and Phillips's
largest) ; ad. <J
Kalutara (Brit. Mus.) ; ad. <J.
Anasigalla (Brit. Mus.) ; ad. 6*.
Matara District, S.P. (Hill's
smallest) ; ad. g
Phillips's average of 7 ; ad. 3 .
Kalutara (Brit. Mus.) ; ad. $.
Kalutara (Brit. Mus.) ; ad. £.
Hiyere (Hill's largest) ; ad. $ .
Phillips's largest ; ad. °
Matara (Hill's smallest) ; ad. $
Phillips's average of 9 ; ad. $ .
The individual variation in size in both sexes is considerable.
I have assumed that the smallest # and $ from Matara
recorded by Hill are adult. It is curious that their three
dimensions are exactly the same ! The tail may be from
about 4 to 10 in. longer than the head and body.
According to Hill and Phillips this race is restricted to the
wettest part of the lowlands on the south-west and south
of the island, between the coast and the mountains, where
the average rainfall is from 75 to 200 inches per annum owing
to the south-west monsoon, thus resembling the Malabar
tract of India, where the equally black Leaf-Monkey (K.johnii)
occurs. In the Udugama Hills it ascends to 3,000 ft.
It is generally found in the forests, wandering about in
the tree-tops in troops consisting of 30 or 40 individuals
of both sexes and all ages ; but in the smaller jungles and
near villages the troops are only up to ten or a dozen in number.
In broken ground it may be seen clambering about the
boulders. In places where it is left unmolested it is com-
paratively fearless ; but where it is shot for food or its skins
it becomes timid and goes crashing away with prodigious-
leaps through the forests or hides in the foliage at the approach
of man. It feeds upon leaves, fruits, flowers, fern-fronds,
and seeds, possibly, also, to a certain extent on tree-frogs,
* At least in my skins. According to Phillips the tail becomes
reddish-bro-wn towards the tip.
KASI. 153-
since a young one in captivity ate them with avidity. Its
characteristic cry is a loud, oft-repeated " hooh ! hooh !
hooh ! " heard in the early mornings, but a shrill shriek or
bird-like twittering is also uttered.
No special breeding season is known, but the young are
apparenly born mostly in February and March.
17 b. Kasi senex nestor (Bennett).
Semnopithecus nestor, Bennett, Proo. Zool. Soc. 1833, p. 67.
Pithecus vetulus nestor, Hinton, Ann. Mag. Nat. Hist. (9), xi,
p. 510, 1923 ; Phillips, Ceyl. J. Sci. (B), xiii, p. 274, 1926.
Pithecus vetulus philHpsi, Hinton, Ann. Mag. Nat. Hist. (9) xi,
p. 510, 1923.
Pithecus senex nestor, Poeock, Journ. Bomb. Nat. Hist. Soc. xxxii,
p. 502, 1928.
Pithecus vetulus nestor. Hill, Ceyl. J. Soi. (B), xxiv, p. 62, 1934 ;.
Phillips, Man. Mraom. Ceyl. p. 18, 1935.
Kasi vetulus nestor, Hill, Ceyl. J. Sci. (B), xx, p. 127, 1936.
Vernacular. — Vandhura (Sinhalese) ; Mundi (Tamil).
Locality of type of nestor, Ceylon, probably Bayigam
(Phillips) ; of phillipsi, Panadura.
Distribution. — The low-country wet zone of the Western
Province.
Very like vetulus in the size and consp'cuousness of the
rump-patch and the extensive paleness of the tail distally,
but distinguished by the general hue of the back being
drabby-brown or greyish-brown and not so sharply contrasted
with the rump-patch as the black of vetulus ; the crown and
nape, too, are paler, not so deep a brown as in vetulus. It is
also rather smaller and lighter in weight than vetulus.
Approximate measurements (in English inches) of specimens-
in the British Museum and others recorded by Phillips and
Hill :—
Head and
Locality and sex. body. Tail. Foot.
Kottawa (Hill's largest) ; ad. (J 22 22+ 6
Phillips's largest ; ad. <J 20£ 30j 7£
Panadura(typeofj>W«ps?",Brit.Mus.);;? 19J 284 6i
Phillips's average of 6 ; <J 20 26 6£
Gonabendaduwa (Brit. Mus.) ; ad. $ . . 22| 30f 6|
Godigamuwa (Brit. Mus.) ; ad. 9 21 i 28 6
Godigamuwa (Brit. Mus.) ; ad. $ 19J 25£ 6J
Phillips's average of 11 ; 9 20J 26£ 6
According to these dimensions females are at least as large
as males. Hill's largest $ from Kottawa has an exceptionally
short tail. It died in captivity, and if the tail-measurement is
accurately recorded it seems probable that the organ was
imperfect, as is often the case in captive monkeys. Phillips
recorded the weight of a large captive <$ as 8| lb., the average
of 2 gg as 7| lb., and the weight of a small captive $ as just
under 7 lb.
154 C0L0BIDJ3.
The range of this monkey is the low- country wet zone
north of the Kaluganga, which separates it from vetulus,
where the average annual rainfall is typically from 75 to 100 in.,
but reaches 200 in. in the foothills. On the coast it occurs
at least as far as Colombo, but inland it ranges still farther
to the north.
So far as is known its habits do not differ from those of
vetulus.
17 c. Kasi senex senex (Erxleben) *.
Cercopithecus senex, Erxleben, Regn. Anim. p. 24, 1777 ; and of
several subsequent authors under various generic names,
including Semnopithecus senex, Blanford, Mamm. Brit. India,
p. 35, 1888 ; Pithecus senex senex, Poeock, Journ. Bomb. Nat.
Hist. Soe. xxxii p. 498, 1928.
Presbytia albinus, Kelaart, Prodr. Faun. Zeyl. p. 7, 1852.
PUhecus philhricki, Phillips, Ceyl. J. Sei. (B), xiv, p. 57, 1926.
Pithecus senex philhricki, Phillips, Ceyl. J. Sci. (B), xv, p. 122,
1929.
Pithecus vetulus philhricki, Hill, Ceyl. J. Sei. (B), xix, p. 74, 1934 ;
Phillips, Man. Mamm. Ceyl. p. 23, 1935.
Kasi vetulus philhricki, Hill, Ceyl. J. Sci. (B), xx, p. 129, 1936.
Vernacular. — Kalu vandhura (Sinhalese) ; Mundi (Tamil).
Locality of type (not preserved) of senex, " hills of southern
Ceylon " ; of albinus, Matale, C.P. ; of philhricki, Kantalai,
E.P., near Trincomalee.
Distribution. — The hills east of Matale and Madulkelle
up to 5,000 ft. ; also the low-country dry zone of the N.C.P.,
N.W.P., E.P., and C.P., " right across the Island in its northern
lowland portion south of lat. 8° 30', from Marich-chukkaddi
on the west to Mlaveli on the east coast " (Hill).
* The name senex, although admitted as valid by all authors, was
discarded by Hill in 1934 as a nomen nudum, which it certainly is not,
since it was given to monkeys from the hills of Ceylon described by
Knox as " milk white." Hill's real reason apparently was its ascription
to abnormally coloured individuals ; and for the same reason, with
which no systematist will agree, he rejected atbinus Kelaart, usually,
if not always, regarded as a synonym of senex. When dealing with
philhricki in his monograph of these monkeys in 1934, he said that
since Kelaart's white monkey (alhinus) came from Matale it was
undoubtedly an albinistic specimen of philhricki, and that those seen,
and described, by Robert Knox were probably also of this type.
I entirely agree, and can see no escape from the dropping of philhricki
as a synonym of albinus, since philhricki, and no other race, occurs
near Matale and is known from Phillips's records to be liable to albinism,
at least at Gammaduwa. Also I see no valid reason for dissenting from
the traditional identification of albinus with senex, which, as the oldest
name, has to be assigned, even arbitrarily if necessary, to one of the
races of this Ceylonese species. Hill has done a great service by showing
to which it belongs. I may add that when Phillips described philhricki,
which was unknown to me when I published my paper in 1928, he was
not aware of its occurrence in the hills by Matale, the type-locality of
albinus. ' '
KASI. 155
Distinguished from vetulus and resembling nestor in the
generally brown hue of the dorsal surface, but differing from the
latter, at least typically, by the comparative inconspicuousness
of the rump-patch, which is only a little lighter than the area
in front of it. It is also larger than nestor and than vettilus,
■on the average, in all its flesh-dimensions.
The only skins of this race that I have seen are three from
Kantalai, including the type, presented by the Colombo
Museum, -which have the rump-patch comparatively incon-
spicuous, and two unmeasured skins from Kala Oya, N.C.P.,
presented by Sir F. Colyer. These have the rump-patch to
all intents and purposes as well defined as in nestor, and I
should have assigned them to that race but for Hill's record
of " philbricki " from Kala Oya in 1936, based on seven
specimens. But his measurements of Kala Oya specimens
suggests that they are on the average rather smaller than
typical members of this race. No doubt Kala Oya is one of
the localities where the two races blend. In 1936 he sum-
marized the characters of the race by saying that it L " is
distinguished mainly by its large size and very long white-
tipped tail. The rest of its characters overlap those of other
races." In none of my skins is the tail white-tipped. The
extreme end is pale tawny, the pale area being less white
^nd less extensive than in vetulus and nestoi .
Flesh-measurements (in English inches) : —
Head and
Locality and sex. body. Tail. Foot.
Palonnaruwa, N.C.P. (Hill's largest) ;
ad. <J 26£ 33 7£
Palonnaruwa, N.C.P. ; ad. $ 24J 331 "'
Phillips's largest ; ad. <J 24J 33| 7
Kantalai (type of philbricki) ; ad. g . . 24 35 7$
Phillips's average of 4 ; ad. <J 23$ 33$ 7
Phillips's largest, from Sigirya C.P. ;
ad. $ 25$ 30| 7
Kala Oya (Hill) ; ad. ? 20 32§ 6*
Kantalai (topotype) ; ad. $ 19$ 27 6
Phillips's average of 3 ; ad. ? 2lJ 31 6J
The weight is variable. Phillips recorded a young <£,
from an unstated locality, as 22 lb., and reasonably surmised
that an adult <$ would scale 25 lb. But Hill's heaviest <J
from Kala Oya was about 16 lb., 2 $? from the same locality
were about 15| lb. and 9| lb. respectively, and 4 adult <J<J from
Palonnaruwa were about 13J lb. each.
This race is the only one of the four that inhabits the same
Area of Ceylon as the Langur (S. entettus ther sites). As
indicated above, it has a wide and varied range. According
to Phillips it spends much of its time sitting high up in the
branches of leafless trees or climbing leisurely from bough
to bough. In the lowland jungles those he encountered had
156 colobid^;.
little fear of man ; but in the hills and jungles at Gammaduwa
and elsewhere, where it has been hunted for its flesh and skin,
it is very wary and shy.
17 d. Kasi senex monticola (Kelaart).
Presbytia oephalopterus var. monticola, Kelaart, Journ. As. Soc
Ceyl. ii, p. 321, 1850, and Prodr. Faun. Zeylan. p. 2, 1852.
Presbytia ursinus, Blyth, Journ. As. Soc. Beng. xx, p. 155, 1851 ;
and of many later authors, including Blanford.
Pithecus senex monticola, Pocook, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 502, 1928.
Pithecus vetulus monticola, Hill, Ceyl. J. Sci. (B), xix p. 68, 1934 j
Phillips, Man. Mamm. Ceyl. p. 21, 1935.
Kasi vetulus monticola, Hill, Ceylon J. Sci. (B), xx, p. 128, 1936.
Vernacular. — Maha vandhura (Sinhalese) ; Mundi, Periya
mundi (Tamil).
Locality of the types of monticola and ursinus, Nuwara
Eliya.
Distribution. — The central hill ranges above 4,000 ft.
Generally resembling normally coloured specimens of
senex, exemplified by skins described as typical " philbricki,"
,1/ > - ■*". i.
Fig. 46. — Head of the Bear-Monkey (Kasi senex monticola) from
Hakgalla in TJva, 5,600 ft.
in its tolerably uniform brown hue, with the " cap " and
rump-patch at most only a little lighter and contrasted, and
with the white whiskers long and conspicuous, but distinguished
by its long, shaggy, thick, luxuriant coat, relatively shorter
tail, and more massive build.
Of this race, sometimes called the Bear-Monkey, from an
example of it being mistaken by Major Forbes for a Sloth-
Bear, the British Museum has a skin in poor condition received
from Kelaart, one from Hakgalla, 6,210 ft., one from Pattipola,
6,200 ft. (E. W. Mayor), and one from Adam's Peak (S. B.
KASI.
157
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"Sot
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body.
Tail.
Foot.
Weight,
23|
24f
7
20
23f
28
«f
—
22i
28
6f
20
21*
27
6*
—
23f
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20*
22*
25£
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—
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16*
22*
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158 COLOBID-ffi.
Bell) without record of altitude. This is blacker than the
last two. Kelaart got his type from 6,185 ft. in Nuwara Eliya,
and Hill recorded a specimen from the Elk Plains, 6,000 ft.
Flesh-measurements (in English inches) and weights
(in lb.) :—
Head and
Locality and sex.
Pattipola (Brit. Mus.); ad. <? .
Bogawantalawa (Hill) ; ad. (J .
Hakgalla (Hill and Phillips's
largest) ; ad. $
Phillips's average of 2 ; ad. $ .
Phillips's largest ; ad. <j>
Bogawantalawa (Hill) ; ad. § .
Hakgalla (Brit. Mus.) ; ad. ?.
Phillips's average of 3 ; ad. <j> .
It will be noticed that Phillips's largest $ has exactly the
same dimensions as Bill's $ from Bogawantalawa. They
were probably taken from the same specimen, differently
sexed by the authors.
According to Phillips this monkey is now found only in.
the jungles of the high mountain ridges and in the forests
around Hakgalla, the Bopats and the Horton, Moon and
Elk Plains.
Owing to the persecution to which it has been subjected,
largely for its luxuriantly coated skin, it is not only extremely
timid of man, but is in some danger of extermination. It has
already been killed out in certain districts where it was formerly
known, and is rare in localities where it still survives. Its
habits are the same as those of the other races. It lives
in troops among the tree-tops, and on the approach of man
either retreats precipitately through the forest or hides in
the foliage, concealed by its sombre hue, its white whiskers
blending with the clumps of lichens growing on the branches.
Soon after dawn, especially in the north-east monsoon, its
presence in troops is indicated by its loud cry, " hooh t
hooh ! hooh ! " echoing across the valleys from the high
jungles (Phillips).
Genus PRESBYTIS Eschscholtz.
Presbytia, Eschscholtz in Kotzeb. Reise, p. 533, 1821 ; and of many
subsequent authors in part.
"The Aygula-grenxp of Pithecus," Pocock, Joura. Bomb. Nat.
Hist. Soo. xanrii, p. 675, 1928.
Presbytia, Pocock, Proc. Zool. Soc. 1934, p. 896 (sm&u stricto) ;
Hill, Ceyl. Journ. Sei. Zool. & Geol. xx, pp. 116-23, 1936.
Type of the genus, mitrata Esch. (=Simia aygula Linn. ;
see Thomas, Proc. Zool. Soc. 1911, p. 128).
Distribution. — From Tenassebm and S.W. Siam, south-
eastwards to Borneo.
PRESBYTIS. 159
Distinguished from Trachypithecus and Kasi by the sum of
a number of characters of which one or more may fail when all
the species of the genus are considered, but which hold good,
so far as is known, in the case of the British Indian species
of the two genera.
The newly-born young is of the " crueiger " type, the
general whiteness of the colour being relieved by a dark
spinal stripe, whence on the shoulders a dark stripe extends
on each side to the arms. In the adult the inner side of the
thigh is conspicuously white and sharply marked off from the
darker hue of the outer surface.
In the adult $ the perinseiim is comparatively long, as
long as the vulva, which is situated lower down between the
callosities ; the clitoris does not lie between the labia of the
vulva, but is external to it, below and sunk in a small pit
in the centre of a rounded swelling which projects below the
callosities*.
The skulls of both genera are subject to a good deal of
individual variation, but on the average the cranium of
Presbytia is less sharply constricted behind the orbits, has the
occipital region more inflated, with a weaker crest, the brow-
ridges weaker, with the upper edge of the orbits straighter,
less arched, giving a " frowning " aspect to the brow, the
nasal bones prominently convex, the anterior nares broader
below, less elongated, the muzzle more abbreviated, and there
is a well-marked emargination where the zygomatic arches
join the maxillae ; the mandible finally is not so robust,
especially in its postdental portion. (See also Shortridge's
note on the appearance of the face in the only British Indian
representative of Presbytia, recorded on p. 163.)
18. Presbytis femoralis (Martin).
Sermwpithecus femoralis, Martin, Charlesw. Mag. Nat. Hist, (new
series) ii, p. 436, 1838 (first description of the species named
femoralis, but not described by Horsfield in Appendix to ' Life of
Sir S. Raffles,' p. 642, 1830.)
Semnopithecus or Presbytis femoralis of Anderson, Blanford, and
other authors, including Miller, Smiths. Misc. Coll. lxi, p. 28, 1913.
Piihecus femoralis, Pocock, Journ. Bomb. Nat. Hist. Soc. xxxii,
p. 675, 1928, and Proc. Zool. Soe. 1935, p. 900 (under Presbytis).
Locality of the type, Singapore (selected by Miller).
Distribution. — From Tenassekim and S.W. Siam through
the Malay Peninsula to Sumatra, the Rbio Archipelago, and
Natuna Island.
Hair of the forehead typically parted by a pair of whorls
from 20 to 24 mm. behind the brow, rising into an erect
* The characters of the $ external genitalia have only been observed
in one specimen of P. melalophus from Sumatra, and were described
and figured in my paper in 1934.
160
COLOBID.3E.
crest between them and projecting forwards as a long fringe
■over the brow and forming a high, thick tuft or crest on the
crown. Colour of the upper side varying from very dark
brown to pale sepia-brown ; the buttocks may be brown like
Tthe back or whitish and sharply contrasted with it ; the
Kg. 47. — A & B. Front and side views of skull of adult cj Presbytia
jemoralis keatU from Bankachon, Tenasserim. x f.
"underside similarly varies from, brown to white, but the
underside of the tail is always dark, hardly paler than the
upper side, and never strongly contrasted with it ; the limbs
• externally become blackish towards the hands and feet,
PEESBYTIS.
161
but the inner side of the arm may be whitish from the axilla
to the elbow, and the inner side of the thigh is always white
and sharply defined from the dark outer side, the white frequently
extending as a narrow stripe between the knee and heel.
The face is black with the lips flesh-coloured and clothed
with white hair, and the eyelids are also to a certain extent
pale.
18 a. Presbytis femoralis keatii Robinson & Kloss.
Presbytia neglecta keatii, Robinson. & Kloss, Journ. Fed. Mai.
St. Mus. iv, p. 174, 1931.
Pithecus femoralis keatii, Wroughton, Journ. Bomb. Nat. Hist.
Soc. 1915, p. 702 ; Poeock, Journ. Bomb. Nat. Hist. Soc.
xxxii, p. 676, 1928.
Presbytis femoralis keatii, Poeock, Proc. Zool. Soc. 1935, p. 902.
Presbytis robinsoni, Thomas, Proc. Zool. Soc. 1910, p. 634.
Vernacular. — Lutong-koka (Tenasserim) .
Locality of the type, Trang, in northern Malay Peninsula.
Distribution. — From the northern Malay Peninsula (Trang
and Perak) to Tenasserim and S. Siam, west of Bangkok.
Fig. 48. — Inner surface of leg of Presbytis femoralis keatii from Malewoon,
Tenasserim.
Resembling typical femoralis from Singapore and other
districts in the southern portion of the Malay Peninsula
in being entirely darker or lighter brown above from the brow
backwards and on the outer side of the limbs and on the
vol. i. ' M
162 coloeid^:.
buttocks and outer side of the thighs ; but distinguished by
the whole of the underside from the throat to the inguinal
region, being dark brown, without a median white stripe,
and with the narrow stripe on the inner side of the leg usually-
extending to the ankle, whereas in typical femoralis the
underside is dusky greyish-black, relieved by a narrow
white stripe running backwards from the chest, and the white
on the inner side of the hind leg passes only a little way
below the knee.
As I recorded in 1934, two partially albino examples of
this race were collected in the Malay Peninsula. One of them,
from Trang, the type-locality of keatii, Thomas described as
a distinct species, P. robinsoni. It is mostly covered with
* I
Fig. 49. — Head of Presbytia femoralis keatii from Malewoon,
Tenasserim.
a, mixture of grey and white hairs above, but has the hands
and feet dark brown and the end of the tail greyish-brown.
The other, taken at Bandon, in association with typical keatii,
is even paler, being mostly creamy-white, with the hands
and feet pale buff-brown, darkening on the fingers and toes.
Not improbably similar mutants will be found in Tenasserim.
The flesh-measurements (in English inches) of two typically
coloured examples, collected by Shortridge in Tenasserim, are
as follows : —
Malewoon, ad. <J : head and body 22§ ; tail 31f ; hind foot 1\.
Bankaehon, ad. $ : head and body 22£ ; tail 32f ; hind foot 1\.
The weight of the $ was 15£ lb., of the $ 14£ lb.
LEMUROIDEA.
163
The equality in size of the body-measurements between
the sexes is borne out by the skulls, as the following
dimensions (in mm.) show : —
Locality and sex.
— ^
o-"
E-i
Oondylo-
basal
length.
Zygomatic
width.
Orbital
width.
Maxillary
width.
Upper
molars.
(4
ea
•■3 5?
a
Malewoon, Victoria Point, Tenas-
serim ; ad. c?
Bankaehon, Victoria Point, Tenas-
serini ; ad. £
96
93
72 ! 72 \ 64
— 1 76 ! 63
1
72 70 i 62
26 25
29 24
23 24
69
70
69
Malewoon, Victoria Point, Tenas-
The largest known skull, that of an adult $ from west of
Bangkok, in Siam, has a total length of 101 mm. and a condylo-
basal length of 82 mm.
Next to nothing of the habits of this Leaf -Monkey has been
noted, but they probably differ in no respect from those of other
species of the family. The large male from Siam, west of
Bangkok, was said by its collector, K. G. Gairdner, to repre-
sent a purely mountain form ; but the specimens collected
by Shortridge came from comparatively low levels in Tenas-
serim, where it is " apparently not at all plentiful and rather
shy." On freshly-killed specimens Shortridge observed that
the " facial area is very small and quite unlike that of Burmese
and Indian Langurs. The mouth is pinkish-white, but the
orbits are dusky flesh-coloured and not a conspicuous feature."
Suborder LEMUROIDEA.
Distinguished from the Pithecoidea (p. 14) by retaining
the rhinarium, or moist area of naked skin round the nostrils,
with its extension as a narrow strip, or phiUrum, to the edge
of the upper lip, dividing it into a right and left portion, and
by the adherence of the lip in the middle to the gum so that
it is not protrusible. The hands and feet have the fourth
digit never shorter than the third, in almost all cases longer,
and the second digit of the foot is armed with a distinct
claw, quite different from the flat nail of the rest of the
digits. The tongue has a well-developed, underlying lamina,
with serrated free end, the sublingna.
The skull has the orbit communicating with the temporal
fossa, there being no bony partition between the two. The
lower canine is reduced in size, incisiform, and projects forwards
alongside the similarly projecting incisors, the six teeth
m2
164 LOEISID^l.
combining to form a comb-like structure, and the two median
upper incisors are separated by a distinct space.
The Lemttboidea and the suborder Ohieomyoidea, contain-
ing the aberrant Aye- Aye (Chiromys or Daubentonia), which
has rodent-like dentition and other peculiarities, constitute
the Strepsirhine division of the Primates, distinguished from
the Haplorhine division, containing the Tabsioidea and
Pithecoidea, by the structure of the nose and the non-
protrusible upper lip, by the absence of the postorbital parti-
tion in the skull, and by the peculiar arrangement of the lower
incisors and canines. These teeth act as a comb for scraping
the fur and skin, and the function of the sublingua is the
removal of particles from the teeth. The claw on the second
digit of the foot also acts as a scratcher and cleaner of the fur.
The Asiatic LEMtJROiDEA belong to a single family.
Family LORISID^.
Distinguished from the families Lemuridse -and Indrisidse,
which inhabit Madagascar, by the tympanic bone being large
and external to the bulla, of which it forms the outer wall,
not small annuliform and enclosed within it. Also by the
clitoris being traversed throughout by the urethral canal,
which opens at its tip, instead of openly grooved behind, with
the orifice of the urethra at its base.
The tail is very short or absent. The hands and feet have
the first digit (pollex and hallux) relatively very large and
capable of considerable backward extension ; the second digit
is abbreviated, the remaining three being strong, these features,
associated with a short, broad, well-padded palm and sole
and short hairy heel, combining to give a tenacity of grip
unequalled in other families of Primates.
Mammas 4, tolerably widely spaced on the hinder part of
the chest and the fore part of the abdomen.
The skull has the muzzle moderately elongated, with its
upper and lower edges nearly parallel and the anterior nares
terminal ; the orbits are expanded, with thickened rims,
the cranium has a pair of strong temporal ridges, which only
occasionally coalesce, and the occiput does not project far
beyond the condyles. The dental formula is i. §, c. \, pm. |>
m.\; the first lower premolar is large and canine-like, closing
behind the upper canine.
To this family also belong the two genera of African Pottos
(Perodicticus and Arctocebus), distinguished from the Asiatic
by, among other characters, almost complete suppression
of the second digit of the hand. A closely related family is
the Galagidae, comprising the African Galagos, which have
PLATE VII.
MAMMALIA.
Photo W. S. Berridge
Slow Loris (Nycticebus coucang).
Photo F. W. Bond.
Slender Loris (Loris tardigradus).
NYCTICEBUS. 165
the tail and the heel very long, in conformity with prodigious
leaping capacity, in great contrast to the complete inability
to leap exhibited by the Lorises and Pottos.
The two Asiatic genera may be briefly diagnosed as follows : —
ji. Ears small, limbs stout, tail prominent ;
skull with shorter snout and palate, narrower [E. GeoSr., p. 165.
orbits, etc Nyoticebtjs
■6. Ears moderately large, limbs very slender,
tail typically suppressed externally ; skull
with elongated snout and palate, broader [p. 174.
orbits, etc Lokis E. Geofir.,
Genus NYCTICEBUS E. Geoffroy.
Nycticebua, E. Geoffroy, Ann. Mus. Hist. Nat. Paris, six, p. 162,
1812 ; and of subsequent authors.
Type of the genus, bengalemsis E. Geoffroy.
Distribution. — S.E. Asia from. N.E. Bengal, Assam, Uppek
Bukma, and Laos southward to Annam, the Malay Peninsula,
and through the Sunda Islands to Borneo.
Body robust ; limbs stout, with powerful hands and feet
and thick digits ; head rounded, the muzzle rounded, with
the upper jaw and nose not very noticeably projecting beyond
the lower jaw ; ears reduced, nearly concealed in the fur,
the margin rounded, the supratragus valvular and surmounted
by a small supplementary ridge and small pocket-like sub-
marginal depression just above the position of the suppressed
antitragus ; tail always present, but conical and short, from
$ to 1 in. long. Skull with interorbital septum moderately
broad ; orbits expanded, but not so wide across as the
zygomatic arches ; the nasals and premaxillas only slightly
produced beyond the level of the upper incisors ; the palate
not prolonged beyond the level of the last molars, and the
median upper incisors larger than the laterals, which may be
absent.
Until 1902 the Slow Lorises were cited by all writers,
except a few of early date, as Nycticebws tardigradus. But
in that year Stone and Rehn (Proc. Acad. Nat. Sci. Philad.
lv, p. 141) pointed out that the name tardigradus Linn,
belongs to the Slow Loris of Ceylon. They substituted
couoa/ng Bodd. for the Slow Loris, admitting a considerable
number of local races, which, with many others subsequently
described, were cited as " species " or " subspecies," according
to the fancy of authors. But there is abundant evidence,
in my opinion, that many of these so-called " species " or
" subspecies " were based on purely individual differences,
often due to seasonal changes in the coat and colour, and
I here adopt the conclusion that there is but one species of
Nycticebus, for which the oldest known name is coueang.
166 LOEISIDJE.
19. Nyeticebus eoucang (Boddaert).
Tardigradus eoucang, Boddaert, Eleneh. Anim. p. 67, 1785 (Tardi-
gradus preocc). (For other references and synonymies, see
under the subspecifie headings.)
Coat, in good condition, consisting of copious underwool
and abundance of longer, stiffer, but soft contour hairs.
General colour very variable, but the back, in good coat,
always ornamented with a conspicuous dark spinal stripe
extending sometimes from the root of the tail and reaching
the crown, where it may be narrow and cease or broad and
branched, and extend laterally to the ears and eyes, the areas
around those organs being always darkish-tinted ; the forehead
in the middle line, the interocu'lar area, and the muzzle always
white ; the cheeks and temples paler than the crown, some-
times white. When these white areas and the dark stripes
to the eyes and ears are well denned they constitute a charac-
teristic facial pattern or ' ' mask. ' ' But sometimes the extension
of the white areas over the crown obliterates the stripes,
leaving the whole head pale, except for the tinted areas round
the eyes and ears and the median stripe on the crown, and
every stage in the development and obliteration of the darker
markings may be traced.
The three British Indian races of the Slow Loris here
admitted may be briefly distinguished as follows : —
or. Head and neck pale, mostly whitish or grey, the
median stripe on the nape and crown narrow,
with at most faint stripes to the eyes and ears.
a'. Stripe ceasing on the middle of the crown,
cut off from the coloured areas round the [(Fischer), p. 166.
eyes and ears ; size larger bengalensis
b'. Stripe typically bifurcating and passing to
the eyes, stripes from the ears nearly reaching [Elliot, p. 169.
the point of bifurcation ; a little smaller .... tenasserimensis
b. Head and neck mostly well coloured, the stripe
broad on the nape and crown and extended
broadly forwards to the eyes and laterally
to the ears: general colour brighter and size [p. 171.
still smaller eoucang (Bodd.),
19 a. Nyeticebus eoucang bengalensis (Fischer).
Loris bengalensis, Fischer, Anat. Maki, p. 30, 1804 ; Tiedemann,
Zool. i, p. 334, 1808.
Nyeticebus bengalensis, Geoffroy, Ann. Mus. d'Hist. Nat. xix»
p. 164, 1912 (in part).
Nyeticebus tardigradus, Fischer, Syn. Mamm. p. 71, 1829 ; and
of many later authors, at least in part, including Blyth, Jerdon,
Anderson, Blanford, and other writers on Indian Mammals
till 1902 (not Loris tardigradus, Linn.).
Nyeticebus cinereus, M.-Edwards, Now. Arch. Mus. Bull, iiiv
p. 9, 1867 ; and of Anderson (1881), Elliot (1912), Osgood (1932)
as a subspecies.
KYCTICEBTJS. 167
Nycticebus incanus, Thomas, Ann. Mag. Nat. Hist. (9) viii,
p. 627, 1921.
Nycticebus bengalensis, Thomas, Journ. Bomb. Nat. Hist. Soo.
xxviii, p. 433, 1922.
Type, " Le paresseux pentadaetyle du Bengale," Vosrnaer,.
Deseript. Anim. p. 18, pi. 6, 1770.
Locality of the type of bengalensis, Bengal ; of cinereus,
Bangkok ; of incanus, Kyeikpadein, Pegu.
Distribution. — Assam*, Teppebah, Chtttagong, Uppeb
and Lowee Btjbma, Siam, Laos, and Annam.
Distinguished by its larger size and pale head, the median
stripe ceasing in the centre of the crown and not extended
forwards or laterally to join the coloured areas round the eyes
and ears ; at most faint, abbreviated stripes extending upwards
from those areas towards it.
General colour vary variable. Head, nape, and usually
at least the fore part of the shoulders white, cream or grey,
except for the median brown stripe, which ceases in the middle
of the crown, and for faint stripes from the eyes and ears,
which never meet the median stripe. Spinal stripe broad or
narrow, varying from blackish-brown to ochreous-brown,
sometimes traceable to the rump, sometimes not beyond the
middle of the back, commonly more or less emphasized
anteriorly by grey or silvery hairs continued from those of the
shoulder. Planks and back varying from sepia-brown
through all shades to ochreous, buffy or dark grey. Arms
to elbow varying with the tint of the shoulder from brownish
to whitish, below elbow to hand typically paler, whitish ;
legs varying from brown to nearly white ; hands and feet
always quite pale. Lower side varying from whitish to ashy-
grey or pale buff. The wool of the dorsal side slaty at base.
Li a large series of skins ranging from the Naga Hills to-
Lower Pegu and thence eastward to Siam, Laos, and Annam
no two are exactly alike, even from the same district, the
variations in colour being mainly due to the condition of the
coat, whether fresh or faded, and especially to the presence
or absence of the contour hairs, which when moulted leave
the coat entirely woolly and when abundant give a silvery
sheen to the back, particularly alongside the dark spinal
stripe, which they throw into strong relief.
Vosmaer's figure and description of the type show that the
* Annandale (Proc. Zool. Soo. 1908, pp. 888-9) recorded, on the
evidence of La Touche, the existence in the Lushai Hills of a Lemur
he considered to be undescribed because it had a bushy tail ; and in
support of his belief he published a photograph of the animal suspended.
Sloth-fashion, from a branch. The photograph clearly _ represents
a typical example of N. coucang bengalensis. A bushy tail up to an
inch in length, not including the terminal hairs, is a well-known feature
in Nycticebus.
168 LOEISID^B.
general colour was " grey or clear ashy-yellowish," a little
richer and more ochreous on the flanks and thighs. Two
skins from the Naga Hills are better coloured, the whole
of the upper side behind the shoulders, apart from some grey
Lairs alongside the spinal stripe, being pale ochreous-brown.
One of them, labelled " Naga Hills," has the abdomen buffy
or pale brown in the middle, whereas the other, labelled
" Champang," which is a little brighter on the flanks and rump,
has the abdomen mainly whitish. A skin from Bassein in
Lower Pegu (November 22) closely matches those from the
Naga Hills, but has the flanks paler, thus approaching the
type of bengalensis, and one from Kindat, on the east
bank of the Chindwin (December), differs from the skin
from the Naga Hills in having the nape and shoulders ashy,
the flanks and rump brown. These skins are in good coat,
but in one taken 50 miles north of Pegu (June) the coat is
all woolly, without contour hairs, the whole of the back
being ochreous or buffy, without silvering, the arms and legs
paler and the belly pale buff. The type of incamis, from
Kyeikpadein, Pegu (August) is, on the other hand, mostly
grey, with no ochreous or brown except a little on the rump,
the arms, legs, and underside being ashy, but there is some
silvery-grey on the head and alongside the spinal stripe.
The type of cinereus, from Bangkok, was described as clear
ashy-grey mixed with golden-brown on the back and rump ;
but a co-type in the British Museum is in moult, and is mostly
covered with soiled grey wool, but some ochreous-brown new
hair erupting in patches on the flanks suggests that the skin
would have resembled those from the Naga Hills when the
coat was fresh. A skin from Taungyi in Thaton closely
matches this co-type of cineieus. Anderson identified as
N. tardigradus var. cinerea skins from Goalpara in Assam,
from Tipperah, and from the Kakhyen Hills east of Bhamo.
A skin from Xien Quang Koo, Laos, named bengalensis
by Thomas, was determined by Osgood as bengalensis cinereus ;
and a much more extensively grey skin from Lao Ba in Annam
was referred by both these authors to cinereus. I am unable
to find any reliable character by which cinereus can be dis-
tinguished from bengalensis.
The skull is the largest of the genus, with the condylobase
exceeding 60 mm. In the co-type of cinereus, ? adult $, from
Bangkok, it is 61 mm., only 1 mm. shorter than in the skull
from Toungoo, and the cranial ridges meet to form a low
sagittal crest as in the # from the Naga Hills. But in another
adult,probably £, skull from Raheng the condylobase is 63 mm. ,
but the total length is 68| mm., 4| mm. longer than in the
skull from Bangkok, and the cranial ridges form a crest 4 mm.
iigh. This suggests that the occipital and sagittal crests
NYCTICEBUS.
169
increase with age after the condylobase has attained its full
length. There is no difference in size between <$ and 2 skulls,
but the cranial ridges seem to be farther apart on the average
in the 2.
19 b. Nyctieebus coucang tenasserimensis Elliot.
Nycticebus tardigradus, Blanford, Mamm. Brit. Ind. p. 45, fig. 12,
1888 (Tenasserim variety).
Nycticebus tenasserimensis, Elliot, Bev. Primates, i, p. 25, 1912.
Locality of the type, Amherst, N. Tenasserim.
Distribution. — Tenasserim and S.W. Siam.
Distinguished from N. coucang bengalensis by the bifurca-
tion of the stripe on the crown to join the dark area round the
Fig. 50. — Tickell's Slow Loris (Nycticebus coucang tenasserimensis) from
Amherst. (Adapted from Tiekell's figure, but with the frontal
stripes too pronounced.)
eyes, by the extension of the stripe above the ear almost to the
point of bifurcation, and by the dark hue of the side of the
neck, which is the same tint as the upper arm and confluent
with the patch over the ear.
The type of this provisionally admitted race is the figure
of an adult <J Slow Loris from Amherst recorded by Tickell
170 LORISID.SI.
in his MS. A considerably altered copy was published by
Blanford as representing the Tenasserim variety of N. tardi-
gradus, and on this evidence Elliot established his " species."
Quoting Blanford, he said the general colour was " pale
rufescent." This is incorrect. In TickelTs description it
is said to be " pale tawny, a little dulled with greyish-
brown, becoming greyish-white on the hands and feet, and
almost greyish-white on the occiput and nape, with the
middle of the back, the rump, the arm to the elbow, and
the thighs orange tawny. . . ; the dorsal line is continued
from the crown as two faint lines to the space round the
eyes ; the sides of the neck, like the shoulders, are reddish-
tawny, and from the same tint enveloping the ears passes
a stripe which nearly reaches the vertex of the crown."
The description and figure of this specimen agree very closely
with typical N. couoang bengalensis, except in the particulars
mentioned in the diagnosis. In these the specimen approaches
the next race, but the stripes on the head are not so broad and
pronounced, the whole crown being whiter. It is a transitional
form. Tickell stated that this Loris is found throughout
Tenasserim and in Arakan as well, but no doubt he did not
distinguish the Amherst specimen he described and figured
from the Burmese Lorises I identify as bengalensis.
A specimen from Mergui Town (Shortridge) (April) is
referred to this race. The coat is all woolly, and the general
colour above is drabby- brown from the shoulders to the
elbows and rump, with the legs a little greyer ; but the cheeks,
temples, and the whole of the neck are deep ashy-grey, as if
the hairs had lost their white tips. It differs from the type
of tenasserimensis in the stripes from the eyes just failing to
reach the stripe on the crown.
At Koh Lak in Peninular Siam Kloss (Journ. Nat. Hist.
Soe. Siam, ii, p 77,. 1916, and p. 289, 1917) collected two
Slow Lorises which he identified as N cinereus. One of them
agrees almost exactly in colour with examples of N. coucang
bengalensis from the Naga Hills, but has a smaller skull,
measuring only 60 mm. in total length, the same as in many
skulls of the next race from, the Malay Peninsula. The
other, in the extension of the cranial stripe to join the coloured
areas round the eyes and ears, comes close to the type of
N. c. tenasserimensis, and its skoll, measuring 62 mm. in
total length, is almost the same as in the example assigned
to N. c. tenasserimensis from Mergui Town.
The skull, judging from the scanty records, is a little
smaller on the average than in bengalensis, and thus approaches
skulls from the Malay Peninsula, in five of which the condylo-
basal length ranges from 54 to 56 mm. The measurements
of the skull of the type are taken from Tickell's figure, drawn
natural size.
PLATE VIII.
MAMMAL
A. Face of Nycticebus coucang bengalensis.
B. „ Nycticebus coucang tenasserimensis.
C. „ Nycticebus coucang coucang.
NYCTICEBUS. 171
19 c. Nyetieebus coucang coucang (Boddaert).
The tailless Macauco, Pennant, Quadr. ed. i, p. 212, no. 128,
pi. 26, 1781.
Tardigradus coucang, Boddaert, Elench. Anim. p. 67, 1785 *.
Nyetieebus coucang, Thomas, Journ. Bomb. Nat. Hist. Soc. xxviii,
p. 433, 1922.
Locality of the type not recorded, but, according to Thomas,
Java.
Distribution. — The Mergui Archipelago, Malaya, Sumatra,
Java.
Distinguished from the preceding two races by having
the head, nape, and shoulders better coloured, not so white,
■with the stripe on the crown broad and dividing into four
broad branches extending respectively to the ears and eyes.
The size, also, both of the body and of the skull, is a little
smaller, and the general colour on the average brighter.
The only British Indian representative of this race known to
me is a (J from King Island, Mergui Archipelago (Primrose),
(December). The general colour is rich, rusty-ochreous
above and below, becoming gradually paler distally on the
arms and legs, but not white on the hands and feet, and with
the chest a little greyer than the belly ; the spinal stripe
is well defined by patches of very deep brown on the fore
back and between the shoulders, but is paler and redder
on the nape, which is buffy-grey laterally ; the centre of the
crown is ochreous, and the stripes are defined by ashy-grey
on the forehead and cheeks.
Th : s specimen closely matches an example from Perak
in the Malay Peninsula, except that the sides of the neck
are greyer and the crown less extensively reddened. This
specimen belongs to the form usually quoted as malaianus
Anderson, which, in my opinion, is indistinguishable from
javanicus Geoffroy.
According to Blyth (Journ. As. Soc. Beng. xvi, p. 735, 1847)
this " darker coloured, more rufous Malayan form," which he
cited as his var. B of N. tardigradus, was brought to Calcutta
for sale by vessels from the ports of Arakan, Burma, Malaya,
* This species was based by Boddaert upon two distinguishable
forms, " The tailless Macauco " of Pennant and " Le paresseux penta-
daetyle du Bengale " of Vosmaer. When Stone and Rehn revised the
genus (Pioc. Acad. Nat. Sei. Philad. liv, p. 141, 1902) they took as the
type of coucang the Bengal Loris described by Vosmaer ; and in this
they were followed by some later authors, including Elliot, until 1922,
when Thomas pointed out that in accordance with the generally
accepted custom of selecting the form, first quoted under the name of
a species as its type, the " tailless Macauco " of Pennant is, ipso facto,
the type of coucang, and this Loris is not the larger, white-headed
northern form, bengalensis, but the smaller, more southern form with
the well-coloured head.
172 LORISIDJE.
and Singapore. He added that it is rare in Eastern Bengal,
but I can find no evidence that he had a specimen from that
district. When Anderson described it as N. tardigradus var.
malaiana, based on specimens from Malacca and Penang,
he said it occurs in Cblttagong and Arakan, but he cited no
specimens in support of this statement (Cat. Mamm. Calc.
Mus. p. 95, 1881). I do not believe it is found so far north.
The only available flesh-measurements (in English inches)
and the weights (in lb.) of these British Indian races of the
Slow Loris are as follows : —
Head and
Locality, name, and sex. body. Tail. Foot. Weight.
Kindat (bengalensis) ; ad. <J... 14f $ 3— 2£
50 m. N. of Pegu (bengalensis) ;
ad. (J 14$ 1 3 —
Laos (bengalensis) ; ad. <J 14 $+ 2f —
Amherst (tenasserimensis type) ;
ad. ^ 12£ — — —
Mergui Town (tenasserimensis) ;
ad. <J 13£ f 2f 3£
Koh. Lak, Siam (tenasserimen-
sis); ad. tf 11 * 2f —
These measurements suggest that bengalensis is a little
larger than tenasserimensis. The specimen assigned to
typical coucang from King Island was unmeasured, but six
adult $ examples from various localities in the Malay Peninsula
range in length of head and body from 11 to 12| in., almost
the same as in the three recorded examples of tenasserimensis.
Habits. — Very little is known about the breeding of
Nycticebus, except that a single young one is, usually at all
events, born at a time. In general habits there is probably
no difference between the various races ; but accounts have
mostly been taken from the behaviour of captive specimens,
observation of wild individuals being almost impossible owing
to the species being exclusively nocturnal and an inhabitant
of dense forests, where it seldom leaves the trees. According
to Tickell it is slow in its movements in trees, but climbs
readily with a tenacious grip, and on the ground progresses
with a wavering kind of trot. By day it sleeps rolled up in
a squatting attitude, with its head and hands buried between
its thighs. When first caught it is savage and bites quickly
and severely, with a grunt or growl. It feeds on plantains
and other fruits and greedily devours insects of many kinds.
In captivity it may become diurnal. One that Tickell had
attempted to catch a cockroach some 10 or 12 inches distant
-on the floor of a room. The Loris stealthily stalked the
insect till within striking distance, then, raising itself on its
hind legs, " flung itself " at its prey with a swift grab — but
missed it.
NYCTICEBtfS.
17a
g
s
10
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&
CS
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<3
o
1
a
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to
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174 LOEISIDJE.
This account has since been confirmed and extended.
Shortridge recorded that the specimen he collected on Mergui
Island, where the animal is apparently fairly well known,
was very savage, growling like a cat when approached, and
always ready to attack anything placed near it. According
to Mackenzie the animal moves along a branch in a most
peculiar way, very slowly and deliberately, putting the hind
foot of one side right up to the hand, then moves the hand,
then the hind foot of the opposite side, then the hand, and so
on, bending the back the while in a most extraordinary way.
A specimen from Thaundaung, kept in captivity, never
voluntarily came to the floor of its cage, but would hang
from a branch by its hind legs to pick up anything and sometimes
to drink. It stalked prey in the way described by Tickell,
then seized it with a sudden pounce, usually grabbing it
between the fingers and palm, sometimes stretching to its
fullest extent to reach it, but always retaining the hold of its
hind feet. It ate rice, fruit, insects of many kinds, its favourites
being Longicorn beetles, cockroaches, crickets, and grass-
hoppers, and it was seen to catch moths on the wing. The
stomach of a specimen captured near Kindat contained leaves
and shoots, a beetle, and a piece of bone, apparently of a bird.
The Chins, Mackenzie tells us, use the fur to stop bleeding,
and say it is very efficacious.
Genus LORIS E. Geoffroy.
Loris, E. Geoffroy, Mag. Eneycl. i, p. 48, 1796 ; and of all recent
authors, including Osman Hill, Ceyl. J. Sci. (B), xviii, pp. 89-
132, 1933.
Type of the genus, gracilis, E. Geoffroy (=tardigradus
Linn.).
Distribution. — S. India and Ceylon.
Distinguished from Nycticebus by its more slender build
and relatively much longer and thinner limbs, smaller hands
and feet and less powerful digits, the hands smaller as com-
pared with the feet ; nose projecting prominently beyond
the lower jaw ; ears similar in structure, but less reduced,
not so concealed by the fur, with the supratragus and flap
above it larger, more completely valvular, the flap which
bears the deeper pocket-like depression larger, the margin
of the pinna not so rounded, more or less angled anteriorly*,
tail only very occasionally a few mm long. Skull with
interorbital septum very narrow, orbits more expanded,
* Osman Hill was the first to dseribe the ear of Loris, taut I cannot
agree with his estimate of its peculiarities ; and the figures he published
of the ears of Nycticebus, Loris, and Tarsius are quite unlike the ears
of the examples of those genera I have seen.
LORIS.
175
exceeding the width, across the zygomatic arches ; nasals
and premaxillse produced in subtubular form beyond the
level of the upper incisors ; the palate produced beyond the
level of the last upper molars, and the upper incisors subequal
in size.
Like Nycticebus, Loris contains a single species.
Vfe'
Fig. 51.
A. Side view of head, with ear exposed, of a Slender Loris (Lorh
tardigradus) from Ceylon.
B. The same of a Slow Loris (Nycticebus coucang).
C. Rhinarium, from the front, of Slender Loris.
D. The same of Slow Loris.
E. Palmar surface of right hand of Slender Loris.
F. Sole of the foot of the same.
20. Loris tardigradus (Linnaeus).
Lemur tardigradus, Linn. Syst. Nat. ed. 10, p. 29, 1758. (For
synonymy and bibliographical references, see under the sub-
species.)
Colour more uniform than in Nycticebtis, the dorsal side, in
good coat, varying from dark grey with silvery contour hairs
to earthy-brown, the ventral side from white to bufiy, the
176
LOBISIDiG.
dark spinal stripe never strongly pronounced, very frequently
traceable, at least on the back, behind the shoulders, but
often undeveloped ; the head -with the crown coloured like
the back, but, as in Nyctkxbus, the muzzle and the area between
Fig. 52. — The Slender Loris (Loris tardigradus).
the eyes are white and the area round the eyes dark, blackish
or brown ; but the colour may vary considerably when the
coat is in moult.
Key to the Races of L. tardigradus.
. General colour above, in good coat, dark grey,
considerably silvered, below typically white,
at least the tips of the hairs on the cheeks
white like the interorbital stripe and em-
phasizing the ocular patch.
a.' Hairs of abdomen and cheek white through-
out, or of abdomen faintly grey at base,
c. Size largest ; in a large percentage of
specimens the hairs of the abdomen [Cabrera, p. 177.
white to the base lydekkeriamus
c'. Perhaps on the average smaller, with a
larger percentage of specimens with
hair on abdomen faintly grey at the
base nordious Hill, p. 182.
&'. Hairs of cheeks and abdomen sooty in basal fKps, p. 184.
half grandis Hill & Phil-
. General colour above browner, more ochreous
in. the pelage and less silvering on the average ;
cheeks darker than interocular stripe, the
ocular patch less emphasized. [Wroughton, p. 180.
d. Larger, a little paler brown maUxbaricus
d'. Smaller, a little darker, more rufous-brown, tardigradus (Linn.),
[p. 181.
LORIS. 177
A point of interest connected with these races is the occur-
rence both in India and Ceylon of dry- and wet-zone forms,
the wet-zone form in each case being more richly tinted,
more rufous or ochreous than the dry-zone form, and also-
a little smaller.
20 a. Loris tardigradus lydekkerianus Cabrera.
Loris tardigradus typicus, Lydekker, Proc. Zool. Soo. 1904, p. 345-
Loris lydekkerianus, Cabrera, Bol. Soc. Esp. Hist. Nat. 1908,
p. 135 ; Thomas, Aim. Mag. Nat. Hist. (8) i, p. -469, 1908 ;
Wroughton, Journ. Bomb. Nat. Hist. Soc. xsv, p. 45, 1917,
and p. 563, 1918.
Loris tardigradus lydekkerianus, Hill, Ceyl. J. Sei. (B), xviii,
p. 124, pi. 7, fig. 2, 1933.
Vernacular (Shortridge). — Kada-Papa, Adavi-Papa (Kanar-
ese) ; Wanur-Manushya (Marathi) ; Arawe-Papa (Telegu) ;
Kattu-Papa and Kattu-Pullaye (Tamil) ; SMrminda (Dekhani
and Hindustani).
Locality of the type, " Madras."
Distribution. — The Eastern Ghats, westwards to Mangalore
and Mysore. Precise range to the north unknown.
Size, on the average, larger than in other races of tardi-
gradus. Colour of the upper side, in good coat, typically
greyish-brown, silvered to a varying extent, especially on
the fore back and shoulders, and with indications of a darker
median line on the fore back, the fur at the base being deep
slate-grey ; the flanks paler and passing into the usually
clean white abdomen, the hairs of which are, with rare
exceptions, white throughout ; the cheeks entirely white,
like the throat and chin ; the limbs proximally about the
same hue as the upper side, but becoming paler distally,
especially the arms, which are whitish below the elbow ;
hands and feet white above.
The type of this race, labelled '"Madras," and presented
by Edgar Thurston of the Christian College, was probably
brought as a live specimen from the Eastern Ghats.
A good series of skins of this Loris collected by G. C.
Shortridge in the Kolar district of E. Mysore, 2,700-3,000 ft.
(October 12-20), shows some variations from the normal in
colour. One skin is paler, more drabby, with no silvery
tint above, and the belly has a yellowish tinge, with the hairs
pale grey at the base. Another, in bad moulting coat, has also
lost its silver-tipped hairs, and the general hue above is dark
grey and buffy in patches. Two of them have the hind back,
loins, and legs covered with pale dull ochreous or buffy wool,
and sharply contrasted with the dark grey hue of the fore
back and shoulders. Of three specimens collected by Baptista
VOL. I. 3ST
178
LOBISID-ffi.
in the Chettiri Range, Salem, 2,000 ft. (June 8th), one has
a decided bufly tinge in the dorsal pelage, thus approaching
the Malabar race.
Fig. 53.
A. Dorsal view of skull of adult (J Loris tardigradus lydekkerianus
from Mysore, x |.
B. Front view of muzzle of same, showing the subequal incisor teeth.
C. Side view of end of muzzle of same, showing its projection beyond
the upper canine.
D. Dorsal view of skull of adult <J Nyctioebus coueang bengalensis from
the Chin Hills.
E. Front view of muzzle of same, showing the unequal incisor teeth.
F. Side view of the end of the upper and lower jaws of the same.
c, upper and lower canine teeth ; pm, upper and lower first pre-
molar.
■G. Lower teeth of the right side of the same, i, incisors, forming with
c, the canine, the forwardly directed comb ; pm x , canine-like first
premolar ; m„ first molar.
H. Upper teeth of the left side of the same, i, incisors ; c, canine ;
pm 1 , first premolar ; to 1 , first molar.
LOEIS. 179
There is very little difference in size between the sexes,
the available evidence pointing to slight superiority of the <$.
The flesh-measurements (in English inches) and weights
(in oz.), according to Shortridge and Baptista, are as follows : —
Head and Hind
Locality and sex. body. foot. Weight.
E. Mysore ; ad. <J 10$ 2 J llf
E. Mysore ; ad. «J 9$ 2| 9$
E. Mysore ; ad. (J 10 2i 11
E. Mysore ; ad. ? 9$ 2+ 91
Salem ; ad. <J 9+ 2-! 8~
Salem ; ad. <J 9 2i S
Salem ; ad. 2 8$ 2 8
One <$ weighed by Shortridge was 12J oz.
Habits. — Shortridge made som einteresting observations on
wild and captive specimens of this Loris in Mysore. The iris
of the eye is chestnut, the pupil contracts to a nearly vertical
slit, and when expanded shines with a golden-coppery light.
Wild specimens in foliage are strikingly like the little Spotted
Owl (Athene brama), have the same habit of swaying the head
and, when fighting, utter a similar screech. They also make
a faint chattering noise and a low growl when irritated.
Their quarrelsomeness when kept together in a cage shows
that they are not naturally gregarious. They -would occa-
sionally drop to the floor of the cage from a branch 3 or 4 ft.
up, but are quite incapable of jumping even the shortest
distance from branch to branch. Their movements along
a branch are usually slow and deliberate, but much less so
than in Nycticebus. On the ground the movement is usually
exactly like that of a monkey walking slowly, but they
can run on the ground at quite a respectable pace, and
Shortridge thinks that in Mysore they probably travel some
distance from tree to tree. They are unable to swim, and
when put into water merely move their limbs backwards and
forwards without making any progress. They ate cooked rice
and bananas and were particularly fond of grasshoppers, which
they w-ould hold in one hand and munch. In the wild they
are said to feed largely on insects and small lizards. A captive
specimen kept by Kinnear killed and ate the heads of a jerboa
and a gerbille, which were in the same cage. Females with
two young were found on several occasions, and Shortridge
thinks that twins are probably a common occurrence. On
one occasion he captured a female with a three-quarter-grown
young one clinging to her.
il jj
180 LOKISIDiE.
20 b. Loris tardigradus malabaricus Wroughton.
Loris tardigradus, Ryley, Journ. Bomb. Nat. Hist. Soc. xxii,
pp. 284 and 494, 1913.
Loris malabaricus, Wroughton, Journ. Bomb. Nat. Hist. Soc. xxv,
p. 45, 1917, and xxv, p. 563, 1918.
Loris tardigradus malabaricus, Hill, Ceyl. J. Sei. (B), xviii, p. 127,
1933.
Vernacular names in Coorg (Shortridge) . — Hunimunna,
Singalika, Kard-miinishya.
Locality of the type, Huvinakadu Estate, Kutta, S. Coorg,
2,843 ft.
Distribution. — The Malabab district of S. India, the
Wykaad, S. Coobg, and Tbavancobe.
Distinguished from lydekkerianus on the average by its
smaller size, tawnier or more ochreous tinge of the upper side,
slightly darker underside and cheeks, the latter a little darker
than the throat and chin, and the area round the eyes more
rufous-brown.
The general colour is subject to a good deal of individual
variation. The type from Kutta, S. Coorg, has the dorsal
fur tawny-brown superficially, deep slate-grey at the base,
with some silver-tipped hairs on the back, especially on the
shoulders, and also on the hind legs ; the lower side is bufly-
white, the hairs pale grey close to the skin, the throat white,
and the cheeks grey and contrasted. Another from Virajpet
S. Coorg, is more richly tinted than the last, the dorsal pelage
being decidedly ochreous-brown with hardly any silvering.
This specimen is very like some specimens of typical tardi-
gradus. A third, from Trivandrum, is darker, not so tawny
as the type, more silvery on the fore back, with the cheeks
paler, less contrasted with the throat, and the hairs of the
Belly whitish at the base. This approaches lydekkerianus
and the northern Ceylonese form nordicus.
The flesh-measurements (in English inches) of three speci-
mens, according to Shortridge, are : —
Head and Hind
Locality and sex. body. foot.
S. Coorg ; ad. g 8f If +
S. Coorg ; ad. ? 8§ If
S. Coorg (type) ; $ 8| If
The weight of the adult S was 6 oz.
Although the flesh-measurements and weight indicate
a smaller race than lydekkerianus, there is very little difference
in the size of the skulls as shown in the table of measurements
(p. 183).
According to a note accompanying Shortridge's specimens,
this race, although apparently rare on the western side of"
India as far north as North Kanara, is probably fairly plentiful
in Coorg ; but it is not easy to find in the thick jungle owing
LORIS. 181
to its nocturnal habits and concealment by day in the thick
foliage of high trees. In some specimens, he observed,
a slender tail, up to 7 mm. long, may be present.
20 c. Loris tardigradus tardigradus (Linn.).
Lemur tardigradus, Linn. Syst. Nat. ed. x, p. 29. 1758.
Loris gracilis, Geoffrey, Mag. Eneyol. (Ann. 4), i, p. 48, 1796 ;
and of most subsequent writers to 1908, including Blyth,
Jerdon, Anderson, and Blanford.
Lemur ceylonicus, Fischer, Anat. Maki, xii, p. 28, 1804.
Loris gracilis zeylanicus, Lydekker, Proc. Zool. Soe. 1904, p. 345*.
Loris tardigradus, Thomas, Ann. Mag. Nat. Hist. (8) i. p. 469,
1908.
Loris tardigradus tardigradus, Hill & Phillips, Ceyl. J. Sei. (B),
xvii. p. 109, 1932 ; Hill, Ceyl. J. Sei. (B), xviii, p. 113, 1932 ;
Phillips, Man. Mamm. Ceylon, p. 31, 1935.
Vernacular. — Geylon Sloth : Unahapuhiva (Sinhalese) ;
Theoangu (Tamil).
Locality of the type, " Ceylon " ; of ceylonicus, Ceylon ;
of zeylanicus (here selected), Peradeniya.
Distribution. — Low-country wet zone of CEYiiOisr.
Resembling malabaricus, but smaller and apparently rather
more rufous-brown above, without silvering, and brighter
below, with the hairs extensively sooty at the base, brighter
or duller yellowish distally, the throat whitish or yellow,
the cheeks dusky and the area round the eyes brown.
This description, taken from two unmeasured specimens
in the British Museum, one being the selected type of zeylanicus
Lydd. from Peradeniya, agrees in essentials with the long
description published by Hill, who states that the lumbar
and sacral regions are sometimes frosted, but less in the 2 than
in the <J, the $ being further distinguished by being larger than
the <J on the average and less rufous in general colour above.
The approximate dimension (in English inches) of his largest
and smallest <J and $ specimens and the averages of several
are as follows : —
Head and
Locality and sex. body. Foot. Ear.
Colombo ; ad. <J S 1* 1 —
Henaratgoda ; ad. (J 7J If 1
Average of 7 ad. <$J 7f l| *
Loe. ? ; ad. O 8£ It *
Colombo; ad. 2 7§ l| 1-
Average of 3 ad. Q 8+ if f
* In his description of zeylanicus the only specimen Lydekker
mentioned individually was a mounted one recently received from
Ceylon. This was marked " type " by Thomas. It is a representative
of nordicus, resembling those collected by Mayor at Anuradhapura.
But it is not the type of zeylanicus, because Lydekker's description was
clearly taken from one of the two or perhaps both the " reddish "
examples of typical tardigradus then in the British Museum, and does
not apply in any way to the mounted specimen, of which the general
tint is brownish-grey with a white belly. Since one of the two specimens
Lydekker described came from Peradeniya, I select it as the type.
182 LORISIDiE.
The weights, according to Hill, are : <J, 3 to 4-| oz. ; $, 3 to
4 oz. ; the average being 3£ oz.
There are no skulls in the British Museum certainly assign-
able to this race, but, according to Hill, <J skulls range from
47 to 50 mm. and $ skulls from 48 to 50 mm. in total length,
the average of seven adult skulls being 48^ mm.
20 d. Loris tardigradus nordieus Hill.
Loris tardigradus nordieus, Hill, Ceyl. J. Sei. (B), xvii, p. 117,
1933 ; Phillips, Man. Mamm. Ceylon, p. 36, 1935.
Vernacular. — The same as for the preceding Ceylonese race.
Locality of the type, Talawa (50 ft.), N.C.P., Ceylon.
Distribution. — The dry zone of the N.P., N.C.P., and C.P. of
Cpylcot from just above sea-level up to 650 ft.
Possibly distinguishable from lydekkerianus by being
slightly smaller and by having the fur in the middle of the
ventral surface on the average greyer at the base, but otherwise
the coloration is apparently the same.
In his key to the races of L. tardigradus Hill distinguished
nordieus, the largest of the Ceylonese forms, from lydekJcerianus,
■the largest of the Indian forms, by the greyness of the basal
third of the ventral hairs, which he alleged to be white through-
out in the Indian race ; but these hairs are occasionally grey
basally in the latter, e. g. in the type, and may be pale through-
out in Ceylonese specimens.
Approximate flesh-measurements (in English inches) are
as follows : —
Locality and sex.
Wilaehcliyia (Willichia) ; ad. g
Sigiriya ; ad. g
Talawa ; ad. $
Tammanewa ; ad. !j>
Talawa ; ad. Q
Talawa (type) ; yg. ad. $> ....
Head and
ody.
Foot.
Ear.
9i-
2*
H
H
It
8|-
2 —
i+
9
2-
H
8f
lt
i*
Si
i£
The average weight of 4 adult $ specimens, according to HilL
is 8 J oz. Both in measurements and weights these specimens
agree very closely with those of lydekkerianus collected by
Baptista on the Chettiri Range, Salem, which from their
locality may be considered almost as topotypes of lydekkerianus-
recorded from " Madras."
Between the skulls of nordieus and lydekkerianus, as entered
in the table, there is practically no difference. But in a long
series of skulls, lent to me by Sir F. Colyer, from Monaragala,
in the dry zone of Uva, the average total and condylobasal
lengths of 6 adult g$ is 51 and 41f , and of 8 adult $$ 49| and
41 mm. Unfortunately there are no skins ; but Hill, who knew
by report of the occurrence of Loris at this locality, thought
LOKIS.
183
S.S
J3
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3
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ri
a
W
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s
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1
o
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«p 45 4s 4-i 43 j j r
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H h is h h "■» '- 13 "3
184 LORISIDJS.
they would prove to be nordicus. If this guess was correct,
the series of skulls from Monaragalla would make the average
size of the skulls of nordicus less than of lydekkerianus. The
skulls in question are intermediate in size between those of
typical tardigradus and of nordicus.
20 e. Loris tardigradus grandis Hill & Phillips.
Loris tardigradus grandis, Hill & Phillips, Ceyl. J. Sci. (B), xvii.
p. 110, 1933 ; Hill, Ceyl. J. Sci- (B), xviii, p. 117, pi. 7.
fig. 1 & pi. 8, fig. 2, 1933 : Phillips, Man. Mamm. Ceyl. p. 34.
pis. 4 & 5, 1935.
Vernacular. — The same as for the typical race, with the
addition of Kalu imahapuluva (Sinhalese) and Kadu-papa
(Tamil).
Locality of the type, Mousakanda in Gammaduwa, 2,200 ft.,
€.P., Ceylon.
Distribution. — " Probably throughout the lower foot-hills
of the mountain cluster of the Central and TTva Provinces "
(Phillips), up to about 3,500 ft.
A little smaller on the average and with a fuller coat
than nordicus, resembling it in dorsal coloration, but with the
basal half of .the hairs of the ventral surface and of the cheeks
deep sooty-grey, and white-tipped hairs extend over the eye
to the median frontal stripe.
This description is taken from the type, a young adult $,
the only example known to me. It bears out Hill's description
of the race except in the dark tint of the hairs of the cheek,
which from his account appear to be white to the base in the
examples he recorded in 1933.
The approximate dimensions (in English inches) according
to Hill are : —
Average of 4 <J<J from Gammaduwa : head and body 8-3- :
foot If. "* °
Average of 4 $ $ from Gammaduwa : head and body 9— :
foot If.
The weights are from 5J to just over 7 oz.
Habits. — No doubt the Lorises of Ceylon resemble the south
Indian form in habits ; but the account given of the latter mav
be supplemented by the observation of Hill and Phillips on the
Ceylonese races. The period of gestation is unknown, but,
as Hill has pointed out, there is evidence that it is over three
months. Lactation lasts for about one year, and the young,
-usually one, but occasionally two, are for a long time, wholly
at first, partly later, dependent on the mother, even availing
themselves of her nursing when as much as half-grown and
able to feed independently. Females have been observed
suckling newly-born young and at the same time nursing the
one to which they had previously given birth.
LOB1S. 185
In captivity, according to Phillips, they show a special
liking for plantains and eat papaws and other fruits, but
seem to prefer insects or animal food of some kind. In the
wild state he thinks they are mainty carnivorous, feeding on
insects, small birds, lizards, and tree-frogs ; but captive
specimens would not touch small mammals, such as mice,
shrews, and bats.
Like Nycticebus they capture prey, after a stealthy approach,
with a lightning grab of both hands, and hold it in a tenacious
grip while devouring it. Everything is consumed — feathers,
scales, and bones of the vertebrates, and wings, legs, and the
horny exoskeleton of insects.
188 CAB.NIVORA
fore foot is also usually hairy between the wrist and the
plantar pad, with the outer carpal pad alone retained, and the
first digit, when present, is raised off the ground. In these
cases the plantar and digital pads alone rest on the ground
and the gait is digitigrade. The Cats, Hyaenas, and Dogs
show the extreme of this specialization of the feet. At the
opposite extreme are the Bears, in which the hind feet are
short and broad and mostly naked from the heel ; the five
toes form a continuous lightly curved line, and all are in
contact with the ground. They are called plantigrade
because, when walking, the entire sole of the foot appears to
be planted on the ground, although in reality both the heel
and wrist are raised to a certain extent. There is ever;'
gradation between these types, intermediate kinds of feet
being called subplantigrade.
The tail is usually long in species which run, jump or climb,
but it may be quite short, and act merely as a covering to the
underlying naked skin, as in bears. In otters it is very thick,
and acts as a swimming propeller.
The anal and genital regions also supply useful systematic
characters in the position of the prepuce, the length of the
perinseum, and the presence or absence of scent-glands
on it. There is also, except in the Bears, a pair of well-
developed glands within the anal orifice, and in connection with
these a pouch or glandular area may be present round or above
the anus.
In the skull, apart from the teeth, three sets of bones or
areas are important, namely, the turbinals in the nasal
passages, the auditory bulla, and the area in front of the
latter low down in the temporal fossa. The bulla may be
composed of a single bone, the tympanic, which primarily
forms the floor of the auditory orifice, but may be developed
into a tube or expanded into a thin- walled cavity ; or it
may be composed of two bones, the tympanic, which forms
its antero-external portion, and the entotympanic, forming
its postero-intemal portion ; where these two bones meet
a partition is developed dividing the cavity of the bulla,
into an outer and an inner chamber.
In the temporal fossa outside the foramen rotundum
there is frequently a bony channel, the alisphenoid canal,
through which a branch of the carotid artery runs. Its
incidence is remarkable, and it has been much used in classifi-
cation. It is probably a primitive character in the order
which has been retained in some families, lost in others (see
figs. 3&4, pp.4& 6).
In his classification of the Indian Cabnivora, Blanford fol-
lowed Flower in assigning them to the suborder Fissdpbdia,
which have normal feet, as opposed to the Potoipedia, in
CAENIVOBA.
189
which, the feet are converted into swimming paddles. The
Fissikedia, which alone are represented in the Indian fauna,
were divided into three main groups, the JUltjkoidea, com-
prising the Cats, Civets, Mongooses, and Hyaenas, the Cykoidea,
the Wolves, Jackals, and Foxes, and the Abctoidea, the
Bears, Raccoons, Weasels, Otters, etc. But quite obviously
from his key this classification is unsatisfactory ; and it
has been abandoned with the discovery that the Dog-family
is closelv akin to those classified under the Arctoidea, that
Pig. 55.
A. Partly diagrammatic longitudinal section of the nasal chambers
of a Cat {Felis).
B. The same of an Indian Jackal (Cams), illustrating the difference
between the iEmroid and Arctoid Carnivora in the arrangement
of the turbinal bones, cr, cribriform plate through which the
olfactory nerves enter the nasal chambers : an, anterior nares ;
pn, posterior nares ; nt, the naso -turbinal, which is a branch of et,
the ethmo-turbinal ; mt, the maxillo-turbinal.
the Hysenas do not differ from the rest of the JSujroidea, as
was supposed, in the structure of the bulla, and that the
turbinal bones, which had not then been studied in this con-
190 JELUROIDEA.
nection, supply a character by which the Carnivora may be
■divided into th.e two groups, ^Elttroidea * and Arctoidea.
The two suborders may be briefly diagnosed and contrasted
as follows : —
a. Ethmo-turbinals very large, occupying the greater
part of the nasal chambers and extending forwards
over and between the smaller maxillo-turbinals,
almost to the anterior orifice of the chambers.
The auditory bulla composed of two bones and
divided by a partition into two chambers. Cow- [p. 190.
per's glands absent ; at most a small penis bone . . iEluroidea,
a'. Ethmo-turbinals excluded from, the anterior orifice
of the nasal chambers by the enlarged maxillo-
turbinals. The auditory bulla composed of a
single bone, the tympanic ; its cavity typically
undivided ; when divided the partition is not
homologous to that of the ^luroid bulla.
Cowper's glands, associated with the generative
organs of the male, present ; a large penis bone in
all Asiatic species Arctoidea.
Suborder JELUBOIDEA.
Key to the Indian Families.
■a. Post-palatine foramina (see p. 5) set far back on the
maxillo-palatine suture ; teeth reduced in number
and highly sectorial, the dental formula being
3 1 3 or 2 X
i. g , c. y , pm. — s— , m. - ; interramal tuft of Pp. 191.
vibrissa absent " Felidae,
a'. Post-palatine foramina in front of the suture ;
teeth not so specialized or reduced in number, the
pm,. s or j , the m. g or y ; the interramal
tuft of vibrissas present.
6. Feet with four toes in front and behind, dog-like
in structure ; auditory bulla without oblique
groove ; jaws and teeth very massive Hysenidse.
b f . Feet with five toes, not dog-like ; auditory bulla
with distinct groove ; jaws and teeth less
massive.
c. Ears tolerably large, with well-developed
bursa and simple supratragus with no valvular
flap above it ; feet compact, with short claws ;
anus not enclosed in a glandular sac ; no bony [p. 330.
tube to the auditory orifice Viverridae,
c'. Ears small and rounded, with vestigial bursa,
a valvular supratragus, and a valvular flap
above it ; feet with freer digits and fossorial
claws ; anus in centre of glandular pouch ;
a well-developed bony tube to the auditory
orifice Herpestldse.
* This name, meaning cat-like, has been objected to, and Hbspes-
toidea, Mongoose-like, substituted on the grounds that the generic
name JEHurus or Ailurus is the admitted name for the Cat-Bear or
Panda, one of the Abctoedba. But for the sake of convenience I here
retain the long-established name MiASBxmrEA., which according to its
meaning is appropriate.
PELIDiE.
191
Family FELIDJE.
In their external characters the Felidse (Cats) come nearest
to the Viverridse, especially to the Linsangs, which have
the retractile claws and claw-sheaths typical of most cats,
the penis short and close to the scrotum, and no perineal
glands. But in the Felidse the plantar pads of the front and
"Fig. 56.
A. Left half of the palate and teeth of a Cat {J?elis Constantino ornata)
from Sehore, C.I. ppf, posterior palatine foramen on the maxillo-
palatine suture ; pm?, second premolar ; pm*, fourth premolar
(carnassial) ; to 1 , first and only molar.
B. Right half of dental portion of mandible. pm t , fourth premolar ;
9%, first molar (carnassial).
C. Outer side of right upper carnassial.
D. The same of right, lower carnassial.
hind feet form compact three-lobed cushions, there is a single
carpal pad on the fore foot, and the hind foot has only four
toes and no trace of metatarsal pads. The typical facial
vibrissa are well developed, with the exception of" the inter -
ramal tuft, which is absent. The ear is tolerably large, has
192 tBiiD^:.
a simple supratragal ridge, and a well-developed marginal
bursa. There are two or three pairs of mammae.
The skull has no alisphenoid canal, the jaws are shortish
and massive, and the auditory bulla is typically high and
rounded. In the mouth there is almost always a large space,
the postcanine space, due to the loss of the true first premolar
and the very small size or absence of the second in the upper
jaw, to the loss of the true first and second premolars in the
lower jaw, and to the uplift of the anterior portion of the
mandible carrying the canines and incisors. This space is
to give penetrating depth to the tusk-like canines. The
single upper larger premolar, morphologically the third,
which precedes the " carnassial " and the two below, mor-
phologically the third and fourth, have triangular compressed
crowns, with a single large pointed cusp and smaller cusps
behind and in front at its base. The crown of the upper
carnassial (#ra 4 ) is also compressed, its outer cutting portion
consisting of three cusps set in a line, the first low, the second
(paracone) high and pointed, the third or posterior (metacone)
lower and more blade-like ; the inner lobe (protocone) of the
tooth is set far forwards, small, and has at most a single cusp,
but in some cases it is reduced to vanishing point. The
single upper molar is a small, nearly functionless tooth with its
long axis set transversely. The lower carnassial, the single
lower molar, is very highly specialized. Its crown carries
two compressed blade-like pointed cusps, the paraconid and
protoconid, but there is no trace of the metaconid on the
inner side of the latter, and the " heel " of the tooth found in
most other Carnivores is at most represented by a small tubercle.
This tooth is always smaller than the upper carnassial.
In the classification of the Felidae a part of the skeleton is
used which, although primarily independent of the skull, has
become secondarily attached to it in the Mammalia. This is
the hyoidean apparatus (see fig. 57), of which the main portion
is a U-shaped bone, the hyoid, embracing the top of the wind-
pipe. The larynx, containing the vocal chords, opens between
the two arms , which are directed backwards . In most mammals
this bone is attached to the side of the back of the skull close
to the mastoid process on each side by a series of three bones
closely jointed together and arising from the forepart of the
U-shaped bone. This is the suspender, or suspensorium,
of the hyoid. Normally the suspensorium holds the larynx
and the back of the tongue close under the base of the skull,
admitting only comparatively slight movement of those
parts in a vertical plane. But in some cases the bones of the
suspensorium are wholly or partly replaced by an elastic
tendon or ligament which allows the larynx to be depressed
or elevated to a greater or less extent.
FELTD^.
193
In a paper, " The Classification of existing Felidte " (Ann.
Mag. Nat. Hist. (8) xx, p. 332, 1917), I made use of the
modifications of this structure in subdividing the family.
Other characters employed were supplied bv the skull and
feet.
Many of the zoologists of olden times could perceive that
the existing Felidse fall into a number of natural groups,
to which generic names were attached, sometimes with no
attempt to diagnose them. But the conception of the affinities
of the species was mostly hopelessly vague, and far too many
names were proposed either in a generic or subgenerie sense.
Fig. 57.
A. Left half of the hyoidean apparatus of a Tiger, showing the ligament
between the upper and lower bones, the ligament in this case very
rauch shortened, thickened, and twisted by methylated spirit.
B. The same of the common House-Cat, showing the complete chain of
bones found in all tho Felidsa except the Pantherinffi.
Admittedly the classification is difficult, and later authors
followed the easier course of assigning all the species but one
to the genus Felis, the only species to which separate generic
rank was given being the Hunting Leopard or Cheetah.
This was the classification adopted by all mammalogists,
vol. i. o
194 3WXDM.
including Blanford, until about twenty years ago. But,
in my opinion it embodies an erroneous conception of the
affiliation of the species, since the Cheetah appears to be
more nearly akin to the ordinary Cats than the latter are to
Lions, Tigers, and their allies, which have a highly modified
hyoidean apparatus, the Cheetah resembling the ordinary
Cats in the normal structure of this system of bones.
The Cheetah, called by Blanford Oynselurus, but now known
as Acinonyx, was, apart from its general build, distinguished
by its so-called non-retractile claws. The main distinctive
character of its foot lies in the complete loss of all trace of
the lobes of skin at the tips of the toes, which in other Cats
to a varying degree constitute sheaths for the claws, protecting
their tips when retracted. In the most perfectly developed
foot in the family there is a pair of these skin-lobes to the
claws of the four main digits, and as a rule in this case the
skin or " web " between the digits is continued up to the
digital pads. But in several of the ordinary cats some of the
lobes may be undeveloped, the others reduced in size and the
webs much less extensive, so that the foot structurally
approaches that of the Cheetah.
The Cats, which are the most highly organized of all the
Caenivoea, are found almost all over the world up to the
limit of tree-growth, except in the Australian Region and
Madagascar. They are essentially predatory, feeding mostly
on freshly-killed mammals and birds ; but many of them
will eat carrion and kill and eat reptiles, frogs, and fishes, and
occasionally insects.
Many are essentially forest species ; others inhabit open
grassland or semi-desert rocky districts, wherever, indeed,
prey is available.
I divide the Felidse into the following subfamilies : —
a. Hyoidean apparatus modified by the conversion
of the median part of the suspender into a long
elastic tendon ; paws with complete claw- [p. 195.
sheaths Pantherinae,
a'. Hyoidean apparatus of the normal mammalian
type, the suspender consisting of a chain of bones
jointed end to end.
6. Paws with more or less well-developed cutaneous
lobes constituting sheaths for the claws when
retracted Felinse, p. 243.
&'. Paws without trace of cutaneous lobes con- [p. 322.
stituting sheaths for the claws Acinonychinse,
PANTHEBINJE.
195
Subfamily PANTHEMME.
The suspender (suspensorium) of the hyoid modified from
the normal mammalian type by the replacement of the median
bone of the chain by a long, elastic tendon conferring great-
mobility upon the larynx and permitting the back of the
mouth to be greatly distended. The tips of the digits of both
fore and hind feet have a cutaneous lobe on the outer and
inner aide of the claws so that the claws, when retracted, are
A, Lower side of right, fore paw of a Leopard with, the toes spread, she
claws extruded, and the hair clipped close to show the pads, the
interdigital webs, and the lobes of skin which act as sheaths to
the claws when retracted.
B. Lower side of right hind paw of the same.
o2
196 EELIDiE.
completely sheathed and the interdigital web extends to the
digital pads.
The structure of the hyoid in this subfamily is associated
with the voice. The sexual call, where known, i. e., in the
Lion, Tiger, Leopard, and Jaguar, is a definite roar — it is not
recorded in the Snow-Leopard — and pleasure or content is
not expressed by " purring " as in the rest of the family.
Key to the Genera of Pantherinse based on Cranial
Characters.
a. The dorsal profile of the muzzle between the
anterior nares and the interorbital area tolerably
straight, not noticeably concave ; the frontal
interorbital area not abruptly elevated ; outer
chamber of the bulla small, the partition close [Oken, p. 196.
to the orifice Pantheka
a'. The dorsal profile of the muzzle decidedly concave
where it passes into the frontal interorbital area,
which is noticeably elevated ; outer chamber of
the bulla larger, the partition remote from, the [p. 239.
orifice TJncia Gray,
Genus PANTHERA Oken.
Panthera, Oken, Lehrb. Zool. 2nd Abth. p. 1052, 1816.
Tigris, id., torn. cit. p. 1066.
Leo, id., torn. cit. p. 1070.
Panthera, Pocock, Ann. Mag. Nat. Hist. (8) xviii, p. 306, 1916.
Type of Panthera, pardus Linn. ; of Tigris, tigris Linn. ;
of Leo, leo Linn.*.
Distribution. — The greater part of temperate and tropical
Asia from the Caucasus and Mongolia to Java and Bali,
and most of Africa except the Sahara ; tropical America.
Dorsal profile of the skull flattish or tolerably evenly
convex, not concave where the muzzle passes into the frontal
interorbital area, which is not noticeably elevated, and the
area behind the elevation less steeply sloped : the basicranial
axis nearly horizontal ; a wide notch between the glenoid
process and the bulla ; bulla with the inner chamber large,
* By some modern authors the three well-known species here assigned
to Panthera are regarded as representing distinct genera under the
names cited. But in the three papers, quoted below, on Tigers and on
the Panthers and Lions of Asia, and published in the Journ. Bomb.
Nat. Hist. Soe. 1929-30, I have shown that the cranial and dental
characters relied on to distinguish them, generically completely break
down when a large series of skulls is examined. It is needless to repeat
the facts. The prevalent external differences between the three
species, when adult, are well known, but are, in my opinion, only of
specific value. Their citation under different generic names completely
obscures their kinship, which is undeniably close.
LU
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Z
z
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PANTHERA. 197
the outer small, the partition between, them close to the
external auditory meatus ; mandible with a sloping, eonvexly
rounded chin.
Key to the British Indian Species of Panthera based on
External Characters.
a. Partem of stripes or spots or both combined,
usually present in the cubs, lost or very nearly so
in the adult, in which the general colour varies
from tawny-brown to greyish ; a black tuft on [p. 210.
the tip of the tail ; male usually with mane leo (Linn.),
a'. Pattern of stripes or spots retained throughout
life ; no black tuft on tail-tip ; sexes alike except
m size.
b. Pattern consisting of vertical black stripes, LP- 19"-
with at most a fe%v faint spots ; a ruff on the tigrix (Linn.),
cheeks as in leo ; size larger
h '. Pattern consisting mainly of irregularly arranged
rosettes composed of from three to fire spots [p. 222.
in a ring ; no ruff on cheeks ; size smaller . . . pardiin (Linn.),
Key to the Species based on Normal Differences in the Skulls.
a. Skull large, in aduit ^J up to about 13 or 15 in. long.
6. Nasals projecting beyond maxillse, frontal area
elevated, facial portion shorter as compared
with cranial and less massive, inferior edge of [p. 190.
lower jaw not convex in the middle tigrls (Linn.),
b'. Nasals not projecting beyond maxillse, frontal
area flatfish, facial portion longer as compared
with cranial and more massive, inferior edge [p. 212.
of lower jaw lightly convex in the middle .... leo (Linn.),
a ' Skull smaller, in adult <J<J up to about 9 in. , typically
in shape and proportions more like that of tigris, [p. 225.
but nasals not projecting so far beyond maxilla? . . pardus (Linn.),
In addition to the three species above mentioned, Panthera
includes the Jaguar (Panthera onca) of America, which is most
nearly related to the Leopard (P. pardus), resembling it in
the nature of its pattern and sexual call, but differing in its
larger size, on the average, relatively bigger head, and con-
siderably shorter tail.
21. Panthera tigris (Lmnseus). The Tiger.
Felis tigrU, Linn., Syst. Nat. ed. 10. p. 41, 1738 ; and of most
subsequent writers.
Panthera tigrti, Pocock, Ann. Mag. Nat. Hist. (S) xviii, p. 306,
1916 ; id., Journ. Bomb. Nat. Hist. Soc. xxxiii, p. 505, 1929.
Typical locality (traditionally fixed), Bengal.
Distribution. — Practically the whole of Asia, in suitable
localities, eastward of a line from the Caucasian area and
Afghanistan to Manchuria and Korea, but omitting the
198 FELIDiE.
Tibetan Plateau, most of India, and extending from Btjhma
•and the Malay Peninsula through the Sunda Islands to Bali,
east of Java, but not found in Ceylon.
The largest of the existing species of Felidse, apart from the
Lion, which it rivals in size, and distinguished from the other
species of Pantherinse by the sum of a number of characters,
more particularly by the highly specialized pattern of numerous,
nearly vertical dark stripes extending from the spine over
the flanks to the belly, generally broken up into smaller,
simple or looped stripes, and always, in normal skins, con-
spicuous against the bright typically reddish or yellowish
tan colour of the upper side and the pale, typically white
hue of the underside. A conspicuous pale, typically white patch
over the eye is also distinctive, and a peculiarity of the
reddish hair of the interspaces is their tolerably uniform hue,
without trace of black speckling, all the black in the pelage
being concentrated in the stripes.
The head is marked with short transverse stripes on the
crown, with longitudinal vibrissal stripes on the white upper lip,
and an irregular pattern — some longitudinal, some vertical —
on the cheeks, which are white in their lower half like the chin
and throat, a long vertical stripe descending in front of the
ear ; the ears are black at the back, with a white patch in the
upper half ; on the nape the stripes are few, thin, and obliquely
longitudinal. On the spine the stripes of the two sides
typically meet and form forwardly jutting angles. The fore
leg externally is generally without stripes, and a considerable
area of the shoulder above it is not infrequently unstriped.
On the hind quarters the stripes are narrower and closer set,
and low down on the thigh they pass into irregular transverse
stripes extending to the hock. The inside of the fore and
iind limbs is white, with a few stripes. The tail is heavily
but irregularly banded, typically loses its bright colour in
the distal half above, and has a black tip, but no tuft.
From sportsmen's records it seems that the tail is, on the
average, less than half the length of the head and body ; in
some cases it is considerably less. On the other hand it is
sometimes considerably more, nearly as long in proportion
as in typical leopards (see Sterndale's records, p. 205).
In both sexes there is always a fringe of hair, varying in
length, which starts just above the ear in front and descends
to the throat, as in the lion ; but, unlike the latter, the hair
on the top of the neck over a wide area grows backwards in
the same direction as that of the head and the spine, only on
the sides of the neck does it usually grow forwards from
a whorl in front of the shoulder and, passing beneath the ear,
meets the backwardly-directed hairs of the cheek at the
fringe ; but the direction here is variable, and may be affected
PANTHERA. 199
by additional whorls . On the nape the hair is always apparently
longer than on the body, and may be long enough to constitute
a small mane.
The skull (see fig. 59) is distinguished by its large size, only
rivalled in the Felidse by that of the lion, from which it is
not always easy to distinguish when a large series of both is
examined. The tiger's skull, however, is, on the average, at
least shorter in its facial and longer in its cranial portions,
measured from the pcstorbital process ; it is usually more
arched, owing to the brow being higher and the sagittal crest
lower ; its nasals are longer, overlap the maxillae behind to
a greater extent, and the anterior nares are narrower ; the
lower edge of the mandible is not convex in the middle, but
is concave in front of the angular process, so that the skull
rests steadily on a flat surface and does not " rock " backwards
and forwards ; finally, the inner lobe of the iipper carnassial
{pnfi) is a little larger.
21 a. Panthera tigris tigris (Linnseus). The Indian Tiger.
JPelis tigris, Linn., Syst. Nat. ed. 10, p. 41, 17.58 ; and of all authors
on Zoology and sport in India and Burma.
Panthera tigris tigris, Pocoek, Journ. Bomb. Nat. Hist. Soc.
xxxiii, p. 505, 1929.
Vernacular. — Bdgh, Sher (female Bdghni Shernii) (H.) ;
Ndhar, Sela-vdgh (H. of Central India) ; Babr (P.) ; Mazar
(Baluchi) ; Shinh (Sindhi) ; Padar-suh (Kashmiri) : Patayat-
bdgh, Wahdg (Mahr.) ; Go-vdgh (Beng.) ; Tut, Sad (Hill tribes
of Rajmehal) ; GartimMla (Kol.) ; Ldkhra (Uraon) ; Krodi
(Kondh) ; Kula (Sonthal, Ho, and Korku) ; PMi (Tam.,
Tel., Mai., and Gond) ; Puli-redda-pffli, Peram-pilli (Tam.) ;
Pedda-puli (Tel.) ; Perain-pMi, Ktidua (Mai.) ; Kuli (Can.) ;
Xdri (Kurg) ; Pirri, Bilrsh (Toda) ; Tag (Tibetan) : TMt or
Tiik (Bhot.) ; Sathong (Lepeha) ; Kek-va (Limbu) : »S*c/n
(Aka) ; Matsd (Garo) ; Kla (Khasi) ; 8a, Bagdi, Tekhu,
Khudi (Naga) ; Humpi (Kuki) ; Sumyo (Abor.) ; Eil, Kliamti) ;
Sirong (Singpho) ; Kei (Manipuri) ; Misi (Kaehari) ; Kya
(Burmese) ; Kla (Talain) ; KM, Botha-o, Tupuli (Karen) ;
Htso (Shan) : Rimau, Harimau (Malay).
Locality of the type, Bengal.
Distribution. — Practically throughout India from the
Himalayas, where it may ascend to 6,000 or 7,000 ft., south-
wards, but not in the deserts of Rajputana, the Punjab,
Cutch, and Sind, and exterminated or very scarce in certain
districts where it was formerly plentiful. Also throughout
Bubma in suitable localities *.
* The Burmese tiger is provisionally identified as P. t. tigris. There
is some evidence that, it may prove to be a distinguishable race ; but
I have not seen sufficient skins and skulls to establish this point.
200
FEUDiE.
One of the largest of the races, only a little, if at all, smaller
on the average than the Mongolian and Manchurian race
(P. t. longipilis), but distinguished from it and from the Chinese
race (styani) and the Persian or Caucasian race (P. t. septentrio-
Tialis) by its shorter and thinner winter coat, and from the
last also by its fewer, more widely spaced stripes. Con-
siderably larger than the three races from the Sunda Islands,
Sumatra (P. t. sumatree), Java (P. t. sondaica), and Bali
(P. t. balica), and also less fully and closely striped.
A good many of the skins of British Indian tigers presented
to the British Museum are more or less faded from being
exhibited or exposed as rugs, but several are unfaded and
exhibit the characteristic bright orange-red hue, and are
practically indistinguishable in colour, although coming
from the following -widely separated localities : — United
Fig. .->9.-
-Skull of Tiger from the Central Provinces, showing the shape
characteristic of the Indian race.
Provinces (Major G. Burrard), Bengal (Col. Sanderson),
Pennghot, Mirzapur (S. Wyndham), Chanda, C.P. (C. E.
Hewetson), Kadra, N. Kanara (N. B. Kinnear), Travancore
(the Conservator of Forests), and the Thaungyin Valley,
Amherst, Tenasserim (E. H. Peacock). The last does not
bear out the supposition that Burmese tigers are richer in
colour than Indian tigers. No two skins are alike in pattern,
the stripes varying individually in thickness, looping, and the
extent to which they are broken up.
Although only two of the skins are dated, many of the others
exhibit interesting differences in the coat. Of the dated skins
a ? from Mirzapore (November) has the coat roughish, but
with hardly any wool and about 18 mm. long on the back,
25 mm. on the nape, and from 45 to 50 mm. on the cheek.
The other, a g from Central India (H.H. The Maharaj Holkar)
(February), has a little wool, a roughish, slightly longer
PAHTHEBA. 201
coat, about 21 mm. on the back, 31 mm. on the nape, and
from 50 to 70 mm. on the cheek. These skins, of different
sexes, are very much alike, allowing for the increase in the
coat from November to February. Neither has what can be
called a mane on the nape. But in two, undated, from the
Nepal Tarai, a tj (Col. R. L. Kennion), with the coat on the
body without wool and only about 12 mm. long, has a distinct
mane, 66 mm. (2| in.) on the nape, shortening to 33 mm.
towards the shoulders, and the cheek-fringe up to 85 mm.
or more (3 J in.). A ? (Sir R. Dane), with a similar short,
clearly summer coat, has the nape-hairs only 25 mm. (1 in.)
and the cheek-fringe 50 (2 in.). Since both these skins are
in summer coat the difference between them in the length of
the hair on the nape and cheek might reasonably be regarded
as sexual, and comparable in a small way to the usual sexual
difference in lions. But this is not borne out by three undated
skins from the Central Provinces (Sir R. Dane). A <J from
Raipur has the coat smooth, without wool, and. quite short,
about 8 mm. only on the back, 17 Trim., not constituting a
mane, on the nape, and about 50 mm. on the cheek. A $ from
Balaghat, on the contrary, has the coat about 18 mm., with
a little wool, on the back, a mane of 56 mm. on the nape and
the cheek-fringe about 62 mm. In this case the differences
are apparently seasonal. The third skin, a <J from Baslar,
has the coat about 12 mm., without wool, the nape-hairs
about 25 mm., and the cheek fringe 50 mm. Unless, as is
possible, there was a mistake in the labelling when the skins
were dressed, it is the $ that has the mane in this ease. The
skulls show the sexes.
Other undated skins from scattered localities show similar
variations indicating seasonal change. A <J from Bengal
(Col. Sanderson) has a sleek coat, with no wool, only about
8 mm. long on the back and 14 mm. on the nape ; whereas
another from Danta Mahi Kanta, Gujerat (Major G. H.
O'Donnell) has a rougher coat, with a little wool, about
25 mm. long, and on the nape about 40 mm. The hair on the
belly also varies greatly, being typically, but not always, longer
than on the body. According to Dunbar Brander tigers in
the Central Provinces rapidly moult the winter coat in March,
there being a great difference between skins at the end of
January and the end of March.
Colour Varieties of Indian Tigers.
Apart from comparatively slight individual differences in
depth of hue, Indian tigers sometimes exhibit partial or
complete albinism. In most of the cases of so-called " white "
tigers the stripes are dark brown or reddish-black, and stand
202 FEUD-ffi.
out boldly against the white ground-colour. This type has
been recorded from Orissa, BUaspur, Sohagpur, and Rewa.
One from Mirzapur (Mrs. Craigie Halkett) is similar, but
has the stripes tan. A wholly white tiger, with the pattern
only visible under reflected light, like the pattern of a white
tabby cat, was exhibited in the Exeter Change Menagerie
in the early part of the nineteenth century and described by
Hamilton- Smith. It was probably an Indian specimen.
Probably similar to this, although the stripes werenot mentioned,
were a couple of young tigers with pink eyes and apparently
" pure albinos," recorded from Cooch Behar in 1922 by
V. N. Narayan*.
Black tigers also have been reported, but no skins have come
to hand, and some of them have turned out to be " black
panthers." One that was sighted near Bhamo, but not
secured, is said to have left " pug-marks " too big for a leopard's !
This was recorded by Mr. Hauxwell (Journ. Bomb. Nat. Hist.
Soc. xxii, p. 788). The story of another that was shot near
Chittagong in 1846, and seen in a state of putrefaction in the
jungle by Mr. C. T. Buckland, is told in ' The Field,' 1889,
p. 73, and a dead one in a similar condition, was said to have
been found in the Lushai Hills (' The Field,' 1928, p. 656) . The
evidence, however, is clearly incomplete. There is no known
reason why tigers should not be black, like leopards and
jaguars ; but at present their existence can be neither asserted
nor denied.
Skulls of British Indian Tigers.
The skulls of British Indian tigers differ from those of the
races occurring in the Sunda Islands in their larger size
and in some other features which need not be particularized :
but there is not sufficient material available of the Chinese,
Mongolian, and Persian races to establish distinctive characters
between the four. In the following table, containing the
measurements of a selection of the specimens in the British
Museum to show the variations in the size, the Indian and
Burmese skulls are treated separately. Although the <J skulls
from the two countries overlap in size, the Burmese are, on
the average, smaller, but there are too few of them to warrant
the conclusion that they represent another race, especially
as the differences between the $ skulls is less marked. There
is no structural difference between the skulls from the two
countries.
* A variety of a different kind, with, the ground-colour normal above
and below, but the pattern brown, and consequently little defined, is
represented by the skin of the south-western Asiatic race which was
procured on the northern slopes of Mt. Elburz by Col. E. L. Kennion.
This was described and illustrated in colour in my paper, 1929.
PASTHERA.
203
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204 FELTD.®.
The Indian tiger skulls vary considerably in length and
zygomatic width, and there is some evidence that the longest
are mainly northern skulls. For instance, in addition to those
enlisted above, one from Bengal (Sanderson) has the condylo-
basal 332 mm., in those from Assam (Marcus Maxwell and
the Maharajah of Gauripur) it varies from 322 to 309 mm.,
the average being 317i mm. ; in one from the Bhutan Tarai
it is 321 mm. ; and in five Nepal skulls it varies from 317 to
305 mm., the average being 309 mm. Even this last average
exceeds by 5 mm. the average of six skulls from the Central
Provinces, but the intergradation is complete. On the other
hand, the zygomatic width in three of the skulls from the
Central Provinces, namely, Osmaston's and those from Jubbul-
pore (Carlisle) and the Bastar State, is actually or relatively
greater than in the more northern skulls. The conclusion
above suggested regarding the superiority in length of northern
skulls seems to be borne out by the measurements in Rowland
Ward's ' Records,' 1928, where the first ten skulls entered in
order of total length came from Kumaun, Nepal, Cooch
Behar, the Duars, Assam, and Bengal, five of them slightly
exceeding my skull from Darjeeling, which is 15 in., the
largest measuring 16 in. and coming from Naini Tal (B. B.
Osmaston). Apart from size, the most noticeable variation
is in the development of the sagittal crest, which profoundly
affects the shape of the dorsal profile of the cranial portion.
Usually it is a thick, three-fold, ridge and quite low, from
about 5 to 10 mm. high in the middle of the crown, but raised
and deep posteriorly, where it runs into the uptilted occipital
crest. In this case the dorsal profile is markedly concave
in front of the occipital crest ; but sometimes the sagittal
crest, e.g., in one of the <J skulls from Gauripur, is as much
as 22 mm. in the centre of the crown, and the dorsal profile
behind the orbits is straight from front to back as in a lion's
skull.
Size of British Indian Tigers.
There has been some difference of opinion about the size
of tigers. Large numbers of both sexes have been measured
by sportsmen ; but in most cases only the total length from
" tip to tip " has been recorded. There are two methods,
caEed respectively " over the curves and " between pegs,"
and these naturally yield discrepant results. Col. C. S.
Stockley, for example, showed that his " large tiger " from
Kheri, entered in the table below, according to its dimensions
" between pegs," was 6 in. longer when the tape was run
over the curves of the head and body. Since the methods
are not always stated the results are unsatisfactory. The skin.
PANTHEKA. 205
from Mergui is the only one in the British Museum in the
series quoted below in which the tail was measured as well as
the head and body. The first seven on the table were taken
between pegs, and probably those cited from Sterndale were
similarly measured, but the evidence that all were full-sized is
inconclusive. That is only certain in the Mergui specimen,
of which I have the skull.
Locality, authority, and sex.
Cen.tr. Prov. (Dunbar Brander) ; <J.
Kheri (Col. C. S. Stoekley) ; <J
'" India " (Col. Stewart Capper) ; <J
"India" (Col. Stewart Capper) ; <$
'" India " (Col. Stewart Capper) ; <J
Bankaohon (G. C. Shortridge) ; (J .
Mergui (G. E. R. Cooper) ; ad. $ . .
•'India " (Sterndale) ; $
"India " (Sterndale) ; ?
Head and
body.
Tail.
Total.
ft. in.
ft.
in.
ft. in.
7 3
2
8
9 11
6 7
2
11
9 6
6 4
2
6
8 10
o 10
5
8 10
5 S
3
8 S
6
3
1
9 1
."> 9
2
7
8 4
5 3£
•2
11
8 U
5 2
3
2
8 4
This table brings out very clearly the great variation in
the proportion of the tail to the head and body.
As regards total lengths " between pegs," Dunbar Brander's
longest cj was 10 ft. 3 in., his shortest 8 ft. 9 in., the average
of many being 9 ft. 3 in. His longest and shortest $ specimens
were 9 ft. 1 in. and 7 ft. 10 in. respectively, and the average
of thirty-nine 8 ft. 4 in. But out of a large number measured
" between pegs " by Gen. R. G. Burton the longest tiger was
9 ft. 8 in. and the longest tigress 8 ft. 6 in. and these were
the only two that reached these dimensions.
Tigers from Burma have been alleged to be smaller on the
average than those from India ; but Peacock gives reasons
for doubting this. I have records of the totallength " between
pegs " of two tigers and two tigresses from that country. The
tiger from Bankaehon, Tenasserim (Shortridge), recorded as
9 ft. 1 in., and said to be considered large for the district,
bears out H. C. Smith's statement that 9 ft. is a good length
for a Burmese tiger But there is no skull to show that the
animal was full-sized. The second tiger, from Pinnwe, Katha
District, near Bhamo (V. H. T. Fields Clarke), known to be
full-sized from its skull, was 8 ft. 4 in., a short beast, but the
skull is nearly 1 in. shorter than an adult <J skull from the
Upper Chindwin (Mrs. Manby). Of the tigresses, both
fully grown by their skulls, one from Mingun, near Bhamo
(Fields Clarke), was 7 ft. 8 in., the other, from Mergui (Cooper),
being 8 ft. 4 in. The average of these two, 8 ft., is a good
length for a Burmese tigress according to H. C. Smith (' Wild
Animals of Burma,' p. 2, 1935) ; but Peacock quotes many
instances of tigers ranging from 9J to over 10 ft. {' A Game
Book for Burma,' p. 160, 1933).
206 FELIDJE.
Compared with the specimens of Indian tigers entered in
Rowland Ward s ' Records ' for 1928, all the above-quoted
specimens are small. Fifty-six Indian tigers in the flesh
ranged from 9 ft. 10 in. to 10 ft. 7 in., and twelve tigresses
from 9 ft. to 9 ft. 10 in. The excess is probably due in a
measure to the animals being especially selected for measure-
ment on account of their size ; but probably most of them
were measured " over the curves."
The standing height at the withers of a living tiger seldom
exceeds 3 ft. I measured many standing against or walking
past graduated scales in the Zoological Gardens, and the biggest,
a ^ of the northern Asiatic race, shipped from Vladivostock
and admitted by all Indian sportsmen who saw him to be
a huge beast, stood 3 ft. 2 in.*
Weights. — Dunbar Brander's heaviest tiger was 512 lb.,
the lightest 353 lb., the average of many being 420 lb. His
average for tigresses was 290 lb., the heaviest being 343 lb.
Sir J. Hewett's heaviest tiger and tigress were respectively
570 and 347 lb. But there are three tigers in Ward's ' Records '
scaling 600 lb. from Cooch Behar, 608 lb. from Gwalior,
and 645 lb. from Kumaun. Shortridge's Bankachon tiger was
382 lb.
The Original Country of the Tigers.
Prom the discovery of fossil remains of tigers in the
extreme north of Siberia in the Pleistocene and the survival
of the species in Manchuria and Amurland, it is inferred that
the species is of northern origin and migrated southwards
to south-western Asia on one side of the Tibetan Plateau
and through China on the other, and thence to Burma and
ultimately to the Sunda Islands. The evidence that it
made its way into India from Burma round the eastern end
of the Himalayas, and not through Afghanistan and Persia,
is supported by one or two facts. There is complete con-
tinuity in distribution and racial characters between the
tigers of India and China and discontinuity in both respects
between those of western India and Afghanistan. In the
Himalayas they are rarer west of the Bhagirathi River than
to the east of it (Burrard), and are not found at all events in
Upper Kashmir, nor are they plentiful in Upper Sind or the
* A dimension interesting to sportsmen is what Bowland Ward calls
" The estimated height at the shoulder." This is taken " between pegs "
when the dead animal is lying on its side on the ground. In the 1928
edition of the ' Eecords ' this dimension ranges in tigers from. 3 ft. 3 in.
to 4 ft. ; but it is not the standing height of the living animal. One
living tiger, however, is said to have stood 3 ft. 10J in. at the shoulder.
If this be accepted, the animal must have exceeded in all its dimensions
the big N. Asiatic specimen above referred to by nearly as much as
an ordinary tiger exceeds a leopard.
PAITTHEEA. 207
Punjab. That the species was a comparatively late comer
into India is attested by its failure to reach Ceylon. (See also
under the account of the Lion, p. 221).
Habits of Tigers.
Tigers live in forests, jungles, and even in scrub or grass-
land, wherever the three essentials for their well-being are
supplied — plenty of game, water, and shelter from the sun.
Being intolerant of heat, they mostly lie-up during the day,
start hunting about sunset, and may be so occupied until
sunrise, often covering great distances in their quest, which is
conducted at a slow, stealthy walk, usually along a jungle
track or nullah if available. The presence of game is detected,
it is said, not so much by scent* as by hearing and sight,
both of which senses are said to be remarkably keen. When
it is perceived, the tiger's approach becomes a silent, gliding
crawl f until within striking distance.
They prey upon game of all kinds — elephant (even adult
cows), bison, buffalo, deer, nilghaie, wild pig, bear, and
porcupines, as well as tame cattle and goats and ponies,
especially if wild game is scarce. They will also eat their
own kind and leopards. Nothing, indeed, seems to come
amiss. In flood time they have been known to devour
crocodiles, water-tortoises, and fish. A hungry tiger will
take frogs, and there is a record of the stomach of one being
crammed with locusts. The droppings of a tiger in the Naga
Hills were full of salt-lick earth, as noted by R. C. Morris.
A great deal has been written about the tiger's method
of seizing and killing its prey. A favourite practice is seizure
by the throat or nape, followed, if possible, by dislocation of
the neck by a wrenching twist. Another, more particularly
adopted in the case of big or powerful beasts like bison and
buffalo, is " hamstringing," which renders the victim incapable
of putting up a troublesome fight. But often to secure the
quarry a leap on the hind quarters has to be resorted to,
and a stroke with the paw is sufficient in some cases. Some-
times tigers hunt in couples, one driving a deer towards the
* According to Dunbar Brander the sense of smell is very poor.
Peacock, however, doubts this, and cites the case of a tiger in Burma
sniffing his foot-tracks although nearly an hour old. Some of the smaller
Cats, with nasal passages no larger relatively than in a tiger, have,
as I know, a keen sense of smell.
t My nephew, Nicholas Pocock, E.C.S., who watched a tigress
stalking a herd of Sambar along a track strewn with dead leaves,
told me she was almost invisible owing to her coloration, and her move-
ments practically imperceptible. Only when he looked from her to
the deer and back again did he notice that she was nearer each time
to her quarry, which did not perceive her until too late.
208 FELma;.
place where the other lurks in wait, and sometimes the hunt
is conducted by a family party.
As a general rule, if not invariably, the tiger starts feeding
on the buttocks of its kill and works forwards, continuing
his meal, which may amount to as much as " three-quarters
of a fair-sized buffalo," until gorged. At sunrise the remainder,
if any, may be hidden from vultures either beneath a bush or
under mouthfuls of grass torn up by the tiger for the purpose,
as seen by Dunbar Brander. After drinking, the tiger sleeps
till sunset, then returns to finish the carcase. But the animal
is not particular to what he kills himself or to the freshness
of the meat. If he comes across a carcase in the jungle he
will eat it even in an advanced state of putrefaction, maggots
and all.
For some inexplicable reason, tigers as a general rule,
it seems, refrain from killing man, as if they had an instinctive
fear of him. But if once the fear is overcome and the facility
with which men and women can be killed is found out, tigers
will thereafter take them on every possible occasion, and may
become a scourge to a district by turning into " man-eaters."
The first attack on a man may be due to any cause compelling
irresistible hunger, such as great scarcity of other game or
some physical defect, like old age, which makes its capture
unusually difficult. Or it may be induced by the ferocity
of a tiger when pairing*. At this time they lose all fear of
man and are particularly dangerous, since they attack at sight
if he happens to be near. Although a man so attacked would
not be killed for prey in the first instance, he would probably
be eaten by the tiger or tigress and the appetite for him excited.
Of their physical prowess there are many records. Half
lifting, half dragging, they can transport a bulky carcase
many times their own weight. A tigress was seen by Dunbar
Brander to drag without apparent effort a half-grown buffalo
up a steep river-bank covered with alluvial soil so soft that
she sank at every step. In Burma, Peacock tells us, a tiger
shifted for 15 yds. the carcase of a bison which thirteen
strongish men could not drag, and he saw another swim
over a river with a medium-sized cow in its mouth. They
are, indeed, strong swimmers, and readily take to water. In
the Sundarbans, according to Hickie, they have been known
to cross tidal rivers some 4 miles wide and running with
* This characteristic was recorded by Dunbar Brander. Many
years ago I noticed the same thing in lions at the Zoological Gardens.
They became savage to a degree when pairing, and would charge the
bars of the cage, growling fiercely at a keeper with whom on other
occasions they were on the most friendly footing. Resentment of the
possibility of interference with the business in hand seemed to be the
•explanation of the change in temper.
PANTHERA. 209*
a tolerably swift current. Their liking for water is further
evinced by their habit sometimes of lying immersed up to the
head in a pond in hot weather. Dunbar Brander saw a tiger
clear a 6 ft. wall " like a greyhound " and a tigress take a gully
19 ft. wide in her stride without checking speed. Having
very little to fear in the jungle, they have less occasion to-
exercise their climbing powers than the smaller species of
Cats ; but, like them, they can climb, and have been known
to do so to fetch men out of trees and to escape when hunted.
Of many records collected by Gen. It. G. Burton one may be
quoted as evidence that they climb as well as ordinary cats.
A tigress enclosed in a shola near Ootacamund ascended a tree-
trunk about 1 ft. in diameter and rising vertically without
a branch for 2o ft. Here she was shot and fell heavily to th&
ground, but, recovering, went again up the tree, despite her
wounds. Tigers are also " treed " sometimes when bayed
by a pack of wild dogs, incidentally one of the few denizens
of the jungle with which they do not know how to deal.
Tigers utter a variety of sounds expressive of different
emotions. The most impressive is a deep-throated, loud,
far-carrying roar, unmistakably like a lion's. This may be
repeated at short intervals and is used as a mate- call*.
There is also the peculiar note very puzzlingly rendered as-
" pook," " moop " or " titting," which has been likened to
the call of a Sambhar, and believed by some sportsmen to be
uttered as a lure to attract the deer. That is probably pure
fancy ; but it is not known what the sound expresses or what
purpose it serves. Both tigers and tigresses when approaching
one another, or even human beings, in a friendly spirit utter
a gentle puffing sound by expelling air in rapidly repeated
jets through the nostrils. When suddenly surprised and
startled they give vent to a loud " whoof." As a rule they
attack prey in silence ; but commonly when charging a man
they utter a kind of barking cough two or three times, which
may rise to a harsh roar. Anger is expressed by a menacing
growl or by wind-like expulsion of air with the mouth open
and the fangs bared.
At pairing time the sexes keep together, a tiger being
sometimes accompanied by two tigresses. The period of
gestation is usually said to be fifteen weeks (105 days), but
Dr. Vevers informs me that in the case of a litter born at
Whipsnade in 1937 it was sixteen weeks (112 days). Before
* la the Zoological Gardens I have heard a tiger aad tigress answering
one another at night from cages some half a mile apart. Inexperienced
people often mistook a tiger's roar for a lion's, and if a tiger started to
roar in the Lion's House the lions would take up the chorus, as is their
eustom when one of their own kind begins.
VOL. I. P
210 FE1XDJB.
the birth of the cubs, usually two or three but occasionally
as many as six in number, the parents separate and the sole
duty of looking after them devolves on the tigress. Cubs
may be bom anywhere in the jungle, but usually an overhanging
rock or a cave is selected as a shelter. About two months
after birth they begin to venture out of the lair and, when
big enough, they accompany the tigress on her hunting, and
may stay with her until about two years old, even after she
has paired again and has another litter on the way. Sometimes
she tolerates their company even longer, and may be seen
with young of different ages. There is some doubt as to the
age at which tigers can be said to be full-grown. Blanford,
on unstated authority, says three years ; Dunbar Brander
says five, adding that they put on muscle even after that ;
but the ability of a tigress to breed long before she is full-
sized was shown at Whipsnade, where one produced a litter
when she was only two years old, as I learn from Dr. Vevers.
Regarding the age to which tigers may live, Dunbar Brander
quotes a case of one well known in a particular jungle as an
adult for fifteen years. Since it was in prime condition
when killed, at the computed age of twenty, it may be inferred
that the animal would not have been far short of thirty if
it had died as the result of old age. This view is supported
by a case recorded by Sanderson, and cited by Blanford,
of a tiger known as a cattle-eater for twenty years which,
when killed, showed no definite signs of decrepitude.
22. Panthera leo (Linnaeus). The Lion.
Felis leo, Lion.., Syst. Nat. ed. 10, p. 41, 1758 ; and of most subse-
quent ■writers.
Pantkera leo, Pooocfc, Ann. Mag. Nat. Hist. (8) xviii, p. 306, 1916 ;
id., Journ. Bomb. Nat. Hist. Soc. xxxiv, p. 638, 1930.
Locality of the type, Barbary.
Distribution. — Within historic times S.E. Europe, S.W.
Asia, Nobthbkn' India, and nearly the whole of Africa,
except the western and central forested area ; but now
exterminated in many districts.
Approximately equal to the tiger in size and in the relative
length of the tail, but distinguished from it by the complete
or almost complete loss of the pattern in the adult*, by the
uniformly darker or lighter tawny hue of the upper side, due
to the blending of blackish and pale speckling of the individual
hairs, by the presence of a black tuft at the tip of the tail,
* On the underside and on the inner side of the limbs spots are some-
times traceable in the adult, especially in the $, and occasionally,
in some B. African lions, at least, a faint pattern of rosettes, showing
a vertical, lineal arrangement, is visible on the flanks from the spinal
■area downwards, and becoming stronger on the belly and limbs.
PAHTHEEA.
211
of a black patch behind the ear, and by the absence of sharp
contrast between the tint of the flanks and belly and of the
outer and inner sides of the limbs. Also by the forward
and upward streaming of the hairs on the sides and top of the
neck to form a higher or lower median erect crest on the nape ;
and in almost all cases by the forward direction of the hairs
Fig. 60. — Upper view of skull of Lion from Amreli.
on the middle of the back from a whorl in front of the loins.
The <J is larger than the $ and, as maturity approaches,
typically develops a mane on the neck, consisting of a median
crest running along the nape from the shoulders to the Grown,
of a fringe on the cheek, and of longer hairs extending over the
sides of the neck to the breast between the fore legs. This is
p2
212 FELID-ffi.
accompanied by a tuft of hair on the elbow and frequently
by longer hairs on the chest and abdomen. But these hair-
growths are very variable in length and luxuriance even in
specimens from the same locality. In the extinct races from
Cape Colony and Algeria the long, thick mane covered the
entire neck and shoulders, and the hairs on the chest and belly
formed a luxuriant fringe ; but in none of the existing races
are such manes developed, and " maneless " lions have been
recorded from East Africa and Persia*.
Newly-born cubs are sometimes uniformly coloured, except
for faint spots on the underside ; but as a rule they have a
distinct pattern on the upper side consisting of rosette spots
like those of a jaguar or leopard, but these spots oom.Tin.only
show a tendency to arrangement in vertical lines and, as a
further stage, to coalesce and form looped stripes like those
of a well-marked tiger. Usually the pattern disappears
in about six months, but is occasionally retained for two or
three years or even more.
The skull typically differs from a tiger's by being lower
and flatter in the frontal region, with the " waist " or post-
orbital region, measured from the frontal process to the fronto-
parietal suture, shorter, so that the facial portion looks longer
and more massive as compared with the cranial ; the anterior
nares are wider, the nasal bones shorter, not projecting
posteriorly beyond the maxillae, and the lower edge of the
mandible is slightly convex in the middle, so that the skull
" rocks " slightly when placed on a flat surface ; also the
inner lobe of the upper carnassial tooth is smaller. These
differences hold good in a great majority of cases ; but tigers'
skulls are so variable that occasionally they are difficult to
distinguish from those of lions. The only difference I have
found to be absolutely constant lies in the mandible, which in
the tiger is slightly concave below, so that the skull never
" rocks " when resting on a flat surface.
22 a. Panthera leo persica Meyer.
Felig leo persicus, Meyer, Diss, inaug. de gsnere !Felium, Vienna,
p. 6, 1S26 ; Fischer, Syn.Mamm. p. 197, 1829 ; and of some later
authors.
Felis leo bengalensis, Bennett, The Tower Menagerie, p. 1, 1829
(nom. preocc).
Leo asiatious, Jardine, Nat. Libr., Felinae, pp. 121 and 266, 1834.
Felis leo goojratensis, Smee, Proe. Zool. Soo. 1833, p. 140, id.,
Trans. Zool. Soe. i, p. 165, pi. xxiv, 1834 ; and with various
renderings by several subsequent authors.
* The mane presents many analogies to the human beard. Its growth
is arrested by castration, and a small one may be acquired by old
lionesses.
MAMMALIA.
PLATE X.
Head of Indian Lion. (Adapted from engraving by Bennett of specimen
from Hariana exhibited in the Tower of London.)
The " Maneless Lion of Gujerat." (Drawn from one
of Capt. Smee's specimens.)
PANTHERA. 213
Felis leo indicus, Blainville, Osteogr. Mamm. Atlas, FeKs, pi. vi,
1843.
Felis leo peraicits, asiaticus, and goojrattensis, Matschie, SB. Ges.
Nat. Fr. Berlin, p. 94, 1900.
Panthera leo persica, Poeoek, Journ. Bomb. Nat. Hist. Soe. xxxiv,
pp. 638-65, 1930*, and xxxviii, p. 382, 1935.
Vernacular. — Sher, Babar-sher, Singh (Hindi) ; Untia-
bdgh (Camel-tiger) (Guzerati) : Sdwach (Kattywar) ; Shingal
(Bengali) ; Silh or Suh £, Siming 2 (Kashmiri) ; Bastar
(Brahui).
Locality of the type of persicus of Meyer, " Persia '* : of
Fischer, Teheran (ex Temminck, Mon. Mamm. p. 86, 1827) ;
of bengalensis, Hariana (according to Blyth) ; of a&iaticus,
Bussorah or Basra (but probably based on the same specimens
as Fischer's persicus) ; of goojratensis, Ahmedabad : of
indicus, India.
Distribution. — Formerly Persia, Mesopotamia f , no doubt
Baluchistan, and Nobthekk India from Sind in the west
to Bengal in the east and from Rampur and Rohilkund in
the north to the Nerbudda in the south. Now restricted to
the Gir Forest in Kathiawar.
Distinguished, on the average at least, from existing African
races by the sum of a number of characters : — The skull
has the auditory bullae less inflated, the postorbital area,
or " waist," measured from the frontal process to the fronto-
parietal suture, shorter and the infraorbital foramen typically
divided into two by a bridge of bone ; in external characters
the tail-tuft is larger, and the elbow-tuft and belly-fringe
are comparatively well developed in assoeiatkm. with a poorly
developed mane.
The coat, according to the season, may be short and sleek
or tolerably thick and long, long enough to be brushed in
all directions. The general colour is very variable, ranging
from ruddy-tawny, heavily speckled with black, to sandy -
or buffish-grey, sometimes with a silvery sheen in certain
lights and with the black speckling much less in evidence,
* Reasons for the synonymy here quoted were given in this paper
and need not be repeated. It is necessary, however, to add that the
evidence of racial identity between the Indian and the Persian lion is
not complete owing to the scarcity of Persian material. The skull,
for example, of the Persian lion is unknown to me. If ever the Indian
lion is shown to be distinct it will take the name goojratensis, since the
older name bengalensis is unavailable from having been applied to the
Leopard-Cat (F. bengalensis) in 1792.
t Fitzinger (SB. Akad. Wise. Wien, i, p. 362, 1868) stated that this
lion occurred in Greece and Palestine, which he could not have known,
and also in Afghanistan. I am not aware of the evidence of its former
existence in Afghanistan, but it may have been fount! in the southern
parts of that country.
214
EELIDJE.
and below, including the chin, and on the inside of the legs
from buff to nearly white. The mane also varies in luxuriance
and colour. It may consist merely of a crest running along
the nape from the shoulders to the crown and of a scanty
fringe on the cheek and throat, where the hairs are only about
Fig. 61. — Lower view of skull of Lion from Amreli.
4 in. long ; or it may form a luxuriant mat over the summit
and sides of the neck, the longest hairs being nearly a foot in
length. Its colour is usually tawny, with a mixture of blackish
and grey hairs, but it may be tolerably golden-tawny almost
throughout, or it may be heavily blackened along the crest
and low down in front of the shoulder.
PANTHERA. 215
Some recorded flesh-measurements are as follows : —
Head and
Locality, authority, and sex. body. Tail. Total.
ft. in. ft. in. ft. in.
Gir Forest (Lord Harris) ; <$ - - - - 9 ">
Gir Forest (Col. Fenton) ; <J 6 6 2 11 9 5
Gir Forest (Count Seheibler) ;<? 6 6 2 7 9 £
Gir Forest (Col. Fenton) ; <J - - - - 9 1
Gir Forest (Col. Fenton) ; <? - - - - 9
Gir Forest (A. S. Vernay) ; <J - - - - 9 1
Ahmedabad (Capt. Smee) ; J - - - - 8 9J
Fig. 62. — Lateral view of skull of Lion frora'Amreli.
None of the many dressed <J skins that have been measured
exceed the dimensions given above ; but a freshly stripped,
pegged-out skin measured by Lord Lavington was : — Head
216
FELID^.
■and body 7 ft. 6 in. ; tail 2 ft. 8 in. ; total 10 ft. 2 in. This
was no doubt stretched, as Col. Fenton supposed. A dressed
2 skin from Gujerat in the British Museum has the head and
body 5 ft. 4 in., the tail 2 ft. 8 in., and the total 8 ft. From
the evidence quoted above the Indian lion is the same size
Approximately as Central African Lions.
The essential characters of the skull of the lion are
given above. To this it may be added that the skull is
exceedingly well developed muscularly. The <$ from Amreli,
of which the dimensions* are entered below, for instance,
has a more strongly developed sagittal crest than in any
African lion's skuLl I have seen. Although fully adult it is
not an old skull. In connection with that peculiar feature,
the division of the infraorbital foramen into two orifices, it
Fig. 63. — Front view of skull of Lioness from Amreli, showing the dupli-
cation of the infraorbital foramen on the left side.
may be stated that out of thirteen skulls examined, eleven have
it divided on one or both sides, two only resembling African
lions in having it undivided. Sir Richard Owen first drew
-attention to this phenomenon in the skulls collected by
Smee.
The first skull on this list was presented to the British
Museum by the Maharajah of Bhavnagar through the kind
offices of Col. A. H. E. Mosse, LA. ; the second belonged to
a specimen shot by H.H. the late Maharajah Jam Sahib of
* Since sportsmen interested in "records" usually quote the total
length and the zygomatic width in English inches, I may add that
these dimensions in this skull are respectively 13-4 in. and 9-1 in.
It is as long as any recorded skull and a trifle wider.
PANTHEEA.
217
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218 FELmas.
Nawanagar ; the third and fourth were presented by H.H. the
Nawab of Junagadh ; the two remaining skulls of lionesses
are preserved in the Museum of the Royal College of Surgeons.
A Theory aboiii the " Maneless Lion of Gujerat."
When Captain Smee described the Indian lion he un-
fortunately chose for his paper the misleading title " The
Maneless Lion of Gujerat." Although he explained that
"maneless" was used merely in a comparative sense to
indicate, as he erroneously thought*, that his lions differed
from African lions by their smaller manes, the Gujerat lion
was over and over again cited as " maneless " by subsequent
writers. This race indeed seems to have been the source
of the highly fanciful theory put forward by General Rice
to account for the small manes many of them developed.
In years preceding the Mutiny, he wrote, lions were much
more plentiful in that part of India than afterwards and used
to live more in the open plains. No fewer than eighty were shot
in three years by a cavalry officer who chased them on horse-
• back over the open country. Prom being constantly hunted
and persecuted the lions were driven to take refuge in the
forests, where the numerous thorn-bushes dragged out the
best part of their manes until all except very old lions ceased
by degrees to have any manes left.
This notion that the combing action of thorns accounts
for the scantiness of manes in many lions has often been
thoughtlessly quoted with approval as supplying a satisfactory
explanation of the fact. There is not a word of truth in it.
The most that thorns could achieve would be keeping the
mane tidy by the removal of dead, moulted hair which might
for a time adhere to the growing mane before being shed.
They could no more affect its potential luxuriance than the
daily use of a comb can reduce the quantity of living hair
on a woman's head.
The Original Home and Destruction of the Indian Lion.
Prom the discovery of fossil remains of the lion in England,
Prance, and Germany, and from unmistakable sketches of it
left by early man f it is known that in prehistoric times
* His conception of African lions was probably founded on menagerie
specimens, which may grow heavy manes, oruponthe naturally full-maned
lions of Cape Colony or Algeria, which in the early part of the nineteenth
century were not uncommonly exported to Europe for exhibition.
t The only sketch known to me represents the beast as maneless.
This is noteworthy because the Neolithic artist would surely not have
selected a lioness for portrayal and would have depicted such an im-
pressive feature as the mane if is had been present in the lion. Possibly
at that date the lion had not acquired that secondary sexual character.
PAOTHEBA. 21&
the species was widely distributed over the whole of Central
Europe ; and from historical records it is also known that it
still existed in Macedonia at the time of the invasion of
Greece by Xerxes, and was tolerably abundant hi Palestine,
as told in biblical stories. It has long since ceased to exist in
those countries, but in Persia and Mesopotamia it occurred
until comparatively recently, although it is now apparently
extinct there.
At the time of the English occupation of India lions were
abundant throughout the northern portions of that eountry
from Sind in the west to Bengal in the east, and from Hariana,
Rampur, and Rohilkund in the north to the Nerbudda in the
south ; but the last of them are now restricted to the Gir
Forest in Kathiawar, an area of about 500 square miles.
which was set aside as a sanctuary and is owned by
H.H. Sir Mahabat Khan, Nawab of Junagadh. In 1880,
before protection was extended to them, there were said to
be only about a dozen left in that district ; but in 1930 their
number was computed to have increased to 200. This, however,
was probably an exaggeration, since Sir Patrick Cadell guessed
there were not more than seventy or eighty in 1935. The
dates of the extermination of the species in a few districts,
compiled by 1ST. B. Kinnear from various sources, are as
follows : — Palamau 1814, Baroda 1832, Hariana 1834, Ahmecl-
abad 1836, Kot Diji in Sind 1842, Damoh 1847, Gwalior 1865,
Rewah 1866, Abu and Guna 1872, Deesa 1878, Palanpur 1880.
The above stated evidence of the occurrence of lions in
Europe and South- Western Asia in early times supports the
view that the species made its way into India through Persia
and Baluchistan. That it is a comparatively recent immigrant
into India is attested by its restricted distribution in that
country. If India had been its home the lion would almost
certainly have travelled southwards to Cape Comorin and
reached Ceylon before the severance of that island from the
mainland. Probably, too, it would have crossed the Ganges
and Brahmaputra and entered Burma. Still further evidence of
the lateness of its arrival to constitute part of the Indian fauna
is supplied by its absence from the whole of the southern part
of Peninsular Hindustan. The only plausible explanation
of its failure to make its way south of the Nerbudda River
is that it was not given sufficient time. A check was put
upon its movements, and its rapid extermination was started
soon after the administration of India was taken over by the
East India Company.
In all parts of the world occupied by Europeans where
lions occur the disappearance of the lions is merely a question
of time. They are a menace to human life, especially if they
become "" man-eaters," and they ravage the flocks and herds
220 FELID^.
of settlers, finding domesticated livestock easier to prey
upon than wild game. This antagonism between lions and
Europeans, with all their resources for destruction, has led
to the extermination of lions in several areas of Africa where
they were at one time plentiful. Man, too, was doubtless the
principal agent, direct or indirect, aided perhaps by physical
factors lessening the abundance of big game or creating
unfavourable conditions, in the disappearance of lions from
Greece, Palestine, Mesopotamia, and Persia ; and there seems
to be no reason to look beyond man as the prime cause of their
extermination in India. It is only necessary to cite the
following instances attesting extensive slaughter in two
localities. During the Mutiny an English officer shot over
300 lions, fifty of them in the neighbourhood of Delhi ; and
Capt. Mundy in his ' Pen and Pencil Sketches,' 1832, stated
that before his time there were great numbers of lions in the
jungle near Hansi in Patiala, but that owing to the rewards
offered by Government for their destruction and the zest of
English sportsmen they had entirely disappeared from the
district. Incidentally he remarked that crack sportsmen
considered the lion afforded better sport than the tiger because
his attack is more open and his haunts less favourable for
retreat. This opinion of the difference in character between
lions and tigers agrees with that of Major Leveson, " the old
Skekarry," who, after experiences in shooting on foot tigers
in India and lions in Africa, came to the conclusion that the
lion is the pluckier animal of the two. Perhaps he should
have said less cautious and less cunning.
There is no reason to suppose that the two instances quoted
in testimony of the wholesale slaughter of Indian lions in
two districts were in any way exceptional where they were
plentiful. Similar slaughter was no doubt going on elsewhere
at the same time and before and after, resulting in the almost
complete extinction of the Hon in India, as it has been extin-
guished in Cape Colony, Algeria, and elsewhere in Africa.
Stress has been laid on the cogency of the evidence that
the practical disappearance of the Hon in India was due mainly
to the activities of the EngHsh Government and of EngHsh
army officers and civiHans because a famous forest officer
and a distinguished sportsman — the latter anxious to prove
that the tiger has more claim to the title " King of beasts "
than the Hon — put forward the opinion, which others have
accepted, that the tiger was the principal factor in the tragedy.
An obvious objection to this theory is the disappearance
of Hons in Europe and in the countries of S.W. Asia and Africa
referred to where there were no tigers to interfere with
them. But there are other facts to consider. There is
evidence that tigers entered India from the north round the
PANTHERA. 221
eastern end of the Himalayas through Burma, a different
route from that of the lion. Since, moreover, the tiger
spread all over South India, which the lion failed to reach,
it seems likely that the tiger's invasion of the country preceded
the lion's. In that ease the Hon made its way into Northern
India and multiplied exceedingly despite the tiger being
already in occupation of the country. Also the average
difference between the species in habitat makes it unlikely
they were ever brought into serious competition for a livelihood.
There would be no necessary rivalry between them ; and
the view that tigers waged organized warfare against the lions
or that combats between individuals, in which tigers were
victorious, were sufficiently frequent to lessen appreciably
the number of lions may be dismissed as fanciful, because
an encounter would just as likely end in mutual avoidance
as in a fight, and in the event of a fight the lion's chance of
success, so far as anything is known to the contrary, would
be as good as the tiger's. Hence there does not appear to
be a particle of evidence that the tiger played even a sub-
ordinate part in the extermination of the lion in India.
In the old days, as recounted by General Rice, Indian lions
"were sometimes pursued in the open on horseback, but in
thicker cover they were hunted on foot, unless elephants
were used for the purpose. In comparatively recent years
they were also shot on foot in the Gir Forest, as, for instance,
by Col. Fenton and his companion, who employed a gang
of beaters to drive them within range of the places where the
sportsmen were stationed. Some had narrow escapes from
following up wounded beasts ; and finally the killing of an
English officer by a lion led to the prohibition of " foot-
hunting " by those to whom the privilege of shooting in the
Gir was granted. Since then the method has been for sports-
men, safely lodged in a " maehan," to wait for the lions near
a " kill " or to shoot them from these platforms when driven
beneath by beaters.
A great deal has been written about the habits of African
lions, much less about those of the Indian race, and such
accounts as have come down to us are mostly records of
slaughter. But there is no reason to suppose there is any
difference in the two continents.
Except that lions generally frequent more open country and
are less secretive and more regardless of exposure than tigers,
the habits of these two great eats are on the whole tolerably
similar. Being approximately equal in size and strength,
they are alike capable of killing large and powerful game.
Lions, however, except at the breeding season, appear to be
more sociable, judging from accounts by African sportsmen
of two or more full-grown males combining to pull down
222 FEIXDiE.
a. buffalo. It is well known that in Africa lions may become
habitual man-eaters, like tigers, and apparently under the
same circumstances. It is likely enough that in the past
Indian lions did the same ; but at the present time they
seem usually to avoid man, and only to attack him in self-
defence. In the Gir Forest they prey mainly upon the big
game and domesticated livestock. It has been recorded that
they he up during the day in their lairs, and issuing from them
just before sunset make straight for the nearest hamlet of
buffalo-herdsmen. If unsuccessful in their quest they
move on to the next and wait for stragglers from the byres.
Since the cattle are driven out to pasture long before daybreak,
the lions have a good chance of a kill while it is still dark.
In the ease of African lions it has often been stated that
the male, unlike the tiger, stays with the female after the birth
of the cubs and helps her to protect them and later to get
food for them. That the same is true of the Indian lion is
attested by an account given by General Watson of an
unprovoked attack made upon him at Hariana by a Hon
and lioness who were lying up in a patch of thick jungle and
sprang out as he was riding past. He shot the pair and
afterwards found a couple of small cubs in the thicket*.
A lion is in his prime when about five years old ; but
Dr. Vevers tells me of an instance of a lioness that bred
when she was only twenty-two months. Their potential
length of life is probably the same as in tigers. The period
•of gestation is about sixteen and a half weeks (116 days) ;
and the cubs, usually from two to five in number, may be
born with their eyes already open, although frequently they
are closed.
23. Panthera pardus (Linnaeus). The Leopard or Panther.
Felis pardus, Linn., Syst. Nat. ed. 10, p. 41, 1788 ; and of most
subsequent authors.
Panthera pardus, Pocock, Ann. Mag. Nat. Hist. (8) xviii, p. 316,
1916 ; id., Journ. Bomb. Nat. Hist. Soe. xxxiv, p. 64, 1930.
Locality of the type, Egypt (Cabrera and Thomas ; restricted
to the Egyptian Sudan by Holhster).
Distribution. — The whole of Asia south and east of a line
running roughly from Asia Minor and the Caucasus to Man-
■churia and Amurland, and reaching Kangean Island to the
east of Java ; also practically the whole of Africa except
"the Sahara.
Size considerably less f than that of the tiger or Hon, the
* This was in the early twenties of the last century. The two cubs
were hand-reared and sent to England to make history as the first
Indian lions exhibited in this country.
t These remarks apply to Indian representatives of Panthera pardus.
PLATE XI.
MAMMALIA.
Variety of Indian Leopard from Cuddapah.
FAOTHERA. 223
length of the head and body seldom exceeding 4 J ft., -with the
tail considerably over half the length of the head and bod}^,
the hair on the cheeks never growing into a definite fringe
longer than that of the body and, although the hair on the
nape typically forms a median crest from the junction of
convergent streams from the sides of the neck, as in lions,
it never, even in the £, forms anything approaching a definite
mane. The coat varies greatly in length, thickness, and tex-
ture, according to the season, in some races ; the ground-
colour above is also exceedingly variable, from greyish or
whitish-buff, with sometimes hardly a trace of buff, to
olivaceous with a huffish tinge, or to bright reddish-oehreous ;
below, from the front of the upper lip and chin backwards,
including the lower cheek, the lower side of the tail and the
inner surfaces of the limbs, it is generally wholly white, and
sharply contrasted with the tint of the upper side and outside
of the limbs, though the paws may be paler than the rest of
the limbs ; there is a white patch on the otherwise black
back of the ear, but there is no white patch over the eye as
in the tiger. The pattern is very different from that of the
tiger, but not unlike the pattern of some lion-cubs. It-
consists of solid black spots on the head, sometimes for a short
distance behind it, on the outer side of the limbs and on the
belly, which is typically heavily spotted, but elsewhere on the
body the pattern mostly consists of definite " rosettes " which
appear to result from the coalescence of some four or five
smallish solid spots to form definite but irregularly shaped
rings surrounding an area of darker tint as a rule, especially
low down on the sides, than the brighter tint of the interspaces
between the spots. The rosettes vary considerably in size
and spacing, and on the spine, especially over the hind back
and loins, they are usually more elongated and show a tendency
to run in longitudinal stripe-like lines. On the tail the pattern
is less regular and less rosette-like, the individual rosettes
being liable to coalesce more or less towards the end above,
whereas the underside here may be almost wholly unspotted,
so tha. when upturned it looks white.
The normal pattern of leopards, above described, with its
suggestion here and there of longitudinally lineal arrangement,
is much more primitive than the highly specialized transverse
stripes of the tiger.
Variation in the Colour and Pattern of Leopards.
On account of its extensive distribution and capacity for
adaptation to widely different environments the leopard varies
locally in coloration much more than the lion or tiger, and many
geographical races or subspecies of it have been described.
224 FELXB.3E.
Those admitted in the Indian fauna are considered below.
But the species is also susceptible to an unusual amount of
individual variation in colour and pattern in the same locality.
These variations are termed " varieties " or " sports." The
" black " variety is the commonest of them. In this type
the blackness is due to the deposit of black or dark brown
pigment in the hairs that are normally yellow or white, so that
the spots are obscured although probably always visible in
certain lights and generally at least better defined on the
underside because the normally white hair is browner than
the normally yellow hair of the upper side.
There is no doubt that in certain parts of British India
black leopards are much commoner than in others. They
have been recordedfrom Ceylon, and appear to be not uncommon
in parts of South India and Assam. They crop up in Burma,
and the percentage of black specimens seems to increase
southwards through the Malay Peninsula until they become
the dominant, if not the sole, type of leopard found in the
south of that country. There is some evidence that the black-
ness is associated with localities where heavy rainfall is preva-
lent. As a very general rule, at all events in British India,
blackness is a discontinuous variation — that is to say, a leopard
is either " black " or normally coloured, and cubs of the two
types have been found in the same litter. But at Melghat
in the Central Provinces Dunbar Brander saw a dark chestnut
leopard, with black spots, which apparently represented an
intermediate stage.
Leopards sometimes also exhibit the opposite phenomenon,
namely, failure to develop pigment in the normally pigmented
areas. This may result in complete albinism, when the pattern
as well as the interspaces are white. I have seen only one
skin in this condition, and it was said to have come from
Africa. But there is a partially albino skin in the British
Museum which came from Hazaribagh in Bengal (R. E. S.
Thomas). In this the ground-colour is much paler than usual,
almost cream, and the pattern is tan.
The pattern of leopards also sometimes varies remarkably,
as attested by two skins from S. India. In one specimen shot
by F. A. Coleridge at Putnam in Cuddapah the ground-
tint is normal, but the rosettes on the whole of the upper
side have fused into a number of large blotches, with black
rims and enclosing many small spots, separated from each
other on the flanks by narrow pale lines forming a network
pattern, but on the spinal area fusing across the middle line.
This pattern recalls that of some so-called Clouded Leopards
(see p. 247). The second skin was secured by Sir C. A. Souter,
I.C.S., at Kanara. It differs from the last in that the
blotches have fused almost everywhere and additional black
PLATE XII.
MAMMALIA.
Variety of Indian Leopard from Kanara.
PAlsTHERA.
225
pigment has been added; with the result that the leopard is
mainly black above and on the sides, with a few yellow streaks
and spots representing the normal yellow ground-colour.
Although this leopard is nearly black, it is not a " melano "
like the ordinary black leopard. Its blackness is due to an
entirely different process, namely, the distintegration and
fusion of the pattern, and the underside is white*. The skin
of the hind quarters of another leopard of the same general
type as the one from Kanara, but black and marked with yellow
stripes on the sides, was purchased in Madras and presented
to the British Museum by Mr. P. Deraniyagala.
SkuUs of Indian Leopards.
In shape and general proportions the skulls of leopards
typically look like small tigers' skulls. So far as British India
is concerned the difference in size is always well marked, the
largest <§ leopard skull being a good deal smaller than the
Fig. 64. — Side view of skull of adult ^ Leopard from Kumaun, showing
flat dorsal profile. .< J.
smallest skull of a tigress or lioness ; but the small tigress
of Bali, the south-eastern limit of the species, hardly exceeds
a $ leopard in the size of the skull. In general characters
the skulls combine the characters of the skulls of the two larger
species. As a general rule, and for the same reason, the skulls
of British Indian leopards will " rock " to a varying degree
backwards and forwards like a lion's when on a horizontal
plane ; but sometimes, as shown by an adult £ skull from
Ashkote in Kashmir, they rest as steadily as a tiger's. On
the other hand the general shape of the skull, with its more
or less convex dorsal profile, dtie to the elevation of the
* Photographs of this skin and of the one from Cuddapah were*
published in my papr in 1 930, cited above. Sir C. A. Souter informed mo
that when the native who killed this leopard in Kanara applied for the
bounty his applieation was refused because the Commissioner did nnt
recognize the animal a3 a leopard !
VOL. I. q
226 FELLD^.
frontal region, is typically much more like a tiger's than a
lion's skull ; but Col. Stockley sent me the skull of a ^
leopard from the Kumaun Hills which has the dorsal profile
as flat as in a lion. It looks indeed like a diminutive lion's
skull (fig. 64). The nasal bones in leopard skulls may fall short
of the maxillae as in lions or surpass them as in tigers, but they
never surpass them to the same extent as in typical tigers.
The skulls, in short, vary in all details, in the width of the
mesopterygoid fossa, the inflation of the auditory bullae,
the development of the sagittal crest, and the inflation of the
forehead, the highest point of which may be in front of the
postorbital processes or behind them ; and these differences
are not racial, but individual and of no systematic importance.
The differences between <? and 2 skulls are usually well
marked and similar to those of the tiger and lion. The <J
skull is not only larger but is more moulded by the action of
the jaw-muscles, the fore part of the cranial portion being more
compressed so that the postorbital area or "waist" is lengthened
and narrowed, and is about the same width as the interorbital
area. In the 2 the " waist " is typically short and broad,
broader than the interorbital area. The teeth of the 2 are
on the average smaller, the canine particularly being narrower
(shorter) close to the socket *.
23 a. Panthera pardus fusca (Meyer). The Common Indian
Leopard or Panther.
Felis fusca, Meyer, Zool. Ann. i, p. 394, 1794 +.
Felis longicaudata, Valenciennes, C. R. Acad. Sci. Paris, xlii,
p. 1036, 1856 (nom. preocc).
Panthera antiquorum, Fitzinger, SB. Akad. Wiss. Wien, lviii,
p. 47, 1868.
Felis pardus antiquorum, Matsohie, SB. Ges. Nat. Fr. Berlin,
1895, p. 194 (not F. pardus antiquorum Gray, 1827).
* The difference between the <J and <j> skulls of this species was one of
the sources of the belief that two distinct kinds, the panther and the
leopard, live side by aide in the same locality. This opinion dates back
to the time of Temminck. It is quite needless to discuss the point
further, since all zoologists, and probably most sportsmen, now know
that the panther and the leopard are specifically the same animal.
Blanford devoted a good deal of space to the discussion of this topic,
which formerly greatly exercised the minds of naturalists and sportsmen.
He and others also discussed at some length the supposed differences
between "the Asiatic" and "the African" leopard. But it is now
known that there are several local races in both continents, and that
there are no such things as " the Asiatic " and " the African " leopard.
On the material at my disposal I am unable to find any difference between
the typical leopards of India and of Kenya Colony in East Africa.
f The name Felis fusca, the earliest apparently given to an Indian
leopard, was based by Meyer upon the figure and description published
by De la Mefherie (Journ. de Phys. xxxiii, p. 45, pi. ii, 1788) of a black
specimen alleged to have come from Bengal and exhibited in the Tower
MAMMALIA.
PLATE XIII.
A & D. "Waist " and side view of skull of Indian Leopard.
B & C. The same of Indian Leopardess.
PANTHEKA. 227
Felts pardus pardus, Dollman, after Lydekker, Game Animals
of India, p. 314, 1924 ; Rowland Ward's Records, p. 482, 192S
(not F. pardus Linn.).
Panthera pardus fusca, Poeock, Journ. Bomb. Nat. Hist. Soc.
xxxiv, p. 307, 1930 ; Phillips, Man. Mamm. Ceylon, p. 162,
1935.
Vernacular. — Tendwa, Chita, Sona-chita, Ghita-bdgh, Adnata
(H.) ; Palang (Pers.) ; Diho (Baulch.) ; Suh (Kashmiri) ;
Vidua, Srighas (Bundelkand) ; Oorbacha or Borbaclw, (Deccan) ;
Karda, Asnea, Singhal, Bibia-bdgh (Mahr.) ; Tenduioa, Bibla
(Bauris of Deccan) ; Honiga, Kerkal (Canarese) ; Teon-Kula
(Kol.) ; JerJcos (Paharia of Rajmehal) ; Burkal, Gorddg (Gond.) ;
Sonora (Korku) ; Ghiru-ihai (Tarn.) ; Chinna-puli (Tel.) ;
Puli (Mai.) ; Kutiya (Cingalese) ; Bai-hira, Tahir-M, Goial-M,
or Ghor-he (hill-tribes near Simla) (according to Jerdon,
generally known as Lakhar-bagha, a name elsewhere used for
the hyssna) ; Sik (Tibetan) ; Syik or Syiak, or 8ejjiak (Lepeha) ;
Kajengla (Manipuri) ; Misi patrai, Kam-kei (Kuki) ; Hurrea
kon, Morrh, Rusa, Tekhu Khuia, Kekhi (Naga) ; Kya-lak or
Kya-thit (Burmese) ; Kla-preung (Talain) : Eicke-phong
(Karen) ; Rimau-bintang (Malay).
Locality of the type, Bengal.
Distribution. — The whole of Ihdia from Kashmir and
apparently the foothills of the Himalayas in the north to
Cape Comorin in the south and thence to Ceylon : also found
in Bubma.
The coat soft and smooth, at its best seldom exceeding 25 mm.
(1 in.) in length and not thickened with any appreciable
amount of underwool. The general ground-colour of the
upper side bright at all seasons apparently, but varying
a little in intensity from nearly golden to ochreous or orange-
tawny, darker on the back than on the flanks. Pattern also
variable, the rosettes sometimes small and close-set, sometimes
larger and more spaced, their centres typically darker than the
interspaces, but not noticeably in the darker, richer-tinted
skins. The underside and inner surfaces of the limbs white
with black spots.
In 1930 I described briefly all the Indian panther-skins
of this race then available for examination to show their
variations in colour and pattern. Most of them were single
specimens from the following scattered localities : — Ashkote,
Xasair, and Kannah near the western border of Kashmir,
of London. The second name in the synonymy, S. lonpicawtlaia, ww
used by Valenciennes for a supposedly long-tailed leopard from Ceylon
and the Malabar coast. But the name had boen previously appliod by
de Blainville to a specimen of Clouded Leopard (Neofeliti) from Cochin
China. The other names given to Indian leopards wore discussed in
my paper in 1930 and need not be further considered.
o2
228 FEUD.®.
north of Srinagar (Mrs. Entwistle) ; Rhikhikesh, 3,000 ft.,
probably near the Siwalik Hills (Capt. F. S. Tuker) ; Haripur
Kheri in Oudh (A. P. Millard) ; Daltonganj in Western Bengal
(C. A. Crump) ; Mandri on the Tapti, 24 miles east of Surat
(T. B. Prv) ; Byadgi Station, east of Hausbari, 2,000 ft., in
S. Dharwar ; Kolar Town, E. Mysore, 4,026 ft., and Wotte-
kolle, S. Goorg, 2,000 ft. (G. C. Shortridge). Apart from the
skin from Rhikhikesh, which is exceptionally pale, owing,
I am now convinced, to exposure to light as a rug, all these
skins are of the same general type, but differ individually
to a certain extent in tint and pattern. Of particular interest
is the closeness of the likeness both in colour and pattern
between the skins from S. Dharwar and from Nasair in Kashmir,
both agreeing tolerably nearly with the skin from Dalton-
ganj which, coming from Bengal, may be regarded as approxi-
mately a topotype of fusca. The occurrence of the same
leopard, with similar individual variations in tint and pattern,
practically over the whole of Peninsular India is shown by
many specimens received since 1930, namely, from Manikpur,
N.P. (Major G. Burrard) (December) ; from Gir, and Bhav-
nagar in Kathiawar (January and February), and Mahi
Kantha, N. Gujerat (Col. A. H. Mosse, I.A.) "(April) ; from
Mt. Abu, Rajputana, Bhopal and Narsinghgarh in the Vindhya
Hills, Bhandara and the Surguja State, C.P. (H.H. the
Maharaj Kumar, Heir Apparent of Bikanir) ; from Gund-
lupet, S. Mysore, 2,600 ft. (Major E. G. Phythian Adams) ;
and from Hogainakal in the Dharmapuri Range, N. Salem
850 ft. (N. A. Baptista) (August). The last, the only specimen
seen from the Eastern Ghats, was procured on A. S. Vernay's
expedition. The dated skins presented by Major Burrard
and Col. Mosse establish the absence of appreciable underwool
in the tolerably long winter coat. Of six skins from Ceylon,
one from Ambawela is a trifle darker and richer in colour
than any of the Indian skins examined, whereas one from
Kala Oya is a little paler, a third from Hambantota, S.P.,
being intermediate. These were collected by E. W. Mayor.
Two immature skins from Gammaduwa (W. W. A. Phillips) are
about average in tint, and an adult § from Pollaranua, KT.C.P.
(E. C. Fernando) (July), is very nearly as dark and rich as
the Ambawela skin, but agrees tolerably closely with the
richest skins from Surguja State, C.P., India. These sk in a
fit the general description of the Ceylonese leopard published
by Phillips. Possibly the pale skin from Kala Oya represents
the lighter, " almost lemon-yellow variety " he mentioned as
occurring both in the hills and low country, more particularly
in the latter. All the Ceylon skins I have seen have small
dose-set rosettes, smaller on the average than in Indian skins.
PLATE XIV.
MAMMALIA.
Skin of Indian Leopard with exceptionally bold pattern from Rhikhikesh.
PANTHBBA. 229
According to Phillips black leopards are occasionally found
in the island.
There are very few records of Burmese leopards. The only
normally tinted skin I have seen came from Tounghoo (J. M. D.
Mackenzie). In its bright colour, medium-sized rosettes,
and shortish thin coat it is inseparable from several of the
handsomer skins oifu&ca from India. Other skins seen from
Burma were black, namely, from Mogaung, Upper Burma
(Capt. W. Abbey), the Shan States (Poland's Coll.), and
Mt. Popa (G. C. Shortridge).
Most of the recorded flesh-measurements of British Indian
leopards were, unfortunately, taken from " tip to tip " or
" between pegs," an unsatisfactory method because it gives
no idea of the proportion of the head and body to the tail.
There are only a few properly measured skins in the British
Museum ; but Gen. R. G. Burton recorded the dimensions
of several from Berar (Journ. Bomb. Nat. Hist. Soe. xxi,
p. 1063, 1911), and Mr. W. W. A. Phillips has done the same
for Ceylon specimens. The results are entered in the following
table, which shows that the average total length of the males
is a little over 7 ft. and of the females nearly 1 ft. less. Accord-
ing to Dunbar Brander ('Wild Animals in Central India,'
p. 130, 1923) " a fair average male leopard measures 6 ft. 8 in. ;
. . . the large jungle-living animal is anything from 7 ft. 2 in.
up to 7 ft. 9 in., a fair average specimen being 7 ft. 5 in."
Possibly an exceptionally large <J may reach 8 ft. or a little
over.
Head and
Locality, authority, and sex. body. Tail. Total
ft. in. ft. in. ft. in.
Berar (Burton) ; £ 4 6 2 i) 7 3
Berar (Burton) ; £ 4 3 2 7
Berar (Burton) ; £ 4 2 2 6 8
S. Dharwar (Shortridge) ; (J 4 2 3 7 2
Ceylon (Phillips's largest) ; jj 4 8 3 2 7 10
Ceylon Phillips's av. of U (approx.) ; <J 4 2 2 10 6 11}
Berar (Burton) ; „' 3 X 2 6 2
Berar (Burton) ; T 3 7 2 7 ($ 2
Daltonganj (Crump) ; yg. ad. i 3 3 J 2 7 6 1
Toungoo (Mackenzie) ; ad. i 3 10* 2 10} 8 8}
Ceylon, Pollaranua (Fernando); ad. ?• 4 2 !> *5 St
Ceylon (Phillips's largest) ; £ 3 9 2 9 6 6
Ceylon (Phillips's a%\ of 7) (approx.) ; ?. 3 ."> 2 lik fl 11}
The weight of Phillips's largest $ was 170 lb., the average
of that sex being 124 lb. His largest $ was 75 lb., and the
average of 7 $$ (54 lb. The difference between the two sexes
is very remarkable. Dunbar Brander gave the weights of
2 £ leopards from Central India as 152 and 1101b. respectively.
The table of skull-measurements, containing the largest
230
STEUD^.
"8 W
#
w *
.a
FM •s
W
6 o
3
ifl
fei
« n
PAKTHERA. 231
and smallest available, shows that the total length in the
$ varies from, about 9 to 10 in., in the $ from 7 to 8 in. The
only specimen of special interest is the one queried as $ from
Sonnawani Block, C.P. It is unsexed and is not represented
by a skin. Although its condylobasal length and zygomatic
width are exceptionally large for a Q, it is too small for a <£,
but the practical equality in width between the postorbital
and interorbital areas is a <J character, and it may be the skull
of a '" dwarfed " g. There are one or two dwarfed races of
leopard in Africa in which the skull of the adult q has
characters.
The average condylobasal length in 16 adult q skulls from
India is just over 212 mm., and of nine adult $ skulls 172 mm.
In two adult $ Burmese skulls from lit. Popa and Toungoo
respectively the condylobase is 170 mm., in one from the
Ruby Mines (Shortridge) it is 165 mm. They thus agree
with the Indian series. The same applies to Ceylonese skulls.
The largest <£, from Gammaduwa (Phillips), has a condylo-
base of 217 mm. ; in the smallest from S. Ceylon (Bevan)
it is 206 mm., the average of five adult <££ being 2L1J mm.
The condylobase in the largest and smallest $ skulls is 178
and 173 mm. respectively, but the size is more uniform than in
Indian skulls, the average of seven, mostly from 8. Ceylon
(Bevan), being 175 mm.
23 6. Panthera pardus pernigra (Hodgson)*.
Leopardus perniger, Hodgson, Gat. Matnin. etc. Xepal, <«i. 2,
p. 3. and Preface, p. v, 1863.
Locality of the type, Sikhim, (5,000 to 8,000 ft.
Distribution. — Sikkim, Nepal.
Distinguished from the southern race, fu-sca, at least by the
coarser, thicker, more woolly winter coat. In normally
coloured specimens the rosettes are large and stand boldly
out against the paler ground-colour.
Three out of the six skins belonging to Hodgson's collection,
and definitely assignable to this race, are black ; and in his
Catalogue he remarked, " llany more were got. I have
a dozen skins. People say it is a distinct species." Prom this
it may be inferred that a considerable percentage of the leopards
* The type of this race is one of three black leopards from Sikkim
which Hodgson reeordod as a " black variety of Leopard " in the
Preface to his catalogue, giving no description of it, under the name
Leopardus perniger. 1 overlooked the description in the Preface and
also Cabrera's reference to it when I wrote my paper on the Panthers
of Asia in 1930, and in the account of Hodgson's specimens (p. 309)
I identified them as P. p. fiwca, cited Nepal as their locality, and quoted
pernigra as a n»men nudum.
232 FELIDJE.
at 6,000 or S,000 ft. in Sikkim is black. In addition to these
black specimens, two in Hodgson's collection labelled " Nepal "
have similar thick, longish, and rather harsh coats. One
has the ground-tint darkish, nearly rusty-buff, with darker
centres to the rosettes, the belly is pale buffish and the hind
throat decidedly buff. The other has the shoulders and
flanks unusually pale and emphasizing the rosettes, which have
rusty centres. * A third Nepalese skin (R. Everest) is very-
like the last in coat and colour; but a skin from Kaknai,
Khatmundu, 7,000 ft. (Col. R. L. Kennion), in summer
coat, is not distinguishable from the southern race, fusca.
Also apparently referable to pernigra is a skin from Barkat,
Dehra Dun, 5,000 ft. (Capt. F. S. Tuker), December. The
coat, about 30 mm. long, is decidedly thickened with wool,
and the colour is like that of the pale specimens from Nepal,
much paler than in typical fusca, the dorsal area being buffy
and the sides greyish, emphasizing the rosettes, with their
dark ochreous centres. Very similar to the last, both in coat
and colour, is a half-grown specimen from Bharnabari, Bhutan
Duars, 600 ft. (N. A. Baptista), collected in March. If this
identification be correct, the race is not restricted to high
altitudes. Very likely the leopards descend to lower levels in
the winter *.
In the table of skull-measurements (p. 236) I have provision-
ally entered under pernigra, on account of their localities,
there being no skins for examination, a skull from the Kumaun
Hills (Col. C. S. Stockley) and one from Mundiapani in Garh-
wal (B. B. Osmaston). The latter is fully adult, and is
only remarkable for its small size. The skull from the
Kumaun Hills, on the contrary, is very peculiar. It was
referred to above (pp. 225-6) on account of the flatness of its
dorsal profile and generally lion-like aspect. Although not
quite adult, it has an exceptionally high sagittal crest and
salient occipital ridge. The teeth also are unusually large,
partly, no doubt, because they are quite unworn. There is
a skull of fusm, from the United Provinces (R. St. 6. Burke),
with the carnassials 28 and 20 mm. respectively ; but these
teeth are on the average much smaller in Indian leopards
than in the skull from Kumaun.
* A few years ago Col. F. M. Bailey sent to me the skin, of a very
beautiful leopard, richly coloured, and with a full, long coat, the hairs
being ahout 47 mm., nearly 2 in., long, which was killed near Shigatse
in Tibet. On the evidence of a single skin I cannot separate this leopard
fron tiiP race from Shensi, in China, which I described as Panthera
mrdm bedjordi in 1930. It is possible that this leopard may occur on
the northern slopes of the Himalayas and be later recorded as a member
of the British Indian fauna.
PLATE XV.
MAMMALIA.
'— ■* — i\m
Skin of Millard's Leopard (Panthera pardus m/l/ard/) from Kashmir.
FARTHER A. 233
23 c. Panthera pardus millardi Pooock.
Panthera pardus millardi, Pocock, Journ. Bomb. Nat. Hist. Soc
xxxiv, p. 316, pi. viii, 1930.
Locality of the type, " Kashmir."
Distribution. — Kashmir only, so far as is at present known.
Distinguishable at a glance from the typical Indian race
fusca and from the Nepalese and Sikkiin race pernigra by the
entire absence of bright hue in the pelage, the general colour
being dull, dark buffy-grey, almost olivaceous, the pattern
consisting of small close-set rosettes. The coat, too, is much
thicker with underwool than in fusca.
Only two specimens of this race are known, both received
from Sirs. Entwistle and labelled " Kashmir," without further
particulars. They are entirety different in appearance from
the skins, also procured from Mrs. Entwistle from Ashkote
and Nasair in Kashmir, which are identified as fusca- From
the thickness of the coat it may be inferred that millardi
occurs at much higher altitudes in Kashmir than the other
race. The only known skull, that of the type, an adult, %,
shows no peculiarities except that it is unusually large for
a Q, a little longer in its total and mandibular lengths than the
largest 2 of fmca entered in the table.
23 d. Panthera pardus sindiea Pocock.
Fells tulliana, Blanford, Manim. Brit. Ind. p. 69, 1SS8 (in part)
(not tulliana Valenciennes).
Panthera pardm nindica, Pocock, Journ. Bomb. Xat. Hist. Soe.
xxxiv, p. 80, 1930.
Locality of the type, the Kirthar Range on the Sind-
Bahichi border.
Distribution. — The Kirthab Rajstue and S. Wazaristan.
Intermediate in colour between the typical Indian and the
Persian races, not so bright as the former, but brighter than
the latter, and differing from both in its winter coat and slightly
in pattern, the winter coat being thick, erect, and peculiarly
harsh, but not long and tufted, about 26 mm., with the tips
of the hairs curled ; and the pattern consisting of rosettes,
which on the average are larger, more spaced, more annuli-
form, with thin unbroken rims, but no contained spots.
This race is not well known. I have seen only two skins.
The type, a young <J, shot by H. E. Watson in the Kirthar
Range, was identified by Blanford as tulliana and described
as representing " a race inhabiting Persia and found in Baluchi-
stan and the mountains of Sind that differs widely from all
the others [Indian leopards] and is quite intermediate in
coloration and spotting between the leopard and the ounce."
234 FEUD.®.
This description applies well enough to the race described
below as saxicolor, but not to the skin from the Kirthar Range,
which, although undated, was apparently shot in winter,
the coat being thick and harsh, as stated above ; the flanks
are washed with buff, which becomes intensified on the back,
the general hue being much brighter than in the Ounce, but
not so bright as in the ordinary Indian leopard. It is signi-
ficant, however, that Blanford could see the difference between
this skin and skins of the typical Indian race. The second
skin, received since the race was described, was sent to me
by Major D. G. Lowndes from S. Wazaristan. In colour and
pattern it closely resembles the skin from the Kirthar Range,
out the coat is very different, being short and sleek, as in the
July skin of saxicolor from Kushnob, but it decidedly differs
from that skin in its brighter hue and pattern.
The only known skulls of this race, both from the Kirthar
Range, show no structural peculiarities.
Of the habits of this leopard nothing has been recorded.
23 e. Panthera pardus saxicolor Pocock. The Persian
Leopard.
Felix leopardus, P. L. Selater. Proc. Zool. Soc. 1878, p. 289 (not
of Sehreber).
Felis tull-iana, Blanford, Mamm. Brit. Ind. p. 60, 1888 {not of
Valenciennes).
Fells pardwi panthera or tulliana, Lydekker, Game Animals of
India, p. 322, 1924 (not Felis panthera Sehreber).
Panthera pardus saxicolor, Pocock, Ann. Mag. Nat. Hist. (9)
xx, p. 213, 1927 ; id., Journ. Bomb. Nat. Hist. Soc. xxxiv,
p. 77, 1930.
Locality of the type, Asterabad, Persia.
Distribution. — Persia and Balttchist^n'.
Distinguished from the Indian races in both summer and
winter coat by its much paler colour, the general tint being
grey, with at most a pale buff or sandy wash, particularly
on the dorsal area. Also by the very marked seasonal differ-
ence in the coat, which in summer is short and sleek, in whiter
long, thick, tufted, and woolly, the eolour and coat combined
at this season giving the animal a pronounced likeness to the
Snow-Leopard, with which it has more than once been con-
founded (see p. 240).
This race is included in the fauna of British India on the
evidence of two specimens received since it was described.
The skin of an adult $ from Kushnob, Ziarat, about 50 miles
north-east of Quetta (H. J. Todd), July 9, has the summer coat
short and sleek and the general colour sandy-grey, with the
rosettes mostly broken up into five or six spots, the centres
PANTHERA. 235
of those on the spinal area not darker and of the flanks only
slightly darker than the ground-colour ; the underside and
inner sides of the legs white. This skin closely resembles
skins from Persia in summer coat.
A second skin, from Sambaza, Baluchistan, 4,500 ft. (A. H. K.
Sangster), April, has the full winter coat thick, loose, tufted,
and long, about 50 mm. (2 in.) in length, the tail bushy and
the general colour pale grey, with a sandy wash on the back.
It closelyresembles askinfromPalang Kuh,in Seistan, collected
by Col. B. L. Kennion, which in that author's book, '' By
Mountain, Lake, and Plain,' p. 267, 1911, was referred to by
Lydekker as representing an undescribed variety of leopard.
Both are very similar in colour and coat to the Snow-Leopard,
except that the spots are smaller, more numerous, and less
spaced.
The flesh-measurements, presumably, of the ^ from Kushnob,
recorded by Mr. Todd, are : head and body 5 ft., tail 3 ft. 2 in.,
total 8 ft. 2 in. Since the large size is not borne out by the
skull, the dimensions were possibly taken from the stripped
skin. The skull of this race from Kushnob entered in the table
of measurements (p. 236) is only just adult, perhaps not quite
full-sized. It is at all events a good deal smaller than an
adult £ skull from Mishun in Persia (C. E. Capito), in which the
condyiobasal length is 218 mm. and the zygomatic width
135 mm.
This leopard has long been known, and there are earlier
records of it than the one that heads the list of synonyms ;
but it has passed under inadmissible names, being regarded
by some authors as the Snow-Leopard (Uncia) and by Blanford
and Lydekker as identical with the more richly coloured
race inhabiting Asia Minor, Panthera pardus titlliana, the
type of which came from near Smyrna. The two appear to
intergrade in Mesopotamia. I am indebted to Mr. C. E. Capito
and to Mrs. Lane for the opportunity to examine several skins
from the Pusht-i-Kuh Range in Laristan and the province of
Pars in S. Persia, Sir Percy Cox and Major Cheesman also
secured it in Pusht-i-Kuh, where, according to Mr. Capito, it
inhabits caves and gorges in the barren limestone and gypsum
hills down to about (MX) ft. Its habitat appears to be much
the same as that of the Snow- Leopard, and its coloration no
doubt harmonizes with the of the rocky hillsides it frequents.
Origin and Habits.— AH the evidence known to me points
to the conclusion that the leopard, like the tiger and lion,
was of northern origin and migrated southwards to the
countries where it is now found. Fossil remains have
been recorded from Centra! and Southern Europe. It is
236
tfELTDJE.
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PANTHEBA. 237
a more primitive and no doubt earlier type than the
lion and the tiger ; and since its nearest ally, the jaguar,
is an American species, the common ancestor of both
must have inhabited northern Asia at a time when the
two continents were joined by the bridge of land that
formerly separated the Arctic and Pacific Oceans. That the
leopard was already in Hindustan when the tiger reached it
is shown by its presence in Ceylon ; and there is no reason
to doubt that it similarly preceded the lion in its occupation
of the country. By what route it entered India is uncertain ;
but since the resemblance between the typical Indian race
and the leopards of Burma and China is greater than the
resemblance between the former and the leopards of Baluchi-
stan and Persia, I incline to the view that it probably came
into India, like the tiger, by a north-eastern route, perhaps
across the Himalayas, since it occurs in Tibet, north of Sikkim,
as well as to the south of that range. But the known facts
do not justify more than a suggestion on that point.
Although the habitats of the different races of British Indian
leopards vary, the habits of all in a general way are probably
very much the same. Those of the ordinary Indian leopard
are well known, and they differ in certain respects from those
of tigers. Being much less intolerant of the sun than tigers,
leopards are less nocturnal and more often hunt by day,
especially in the event of failure to kill by night, their usual
time of activity. They are, too, less addicted to thick forest
and jungle, and frequent on the whole more open country
such as is supplied by scrub-jungle or rocky hills with bushes,
caves, and crevices for shelter. Many sportsmen have testified
to the obliterative effect of their coloration in foliage or grass,
and Col. Stockley noticed that a leopard that proved to be
of the normal colour when shot looked quite grey by daylight,
and was difficult to see at a comparatively short distance
when passing over the exposed face of a cliff in the Salt Range.
Although unable to kill such large prey as tigers, leopards
similarly prey upon almost all kinds of animals worth killing
that they can seize with safety, ranging in size, as Blanford
tersely expressed it, from an ox to a sparrow. A list of the
animals they feed on would include all the comparatively
defenceless hoofed denizens of India, as well as monkeys,
jackals, porcupines, peafowl, and junglefowl, and to these
Dunbar Brander adds lizards, snakes, and crabs ; but accord-
ing to the same observer they leave alone adult bull nylghaie,
and stag sambhar and swamp-deer (barasingha). On the
other hand, while confirming their avoidance of sambhar,
Col. A. E. Ward states that in Kashmir they easily overcome
fhll-grown stag hangul when hampered by deep snow. Often
238 FELIDiE.
frequenting the neighbourhood of villages and harbouring
in the crops or scrub nearby, leopards may be great pests
to the inhabitants by preying upon calves, goats, sheep,
donkeys, ponies, and other domesticated livestock. Of dogs
they seem to be particularly fond and, being more venture-
some than tigers, will enter bungalows and take them from
under the very noses of their owners. Monkeys may be
seized when feeding or drinking on the ground ; but langurs
up in the trees, where they would be safe if they had the
sense to stay there, allow themselves to be caught either by
falling or jumping to the ground in panic when the leopard
is after them. This has been recorded both by Dunbar Brander
in India and by Phillips in Ceylon.
In relation to their size leopards are as powerful as tigers,
and are at the same time more active and lithe in all ther
movements. Their strength has often been shown in connec-
tion with their not uncommon habit of stowing away their
prey in some place of safety. Of this two instances may be
cited. Dunbar Brander found the carcase of a swamp-deer
lodged by a leopard in a tree, and the leopard above referred
to, watched by Ool. Stockley in the Salt Range, sprang to
a rock 10 ft. above it, carrying a ewe oorial held by the chest
in its mouth. Dunbar Brander thinks the taking of prey into
trees is due to fear of being robbed of it by tigers. To this
might be added hyaenas, which the same author has seen
drive leopards from their kill. But the habit is practised,
according to Phillips, in Ceylon, where there are no tigers or
hyaenas. Hence the habit in India is probably actuated by
the wish to save the prey, or what may be left over for a second
meal, from jackals as much as from tigers or hysenas ; and
jackals are probably the sole factor in Ceylon. Like tigers,
leopards disregard the state of putrefaction of a carcase.
When leopards have found out the ease with which man
can be killed, they may become confirmed " man-eaters."
Once the habit is adopted they become a worse menace even
than tigers, on account of their greater boldness and activity.
They have been known to enter huts and tents at night after
their victims and to take men from " machans " set up in
crop-fields. Blanford quotes from Sterndale and Forsyth
the case of a man-eater at Seon which is alleged to have killed
200 human beings in two years before it was shot. In Ceylon,
according to Phillips, there are very few records of man-
eating leopards, possibly, he suggests, because game may be
more plentiful than in India. There was one well-known case
m the island of a leopard that used to lie in wait for passers-by
on a main road. It was happily shot, however, before it had
accounted for more than twelve human lives.
PLATE XVI.
MAMMALIA.
Skull of Snow-Leopard.
A. Side view. B. " Waist." C. Nasals. D. Left auditory bulla, the dotted
line showing position of partition. E. The same of Common Leopard.
rsciA. 23!>
The attacking roar of a leopard i.s tolerably similar to that
of a tiger ; but the ordinary roar, uttered as a mating call,
is unmistakably different. It is not so loud and resonant,
and consists of a series of about three or four harsh, coughing
barks, with an interval of a second or so between each, and
aptly compared by many to the sound made by the strokes
of a coarse saw through hard, vibrating wood.
The period of gestation, supposed by Blanford to be about
fifteen weeks and by Dunbar Brander twelve, was observed
at Whipsnade. as I learn from Dr. Vevers, to be thirteen weeks
as nearly as possible — to be precise, ninety-two days — in an
Indian leopardess*. As in the lion, but unlike the tiger, the
male stays with the female up to and after the birth of the
cubs which, from two to four usually in number, are born in
some sheltered spot like a cave or overhanging boulder, but
sometimes, according to Dunbar Brander. in a porcupine's
burrow.
Genus UNCI A Gray.
I'ncia, Gray. Ann. Hag. Nat. Hist. (2) xiv, p. 394. 18.5-i ; Pot-ark.
Ann. Hag. Nat. Hist. (8) xviii, p. 306. 1'JltS ; id., Jouru. Bomb.
Nat. Hist. Soe. xxxiv, p. 330, 1930.
Type and only species of the genus, Felis undo, Schreb.
Distribution as below, under the species.
Distinguished from PantTiera by the shape and some other
structural characters of the skull, which is short and wide,
with the orbits high, the frontal interorbital region being
strongly elevated so that the upper profile of the cranial and
facial portions is respectively steeply inclined backwards
and forwards, the facial portion is markedly concave where the
nasals abut against the frontals, and the basicranial axis
slopes noticeably forwards and \ipwards to meet the basifacial
axis at an obtuse angle ; the nasals are short and broad :
their anterior width nearly equalling their median length,
the maxilla is narrow above and the partition of the bulla is
remote from the auditory orifice, so that the two chambers
are subequal in size, with the anterior half of the inner chamber
very narrow ; the occiput has a deep depression dose to the
bulla on each side ; the mandible has the lower edge straight,
the chin nearly rectangular, with the alveolus of the canine
only slightly elevated, so that the post-canine space is moderate
in size.
The skull of the Snow-Leopard differs far more from the skulls
of the lion, tiger, and leopard than these differ from each other.
* In an African specimen, from an unknown locality, it was, curiously
enough. 102 days or 14-& weeks.
240 JTELHUB.
24. Uneia uneia Sehreber. The Snow-Leopard or Ounce.
Felis uneia, Sehreber*, Saugeth. iii, pi. C, 1775, p. 586, 1777 ; and
of most subsequent authors, including Blanford.
Felis irbis, Ehrenberg, Ann. Soi. Nat. xxi, pp. 394 and 410, 1830.
Uneia irbis, Gray, Ann. Mag. Nat. Hist, xiv, p. 394, 1854.
Felis uncioides, Horsfield, Ann. Mag. Nat. Hist. (2) xvi, p. 105,
1855.
TJncia uneia, Pooock, Journ. Bomb. Nat. Hist. Soc. xxxiv, p. 331,
1930.
Vernacular. — Ikar, Zig, Sachak, Sah (Tibetan Bhotia) ;
Bharal he (hills N. of Simla) ; Thurwdgh (Kunawar) ; Stian and
Safed Gheetah (Tibetan) ; Burhel Haye=Burhel Killer (east
of Kumaun), according to Col. A. E. Ward.
Locality of the type of uneia, unknown ; of irbis, the Altai ;
of uncioides, Nepal.
Distribution. — Central Asia, the Altai, and Tibet ; British
India, from Kashmir apparently to the eastern end of the
Himalayas t-
Coat in winter on the upper body thick, tufted, broken, and
from 40 to 50 mm. (2 in.) long, about twice as long on the
belly, but shorter and smoother on the shoulders and nape,
and quite short and smooth on the head, apart from the
cheeks, where it may form a longish, thick fringe ; it is also
short on the limbs, but the tail is uniformly bushy throughout.
General colour of the upper side grey with a cream or buff
tinge, the buff brighter in some skins than others, the individual
hairs being buffish-grey with a black tip and the underwool
dark grey or brownish-grey ; the head is clearer grey, with
some white about the eyes and in the upper lip. The backs
of the ears are black, with a conspicuous white patch. The
pattern consists of solid black spots on the crown, of streaky
spots on the cheeks, but on the nape and shoulders they
are larger and begin to form rosettes ; on the body generally
they typically form large, irregular rosettes, with dark grey
* Xote on the synonymy. — Sehreber gave the name Felis uneia to
a specimen from an unknown locality figured and described by Buffon
as " L'Once " (Hist. Nat. is, pi. 13, 1761). The name irbis was sub-
stituted by Ehrenberg because he disapproved of the use of onca for the
Jaguar and uneia for the Snow-Leopard. The available material is at
present insufficient for the separation of the Himalayan Snow-Leopard
from the one inhabiting the Altai. Hence uncioides is regarded as
a synonym of uneia, although, it will have to be employed if the British
Indian form proves racially distinguishable from the Central Asiatic
form.
t Records of the Snow-Leopard from the Caucasus, Asia Minor,
and Persia, owing to confusion between it and the panther or common
leopard of those countries, were discussed in my paper in 1930, and
need not be repeated. Similar records of it from Manchuria and
Amurland were also no doubt due to skins of the leopard of those parts
-of Asia being mistaken, for it (p. 234).
UNCIA. 241
centres, and two or three sometimes coalesce, forming elongated
blotches, with a tendency to run into longitudinal chains,
especially on the hind back and loins dorsally, where there is
an irregular median black spinal stripe ; these dorsal bands
on the loins may be traceable on the base of the tail, but
posteriorly they break up into separate blotches ; the legs
below the thigh and shoulders are marked with solid black
spots which become smaller inferiorly ; the fore paws are
typically spotted, but the hind paws are spotless, and there is
a mat of dirty whitish hair on the back of the hind leg up to
the hock. Except for some solid spots on the belly, the under-
side is white from the chin backwards, and the underside of
the tail is white throughout. When the coat is loose and tufted
the rosettes become broken up and the pattern obscured.
This is illustrated by a skin from Gilgit, 6,500 ft. (Col. W. F. R.
Trevelyan), dated March 25th, and in full, long winter coat.
In a second specimen, received from the same donor, from the
Ghazi Area, Gilgit, February 1st, the pattern is normally
well defined, the coat being somewhat shorter. The only
other British Indian skins examined are one from Srmagar,
Kashmir (Christie), with shortish coat and distinct pattern,
one from " Kashmir " (Mrs. Entwistle), and one from Simtola
in the Himalayas, which has the coat tufted and the pattern
obscure, irregularly mottled with black and grey. Tibetan
skins are indistinguishable.
The newly-born cubs, as shown by three from Gyantze,
Tibet (Col. F. M. Bailey), are darker than adults, being
brownish-grey in general hue, with the head and fore quarters
confusedly and sparsely spotted, the flanks and thighs marked
with larger spots with paler centres, and the spinal area of the
loins marked with three conspicuous, close-set longitudinal
black stripes.
The size of the Ounce given by Blanford, as head and body
4 ft. 4 in. and tail 3 ft., was apparently copied from Stemdale ;
but I feel sure these dimensions were taken from a flat skin
with the head and body stretched. The total length is the same,
at all events, as in the flat skin of a $ sent by Col. Trevelyan
from Gilgit. Both considerably exceed the dimensions of
three specimens, measured in the flesh and recorded by Col.
A. E. Ward, which are entered below. In the absence of skulls
it is, however, not possible to affirm that Ward's animals were
absolutely full-sized.
Head and
Locality and sex. body. Tail. Total.
ft. in. ft. in. ft. in.
Kashmir; <? 3 8 3 6 8
Baltistan ; <? 3 5 3 6 r>
Baltistan; ? 3 3 2 9 6
VOL. I. K
242
FEUDJE.
According to this table the Snow-Leopard is a little smaller
than the leopard or panther and has the tail relatively longer.
The measurements recorded in the table below show that
the skull of the Snow-Leopard is shorter than that of the
common Indian leopard (see tables, pp. 230 & 236). The skull
of the (J, for example, is about the average length of the skull
of the leopardess. But the postorbital and interorbital widths
are relatively wider. The first three were collected by
Col. W. I\ R. Trevelyan, the fourth by Dr. T. P. Longstaff.
Skull-measurements (
in Trim.) of the Snow- Leopard.
Locality and sex.
i
.2 S
■SIS'
o
si
3
S3
CO >
a
1
1*
II
a*
s
M
"3.3
s.
s
Gilgit, Hunza ; ad. (J . .
185
171
126
46
44
50
126
24
18
Gilgit, Gozi GHzar ; ad. £
175
160
119
46
m
47
119
22£
18
Gilgit, Ish. Koman ; ad. $
171
158
125
47
41
48
119
22
17
GUgit, Bmji ; ad. 9 J . . 168
155
114
47
36
46
115
24
17
Habits. — Near or within British Indian limits the Snow-
Leopard, according to Burrard, ranges from the Hindu Kush
throughout the Himalayas. It is abundant in the Zaskar
Range and occurs in limited numbers a few miles along the
Dauladhar and Pir Panjal Ranges from their junction with the
main Himalayan chain, being everywhere apparently commoner
on the northern than on the southern slopes. It is not found
in scrub or jungle, but on rocky hill-sides above the tree-line
at altitudes as low as 6,000 ft. in winter and as high as 12,000
-or 13,000 ft. in Hummer. That its colour is adapted to that
of its environment no one doubts, and the high position of its
orbits enables it to peer over the edge of a rock to reconnoitre
for prey and locate it with the least possible exposure of the
head before creeping forth to stalk it. It lives on ibex,
bharal, and other wild goats and sheep as well as on musk-
deer, hares, marmots, picas probably, and such game-birds
as the monal pheasant. In Kashmir, according to Ward,
when the upland grazing grounds are open to the flocks it
takes the goats, sheep, and occasionally ponies of the herds-
men. That it may prove a costly nuisance to livestock
owners is shown by Col. R. H. Percy's account of a pair that
for long harassed the farm of the Moravian Mission at Kailing
in Lahul. It lies up most of the day and starts to hunt about
sundown, its generally nocturnal habits, combined with the
MAMMALIA.
PLATE XVIII.
Left. — Flat skin of cub of Snow-Leopard.
Right.— Made-up skin of cub of Common Leopard.
FELINJE. 243
inaccessibility of its home, being the reason why it is so seldom
seen by English sportsmen.
Of its breeding habits nothing seems to have been recorded ;
but the discovery of three cubs at Gyantse, Tibet, by Col. F. M.
Bailey suggests that the litter consists usually of from two
to four.
Skins of Snow-Leopards are greatly in demand in the fur
trade, and the beast is usually secured by trapping in deep
•pits wider at the bottom than at the top and baited with kids.
Subfamily FELINE.
The suspender (suspensorium) of the hyoid of the normal
mammalian type and consisting of a chain of three bones
jointed end to end and holding the larynx close to the base
of the skull, thus restricting its movement. The tips of the
digits of both fore and hind feet furnished at least with a single
•cutaneous lobe protecting the retracted claw on the inner side
•of the second and third digits and on the outer side of the
fourth and fifth ; frequently there is an additional lobe on the
•opposite sides, constituting a complete sheath to the claw, as
in the Pantherinae.
The difference between this subfamily and the Pantherinse
in the structure of the hyoidean suspensorium is associated
with a difference in the voice. The sexual call is not a deep-
toned roar, but a higher-pitched sound, varying according to
the species, where it has been recorded, the most familiarly
known being that of the common House- Cat ; and, as in the
■common House-Cat, pleasure or content is expressed, at
least in some widely divergent species, like the American
Puma and the African Serval, in which it has been heard,
by " purring."
The Fehnse are more generalized in structure than the
Pantherinae and Acinonychinae, and are ancestral to both.
In addition to the genera and species inhabiting British
India, the subfamily contains a large number of differen
kinds found in other parts of Asia, in Europe, Africa, and
America. In the south-eastern parts of Asia there are two
peculiar forms, the Flat-headed Cat {Ictailurus) of the Malay
Peninsula and Sumatra and the Bay-Cat of Borneo (Badio-
folis), which are generically distinct from those of British
India, but the rest of the Asiatic Felinse, as well as the European
and African species with one exception, the Serval {LeptaUurus),
belong to the genera Felis, Caracal, Lynx, and Profelis, repre-
sented in the British Indian fauna. All the heterogeneous
series of Felidaa inhabiting America, except tht> Jaguar (Pan-
thera onca), belong to the Felinse. The largest, dominant,
»2
244 EELIDJE.
and most widely distributed form is the Puma or Cougar
( Puma) , which, has no close kinship with any Old World species * .
Other well-known forms, such as the Ocelots (Leopardus),
and the Eyra or Jaguarondi (Herpailurus), come nearest
to the Leopard-Cats (Prionailurus) of India and Eastern
Asia ; but the only genus which is common to the eastern and
western hemispheres is Lynx, the most northern of all the
genera of lelidse. This genus has a bearing upon the value
attached to the characters I have adopted in this volume and
elsewhere for the division of the Felinse into a number of
distinct genera. It is a species which, apart from the Cheetah
or Hunting Leopard, has been most frequently separated as
a distinct genus from Felis by authors who assigned to the
latter all the remaining species of the family, including even
the Lion, Tiger, and their allies. Yet the Lynx is, beyond
question in my opinion, much more closely akin to typical
Felis than are the Marbled Cat, Leopard-Cat, and others, being
connected with it by such more or less intermediate forms as
the Caracal and Jungle-Cat, whereas there is no such con-
nection between typical Felis and the Leopard-Cat. Hence the
latter must logically be given generic status if that rank is
granted to the Lynx.
In the following analytical keys for the determination of
the genera, the second, based on the skulls, contains the
characters on which they are mainly distinguished. The
first is an artificial arrangement in the sense that Pallas's
Cat (Otocolobus) comes under the section containing species
with which it has no close kinship, the structure of its skull
showing it to be a highly specialized cat related to Felis.
Setting Otocolobus aside, the shape of the ear readily dis-
tinguishes Felis, Caracal, and Lynx from Neofelis, Pardofelis,
Profelis, and Prionailurus ; but the differences enumerated
for separating the four genera included under 6 and the
three included under a' are of specific rather than generic
importance.
Key to the British Indian Genera of Felinas based on the
more obvious external features,
a. Ears low, rounded at summit, usually with
white patch.
b. Ears normally set ; body-pattern funda-
mentally consisting of spots or rosettes
on the flanks, sometimes coalescing to
form large " clouded " blotches, and of
longitudinal stripes on the head and
back, occasionally secondarily lost.
* There is an unmistakable likeness between the pattern of the
newly-born cubs of this species and of the Snow-Leopard or Ounce,
which may point to kinship between them, despite the difference in
the hyoidean apparatus.
FEUXiE. 245
c. Tail not tapering, very long, nearly as
long as the head and body and about
four times as long as the hind foot ;
pattern " clouded " or " marbled."
d. Head long, not rounded ; size large . Neoeeus Gray, p. 247 .
d'. Head short and rounded ; size [p. 253
small Pabdofeus Severtz.,
c'. Tail tapering typically, shorter, at most
a little more than half the length of
the head and body, only occasionally
as much as three times the length
of the hind foot ; pattern of spots
or rosettes, not " clouded " or
•' marbled."
e. Tail conspicuously pale at the tip
beneath ; no conspicuous white [p. 258.
patch on ear Pbokei.is Severtz.,
e'. Tail not conspicuously pale at tip
beneath ; a pale, usually white [Severtz., p. 265.
patch on back of ear Prionaxlubus
b'. Ears set laterally, separated by a very
wide area of the crown ; body -pattern
consisting at most of a few abbreviated [p. 315.
vertical stripes on the loins Otocqlobus Brandt,
a'. Ears high, triangular, narrowed towards the
point, often tufted.
/. At most a small tuft on the ears ; pattern
consisting mostly of transverse stripes
on the sides, sometimes evanescent in
the adult, or of spots which, at least
behind the fore limb, tend to run into
vertical stripes Feus Linn., p. 285.
f. A long erect tuft on the ear, nearly as long
as the height of the ear ; pattern, when
present, spotted, not vertically lineate.
g. Pattern lost on the back and sides ;
tail moderately long, reaching the
hocks ; no cheek-ruff Cabacal Gray, p. 306.
g'. Pattern manifest at least at certain
seasons ; tail short, reaching about
half-way to the hocks ; a distinct
cheek-ruff or fringe Lynx Kerr, p. 310.
Key to the British Indian Genera of Felines based •upon
Cranial Characters.
a. Skull long and low, with short, blunt, widely
separated frontal and zygomatic post-
orbital processes, very long canine teeth,
and large posteanine space Xeofelis Gray, p.. 247.
a'. Skull with long, well-developed postorbital
processes and normal canines and post-
canine space.
b. Outer chamber of auditory bulla small,
the partition lising close to the orifice ;
facial portion of skull shorter, tip of
frontal postorbital process in front of
middle of skull (only slightly in Pardo-
felis).
246 EELID-ffi.
c. Skull high, wide, and short, mastoid
■width more than half condylobasal
length ; posterior edge of palate
deeply notched laterally, mesoptery-
goid fossa lanceolate or ovate in [p. 253.
front Pardofelis Severtz.,
c'. Skull lower, narrower and longer,
mastoid width less than half the
condylobasal length ; a shallow
• emargination on each side of hinder
edge of palate ; mesopterygoid fossa
truncated, with median notch, in
front.
d. Nasal branch of premaxilla slender ;
maxillse not expanded above, and
the nasals not compressed, post-
orbital bar never complete ; ex-
ternal pterygoid crest well [p. 258.
developed Profelis Severtz.,
d'. Nasal branch of premaxilla broad ;
maxillse expanded above and nasals
compressed : postorbital bar com-
plete, usually only in old skulls of [Severtz., p. 265.
one species Prionailurtts
li'. Outer chamber of bulla comparatively or
very large, the partition rising some
distance from the orifice ; tip of frontal
postorbital process about the middle of
the length of the skull.
c Outer chamber of bulla smaller, inner
broad in front*, skull narrower,
mastoid width seldom as much as half
the condylobasal length ; lower rim
of orbit below the level of the summit
of the anterior nares.
/. Nasal branch of premaxilla broad,
posterior edge of palate deeply
notched laterally ; postorbital pro-
cesses broad and ligulate ; meso-
pterygoid fossa with median angular
point in front ; small upper pm
typically retained Felis Linn., p. 285.
/'. Nasal braneh of premaxilla slender
and long, hinder edge of palate not
conspicuously notched laterally ;
mesopterygoid fossa typically with
median notch in front ; small upper
pm absent or early shed.
g. External pterygoid crest a posterior
triangular spike, as in Felis Caracal Gray, p. 30&
g'. External pterygoid crest obsolete. Lynx Kerr, p. 310.
t'. Outer chamber of bulla very large,
inner chamber narrow in front ; skull
broader, mastoid width much more
than half condylobasal length, lower
rim of orbit about on a level with [p. 315
upper ed ge of anter ior nares Otooolobus Brandt,
* In one or two desert species of Felis outside the limits of the British
Indian fauna the outer chamber is very large and the inner is narrowed
in front.
NEOEELIS.
247
Genus NEOFELIS Gray. The Clouded Leopard.
Neofelis, Gray, Proc. Zool. Soc. 1867, p. 265 ; Poeoek, Ann. Mag.
Nat. Hist. (8) xx, p. 343, 1917.
Type of the genus, Felis onacrocdis Horsfield* (=diardi
Cuv.).
Distribution. — From Nepal and Sikkim eastward to S. China,
Hainan, and Formosa, and southward to Burma, Annam, the
Malay Peninsula, Sumatra, Java, and Borneo.
One of the largest of the Oriental Felinse, with the ears
rounded, the head, body, and tail long, the legs short and
stout with broad paws, in which the four main digits are
fully webbed and provided with double claw-sheaths both on
the fore and hind foot ; but chiefly characterized by cranial
and dental peculiarities associated mainly with the fore part
Fig. 65. — Skull of Clouded Leopard (Neofelis nebuiosa), about half
natural size. (From Blanford.)
of the jaws, which carry exceptionally long, piercing canine
teeth, the upper being about three times as long as its basal
width at the socket ; the chin is prominent, flattish, and nearly
vertical in front, nearly rectangular below, and the alveolar
edge of its incisors is raised well above the alveolus or socket
of its canine, so that, when the mouth is shut, the upper jaw-
is uptilted and the alveolar lines of the upper and lower cheek-
teeth are widely divergent in front, producing a large post-
canine space which gives deep penetration to the canine
teeth. The entire skull is long and low, with lightly convex
upper outline, a strong sagittal crest, salient zygomatic
* Zool. Joura. i, p. 542, pi. 21, 1825. F. macroeelis, from Bencoolen,
Sumatra, is a subspecies of nebuiosa. It was the first of the two forms
cited by Gray as representing Neqfelw, and is here selected as the type
of the genus. The second was Leopardm brachyurus Swinhoe, from
Formosa (Proc. Zool. Soc. 1892, p. 352).
248 FELID.&.
arch.es, narrow postorbital and interorbital areas, short,
widely separated frontal and malar postorbital processes,
broad, nearly parallel-sided nasals, slender premaxillse, anterior
nares sloped at about an angle of 45° to the cranio-facial axis,
and the outer chamber of the auditory bulla small as in all
forest cats.
Apart from the special modifications of the fore part of the
jaws and the long canine teeth, which are larger in the male
than the female, the skull recalls the long, low skull of a small
male panther, more particularly in its short, blunt, frontal
postorbital processes and the wide separation between them
and the corresponding processes rising from the zygomatic
arches. It is very distinct from the skulls of the rest of the
Pelinse found in British India and elsewhere.
25. Neofelis nebulosa Griffith.
Felis nebulosa, Griffith, Descr. Vert. Anim. Cam. p. 37, pi., 1821 ;
id., Anim. Kingd. ii, pp. 449-51, pi., 1827 ; and of all recent
authors.
Locality of the type, Canton.
Distribution. — As under the genus.
Ground-colour dark grey or ochreous, often largely obliterated
by the black and dark dusky-grey blotched pattern. Crown
spotted, with a larger patch above the eye, a stripe running
from the corner of the eye backwards over the cheek, another
from the corner of the mouth along the sides of the neck,
one from behind the black ears and two on eacb side along
the nape to the shoulders, sometimes partly fused or broken
up ; similar stripes or elongated blotches continued down the
spine, commonly forming a single median stripe on the loins
and passing as such on to the basal part of the tail ; on the
sides of the shoulders two large blotches of dark dusky-grey
hairs, each emphasized posteriorly by a black stripe which
passes on to the fore leg and breaks up into irregular spots ;
the flanks typically somewhat similarly marked by large
dusky-grey irregular blotches bordered behind by long,
oblique, irregularly curved or looped stripes and in front by
much fainter spots or lines ; on the hind quarters the blotches
are smaller, frequently almost surrounded by nearly horse-
shoe-shaped stripes. These blotches, yielding the clouded
pattern suggesting the popular English name of this
species, are often so large and close set that the intervening
spaces look like pale irregular stripes on a dark ground ; but
sometimes the blotches take the form of large, somewhat
\jaguarine" rosettes, dusky patches surrounded by a black
ring as wide m front as behind. The hind leg is spotted like
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NEOFELIS. 249
the front, and the tail is marked by large, irregular, paired
spots, often coalescing to give an annuliform pattern to the
organ. The front of the upper lip, the underside from the
chin, and the inner sides of the limbs are white, except for two
black collars on the throat and some large black spots on the
chest behind the fore legs and on the inner surface of the
limbs*.
The original figure of the type of this species represented
the ground-colour as brightish ochreous and the pattern as
consisting on the sides of the body of long, widely spaced,
conspicuous black stripes forming the posterior borders of
large dusky patches defined in front by narrower stripes or
spots, the combination suggesting the name Tortoiseshell
Tiger for the species. Canton was probably the port of
shipment of this animal, which was exhibited in the so-called
'' Exeter Change " menagerie. Another, and better drawn,
illustration of it by Landseer, reproduced by Griffith in 1827,
is not so bright a yellow as the first. That the Clouded
Leopards of the Chinese area are at least on the average
lighter and brighter than those of the Himalayas and the
Sunda Islands is borne out by other skins. In the British
Museum there is a native skin, picked up by Delacour and
Lowe at Laoboa, Annam, which is the brightest of all, and
agrees tolerably closely both in tint and pattern with Griffith's
original figure. Also there are four skins from Formosa,
referable to the form named brachyurus by Swinhoe f , which are
tawnier, paler, and less grey than the Himalayan skins I have
seen ; and B. Howell referred to a specimen from Hainan
as " ochreous."
Another synonym of this race is Felis (Necfelis) meZKMatschie
(Arch. Naturg. 1922, pt. 10, pp. 19 and 35), based on a skin
purchased in Canton ; and G. M. Allen has recorded it from
Pokien (Amer. Mus. Novit. no 360, p. 11, 1929).
Apparently a second race is the form described as Felis
* Discussing the alleged but unsubstantiated reports of the occurrence
of true leopards in Sumatra, Mr. E. Jacobson stated his belief that all
the so-called black leopards said to have been shot or seen in that
island were black examples of N. nebulosa { Journ. Fed. Mai. St. Mus. x,
p. 238, 1921).
t This is, I believe, a synonym of nebulosa. Swinhoe described
it as a " species " on the evidenee of the shortness of the tail in the
type, that organ being only a little over half the length of the head
and body of the stripped skin. But in. another Formosan specimen
it is over two-thirds the length. The length of the tail is too variable
to be trusted. The flesh-measurements of an adult & ( ? subspecies) from
Chumpawn in Peninsular Siam (Robinson and Kloss) are : — Head and
body 32 in., tail 30| in., hind foot 7 in., the tail being very nearly as
long as the head and body. No other flesh-measured specimens are
Available for record.
250 TEUCDM.
diardi Cuvier ('Ossemens Fossiles,' new ed. iv, p. 437, 1823),
based on a skin from Java. F. macrocdis Horsfleld, from
Sumatra, is, I believe, a synonym of it. In the type of the
latter the tint is darker and greyer than in nebulosa from
the Chinese area, and the pattern on the flanks is more rosette-
like, the blotches being less elongated, smaller, sometimes
nearly circular, with the anterior rim about as thick as the
posterior, and small black spots involved in the blotches
are more in evidence. I provisionally assign to this race
skins from Borneo and Malaya, which, although resembling
Himalayan skins in general colour, are not quite like them in
pattern. The attempt to define these races is, however,
tentative owing to scarcity of material.
25 a. Neofelis nebulosa macroseeloides (Hodgson).
Felis macrocelis, Tickell, Joum. As. Soc. Beng. xii, p. 814, 1843 ;
Blyth, Mamm. Birds Burma, p. 27 (not of Horsfleld, 1825.)
Felis macroseeloides, Hodgson, Proc. Zool. Soc. 1853, p. 38
(spelt macroeeloides as nom. nud., Hodgson, Calc. Journ. Nat.
Hist, iv, p. 286, 1844, and misprinted macroselloid-es, without
description, Proc. Zool. Soe. 1853, p. 192) ; Blyth, Cat. Mamm.
As. Soc. p. 58, 1863 (macroeeloides).
Felis diardi, Blyth, Proc. Zool. Soc. 1863, p. 183 ; Jerdon, Mamm.
Ind. p. 102, 1867 (not of Cuvier, 1823).
Felis nebulosa, Blanford, Mamm. Brit. Ind. p. 72, 1888 ; and of
later writers on the Indian fauna (not of Griffith, 1821).
Vernacular. — Pungmar, Satchuk (Lepeha) ; Zik (Limbu) ;
Rung (Bhotia) ; Lamchitia (Khas tribe, Nepal) ; Thit Kyoung
(Burmese) ; In-cha (Tavoy).
Locality of the type, of macroseeloides, Nepal.
Distribution. — Nepal, Sikkim, and Bhutan; range to the
eastward unknown, possibly Assam and at least parts of Burma,
but no doubt blending with typical nebulosa in Upper Burma.
At Darjeeling, according to W. H. Matthews, it occurs up to
3,000 ft.
Distinguished from typical nebulosa by its generally darker,
greyer, less yellow hue, and from diardi by the more extensive
blotches of its marbled pattern and possibly larger skull.
Hodgson's skins, including the type, from Nepal, have an
ochreous wash over the generally dark grey ground-colour,
but there is hardly a trace of this in another skm from Nepal,
nor in one from Sik k i m .
Tickell described his specimen from the Snowy Range of
Northern Sikkim as " bistre brown or dull clay brown "
above and as " pale, rufous tawny below and on the inside of
the limbs." TMs skin seems to have agreed with Hodgson's
except in the richer hue of the lower side. Blyth, who in
1863 recorded this animal from Nepal, Sikkim, Bhutan, Tibet,
and Upper Assam, referred to two specimens from Sikkim,
one "folvous" the other "grey." Incidentally he also
SEOFELIS. '2")1
mentioned a " remarkably handsome " skin from the Ya-ma-
doung Mountains, between Pegu and Arakan. This vague
description suggests the occurrence of a bright-tinted Clouded
Leopard, like typical nebulosa, in that part of Burma. 1 have
seen only three skins from Burma. There is one from Bhamo
(C. F. Gilbert), which closely resembles the brightest of the
Nepalese specimens. According to the collector's note this skin
was brought in by natives for the reward, and may have come
from over the Chinese border It approaches typical nebulosa
in colour. The remaining two are native skins picked up by
J. M. D. Mackenzie 8 miles west of Toungoo. They are covered
with dusky olivaceous blotches, separated by narrow pale
interspaces, but on account of their immaturity no importance
can be attached to their generally dusky hue because, according
to Blyth, typical maerosceloides is darker and greyer in hue
when young. They may, however, represent the provisionally
admitted southern form, diardi, which apparently occurs
in Peninsular Siam.
No certainly known flesh-measurements of Himalayan
specimens are available, but the following (m English inches),
taken mostly from skins, may be recorded. To these are
added, for comparison, the flesh-measurements of the example
from Peninsular Siam given by Robinson and Kloss : —
Head and Hind
Locality and sex. body. Tail foot.
N. Sikkim (Tickell), maerosceloides ;
? (J 41 34| —
Nepal (Brit. Mus.), maerosceloides ;
? c? 42 36 —
Nepal (Hodgson), maerosceloides ;
oldo 1 37| 29 —
Sikkim (Brit. Mus.), maerosceloides ;
ad. $ 32 23 —
Nepal (Brit. Mus.), maerosceloides ;
ad. 9 27 24 —
Peninsular Siam (Rob. and Kloss) ;
? drardi ; ad. <J 32 30* 7
The weight of Hodgson's old <$, probably measured in the
flesh, was 44| lb. W. H. Matthews recorded two (£<? as
39 and 42J lb. respectively, the larger being t>5J in. in total
length.
Measurements of the skulls from Peninsular Siam, Sumatra,
and N. Borneo are added for comparison with those of maero-
sceloides to support the view that specimens from the Himalayas
have larger skulls than those from the southern countries
assigned to diardi.
Habits. — The Clouded Leopard is everywhere rare, or at all
events seldom seen by European sportsmen, probably because
it inhabits dense forests and is nocturnal. A good deal of the
material in museums is of native origin, and most of the
252
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information about its habits is of native source, as Blanforcl
said.
From what he was told in Sikkim, Tickell reported it an
frequenting the dense jungles, chiefly near river-banks, in
the valleys north of DarjeeUng, and as sometimes visiting
the villages of the Bhotias and Lepchas to prey upon goats
and pigs. It is savage and wary and, when wounded, may
turn savagely on its assailants. Tickell, not without reason,
ridiculed Raines's statement that in Sumatra the Clouded
Leopard subsists by preying upon the birds it catches in trees
and on the poultry of the villagers. From the deep penetration
of its bite, attested by the long canines and large space behind
them, not to mention its powerful build, this eat is obviously
adapted for preying upon herbivorous mammals of considerable
bulk ; and Swinhoe says that in Formosa it commits great
havoc among deer, no doubt the Formosan Sika. He adds,
however, that it never attacks man unless provoked. A case,
nevertheless, was recorded by A. Brownlow (Journ. Bomb.
Nat. Hist. Soc. xxxii, p. 789, 1932) at Tavoy of a specimen,
which had killed several head of cattle, stalking and attacking
a native boy who killed the beast by splitting its skull with
his knife. But, despite their savage disposition, some Clouded
Leopards are capable of being tamed. A full-grown specimen
that lived many years in the Zoological Gardens, London,
would let its keeper do anything with it. Nothing appears to
be known about the breeding habits.
Genus PARDOFELIS Severtzow.
Pardofdia, Severtzow. Rev. Mag. Zool. (2) x, p. 387, 1858 ,- Poeoek,
Ann. Mag. Nat. Hist. (8) xx, p. 339, 1917 (in part) ; id., Proe.
Zool. Soc. 1932, p. 742.
Catolynx, Gray, Proc. Zool. Soc. 1867, p. 267 (not of Severtzow,
1858).
Type-species of Pardofelis and of Catolynx (Gray), Felis
marmorata Martin.
Distribution. — The Eastepjst Himalayas, Nepal, Sikkim,
Assam, Bubma ; the Malay Peninsula, Sumatra, Borneo, and
Annam, and no doubt in suitable localities in the intervening
countries.
A comparatively small species, about the size of a domestic
cat, but with the ears rounded and the tail very long, about
as long as the head and body, or longer, and over four times
as long as the hind foot. The coat long, full, and soft ; the
ground-colour varying from tawny or brownish-grey to rich
ochreous-brown above, greyish to buff below. The pattern is
individually variable, always consisting of blackish stripes
on the head, neck, and back, of more or less black-edged
blotches on the flanks, and of solid spots on the tail, limbs.
254
eeled.®.
a,nd underside, but the blotches on the flanks are very inconstant
in size ; in their simplest form they are comparatively small,
numerous, and tolerably widely spaced ; frequently, however,
they fuse to form large dark areas defined by narrowish pale
lines, giving the impression of a dark brownish cat with pale
stripes. This phase constitutes the so-called " marbled "
pattern ; but th.e simpler phase is clearly derivable from the
pattern of some of the handsomer examples of the Leopard-Cat,
and, like the latter, the Marbled Cat often has white on the chin
and upper lip, and always the back of the ear, and the tail
uniformly coloured beneath and not conspicuously white at
the end as in Temminek's Cat. The pupil of the eye is circular
when contracted.
The skull differs from that of Prionailurus in the sum of
a number of characters. It is short, broad, strongly convex
Fig. 66.— Skull of the type of the Himalayan Marbled Cat (Pardofelis
tnarmorata charltoni), nat. size. From Darjeeling.
in dorsal profile, not comparatively long and low ; the nasal
branch of the premaxilla is thin, not expanded, the summit
of the muzzle is not compressed above, the maxilla is not
expanded where it abuts against the nasal bone, and develops
no excrescence outside the suborbital foramen ; the meso-
pterygoid fossa is lanceolate, not transversely truncated in
front, and its side margins are thicker ; the occipital region
is much wider as compared with its height, the width across
the mastoids exceeding half the condylobasal length of the
skull, these processes projecting farther beyond the auditory
orifice, and there is a wider shelf-like ridge above this orifice ;
the occipital crest is less steeply inclined, less emarginate
laterally and, like the posterior end of the sagittal crest,
weaker : the mandible has the chin less sloped, more vertical,
and the coronoid higher, with its summit narrower and pro-
jecting farther backwards.
PLATE XX.
MAMMALIA.
Marbled Cat (Por«fofe//s mormorato). (Adapted from a painting by C. E. Swan.)
Rusty Spotted Cat (Prhnailurus rubi S ino S us).
PAEDOFELIS. 23.5
Other cranial characters are the shortness of the nasal bone*,
the wide suborbital portion of the zygomatic arches, the com-
paratively early fusion of the malar and frontal postorbital
processes completing the orbital ring, the wide separation
of the temporal ridges to form a long tongue-shaped area, and
their union, at least in some old males, to give rise to a low-
sagittal crest.
One of the peculiarities of the Marbled Cat is the profound
difference between its skull and that of the Clouded Leopard,
associated with several resemblances in external characters,
such as the feet, which have deep interdigital webs and complete
and double claw-sheaths on all the digits, the long tail, and the
colour and pattern, all probably correlated with arboreal
forest life. The tail, however, is even longer in the Marbled
Cat and the head much shorter and rounder.
Only one species of the genus is known.
26. Pardofelis marmorata (Martin). The Marbled Cat.
Fells diardii, Jardine, Nat. Libr. Mamm. ii, Felirue, pp. 221 and 271,
pis. 21 and 22, 1834 (not Felis diardi Cuv.).
Felis marmorata, Martin, Proc. Zool. Soc. 1836, p. 107 ; and of most
subsequent authors, including Blanford.
Pardofelis marmorata, Pocock, Proc. Zool. Soe. 1932, p. 744.
locality of the type of diardii (Jardine), Java ; of marmorata,
Sumatra*.
Distribution. — As under the genus.
General characters as above, under the genus.
In the typical form of this race, P. ■marmorata marmorata,
from the Malay Peninsula, Sumatra, and Borneo, the coat is
moderately long and thick and the tail moderately bushy ;
the general colour is comparatively dull greyish- brown or
brown, without rich ochreous wash on the flanks and back,
and the pattern is obscure or conspicuous, varying in the size
a,nd confluence of the rosettes as described above.
The following three adult examples, measured in the flesh
(in English inches), give reliable dimensions f : —
Head and Hind
Locality. body. Tail. foot.
Palembang, Sumatra ; ad 18 19| 4f
Babu Tiga, Malacca, ad 20f 2l| 4f
Barito River, S. Borneo, ad 19 18§ 4J-
* Martin's specimen was recorded from Sumatra or Java, but Sumatra
was selected as the type-locality by Bobinson and KIoss (Journ. Fed.
Mai. St. vii, p. 261, 1919. The existence of the species in Java is
doubtful.
t The first two specimens were collected and measured by Robinson
and Kloss. The third, collected hy W. Frost, was measured at the
British Museum. Since the skull of Martin's type from Sumatra is
small and immature, his measurement of the head and body as 23 in.
and the tail 15i must be taken -with reserve.
256 FEUDJB.
The occurrence of this race in the Malay Peninsula suggests
the possibility of its extension into Tenasserim and, perhaps,
other parts of Lower Burma.
26 a. Pardofelis marmorata eharltoni (Gray). The Himalayan
Marbled Cat.
Felis eharltoni, Oav, Ann. Mag. Nat. Hist. (1) xviii, p. 211,
1846.
Oatolynx eharltoni, Gray, Proe. Zool. Soc. 1867, p. 268.
Felis ogilbii, Hodgson, Calo. Journ. Nat. Hist. (8) p. 44, 1847*.
Leopardus dosul and ? duvaucelli, Hodgson, Cat. Hamm. Nepal,
ed. 2, p. 3, 1863 (no description).
Felis marmorata^ Blanford, and other writers on the Indian fauna
(in part, not of Martin).
Pardojelis niarmorata eharltoni, Poooek, Proe. Zool. Soo. 1932,
p. 746.
Vernacular. — Dosal (Lepcha) ; SiJcmar (Bhotia).
Locality of the type of eharltoni, Darjeeling ; of ogilbii and
dosul, Sikkim.
Distribution. — Nepal, Sikkim, Assam, and Upper Burma ;
range to the east and south of Upper Burma unknown.
Distinguished from the typical race by the much thicker
and longer coat and rich oehreous-brown colour, the pattern
less mottled, consisting of large oehreous-brown areas edged
with black behind and sometimes in front, and separated
by a few greyish-buff wavy lines ; the legs the same rich
tint as the body, heavily spotted with black basally ; tail
not so rich, duskier than the body, with the pattern somewhat
obscurely defined.
No flesh-measurements of this race are available , but
they are probably approximately the same as those of the typical
race, since the skulls of the two are alike in size and other
particulars.
Apart from Horsfield's statement that this cat is found
" in the hilly regions of Nepal " f next to nothing is known
about it (Proe. Zool. Soc. 1856, p. 396). There are only three
skins in the British Museum, the type from Sikkim, one
* Felis ogilbii Hodgson has been added to the synonymy of this
race because the dimensions of the skin were given as head and body
18J in., tail 14 in. The tail is shorter in proportion than in the flesh-
measured skins of the typical race quoted above ; but the head and body
in stripped skins is nearly always stretched, and no doubt this was the
ease in the skin in question. But even if it was unstretehed the tail
is much longer in proportion to the head and body than in any known
skin of the Leopard-Cat (Prionailurus bengalensis) . This was overlooked
by Blanford when he identified as the type of ogilbii an unusually red,
artificially-dyed skin of a comparatively short-tailed example of
bengaietmB sent by Hodgson from Sikkim and preserved in the British
Museum. This skin is clearly not the type in question (see p. 272).
t Blanford's assertion that the species had not been recorded from
Nepal is clearly an error.
PARDOFELIS.
•J.51
labelled " Nepal " (Hodgson), and one from. Nam Taniai,
Upper Burma (Kingdon Ward). The last is very richly
coloured, and differs from the type in having the upper lip
and chin ochreous, not white, and the underside washed with
buff. Hodgson's skin is intermediate between the two.
Skull-measurements (in mm.) of Pardofelis marmorata
charltoni.
• _
3
.■§
>> .
J3 .
7S J
•
Locality and sex.
— =P
Sss
m
Bp
3M
!
SH
o
SI
H?
-"•
-"•
^ i °
j "India" ; ad. $ . ..
•J —
—
65
28
13
24
63
13
! "India" ; ad. <J ? ..
. . 92
83
G3
20
13
23
60
ii ; i
9
8
Darjeeling; yg. ad. $">.\ — | S2 61 32 14 231 5S 13 ' 11
„ (type) ; ad. 9 ? 88 : 81 65 30 14 23 3SJ 12 S
The width of the mastoid of these, skulls, in the order of their
arrangement, is 45, 42, 42 mm., in all cases exceeding half the
oondylobasal length. Although the skulls themselves are
Pig. 67. — The Marbled Cat {Felis marmorata). (From Blanford.)
considerably shorter than those of P. bengahnsis horsfieldi
from Almora and the Garo Hills, the mastoid width is greater
Habits. — Except in one or two localities where its natural
habitat has been interfered with by man, the Marbled Cat is essen-
tially a forest species, feeding upon small mammals, like rats
vol. i. s
258 felid^;.
and squirrels, and birds up to the size of pheasants . It is seldom
imported alive for menageries and seldom secured by collectors.
Those working for the Bombay Mammal Survey failed to capture
or to purchase from natives a single specimen in Nepal. Sikkim,
Assam, or Burma*. Robinson and Kloss ascribed the rarity
■with which it is seen to its forest habitat and nocturnal
habits rather than to real scarcity. Hose, however, stated
(Mamm. Borneo, p. 18) that it is found in the " clearings "
in Sarawak, and Frost captured a specimen in a fowl-house
on the Barito River, S. Borneo, in a district which for miles
around and for many years had been cleared of native forest
and planted for rubber and cereals, and the animal was living
on the river cliff, which consisted of rocks overgrown with
scrub and low bush. This is an interesting case of adaptation
to new conditions.
Genus PROPELIS Severtzow.
Profelis, Severtzow, Rev. Mag. Zool. (2) x, p. 386, 1S5S.
Ckrysailurus, id., torn, cit p. 389.
Oatopuma, id., torn. cit. p. 387.
Pyrofelis, Gray, Ann. Mag. Nat. Hist. (4) xiv, p. 354, 1874.
Profelis, Poeoek, Ann. Mag. Nat. Hist. (8) xx, p. 340, 1917; id.,
Proc. Zool. Soc. 1932, p. 752.
Type of Profelis, celidogaster Temm. (—aurata, Temm.) ; of
Chrysailurus, neglecta Gray {—aurata Temm.) ; of Gatopuma,
moormensis Hodgs. (=temminckii Horsf. & Vig.) ; of Pyrofelis,
temminckii Horsf. & Vig.
Distribution. — West and Central African forest region ;
S.E. Asia from Nepal and Southern China southward through
the Malay Peninsula to Sumatra ; ? Java, but not Borneo f.
Resembling the Leopard-Cat (Prionailurus bengalensis) in
its low, rounded ears, in the tail being over half the length
of the head and body, and over twice the length of the hind
foot, and also in the style of its body-pattern, when retained,
but distinguished from the three species of Prionailurus
in external characters by the absence of a distinct white
patch on the ear, by the whiteness of the tail-tip beneath,
and by the characteristic facial pattern described below.
* Two native skins sent by Mackenzie from Toungoo and identified
by the late T. B. Fry as Felis marmorata (Journ. Bomb. Nat. Hist. Soc.
xxx, p. 647, 1929) are young examples of the Clouded Leopard.
f There is no doubt about the kinship between the African and
Asiatic species assigned to this genus. Not only are they similar,
with minor differences, in their skulls, but in their colour phases as
well. In the African Cat (P. aurata) the pattern may be manifest or
obsolete and the colour nearly black, "red" or "grey " ; and since
a specimen in the Zoological Gardens, as I recorded in 1907, changed
from "red" to "grey" the same phenomenon probably occurs in
P. temminckii, although not as yet observed.
PKOFEUS.
259
The skull is distinguished from that of Prionailurus by the
narrow nasal branch of the premaxilla, the less expanded
maxilla where it abuts against the nasals, the less compressed
Fig. 68. — A. Skull of adult cj Fishing Cat (Prionailurus vivarinus)
from Sind. B. Nasal bones of example of the same species from
Nepal. C. Skull of adult $ Temminck's Cat (Profelis temminekii)
from Tejo in the Mishmi Hills. D. Nasal bones of example of the
same species from Myitkyina, Upper Burma.
nasals, more strongly developed pterygoid crest, relatively
wider mastoid width, and by the incompleteness of the orbital
bar at all ages. In this last character it also differs from the
2s
260 FEU.DM.
skull of Pardofelis ; it is also much narrower, longer, and
lower, has the mesopterygoid fossa with its anterior edge
transverse, not ovate or lanceolate, and a shallow emargination
instead of the deep notch on the posterior edge of the palate
•on each side.
27. Profelis temminekii (Horsfield & Vigors).
(For bibliography see under the subspecies.)
Locality of the type, Sumatra.
Distribution. — From Tibet, Szechwan, S. China, and the
Eastern- Himalayas over the south-eastern portions of
Continental Asia to Sumatra.
One of the largest of the medium-sized Oriental species of
Felinse, with the tail a little over one-half to about two-
thirds the length of the head and body, and from over twice
to nearly three times the length of the hind foot. Coat
varying in luxuriance according to the race. General colour
exceptionally variable individually, from very dark brown
to red or grey, at least in the typical race. Pattern either
conspicuous and consisting of stripes on the dorsal area,
rosettes on the flanks, and solid spots on the limbs and below,
or obsolete except on the underside, where some spots are
retained, and on the face. But two features in the colour and
pattern are characteristic. The tail, unless affected by
melanism, is noticeably bicoloured, dark above and whitish
below, the two sharply contrasted, the whiteness below
being particularly conspicuous at the end of the tail, which
has no wholly black tip as in most other species. On the
face there is nearly always a pair of greyish Sands, bordered
with black stripes, running up over the forehead from the
conspicuous whitish patches on the inner side of the eyes ;
on the cheek there is a conspicuous white band edged by the
two black genal stripes and passing from below the eye to
below the ear, and the mystacial area of the muzzle is lined
alternately with narrow black and white lines, with some
brown in front below the rhinarium. The ears are blackish
behind, sprinkled with darkish-grey in the middle and towards
the base, but never have a conspicuous white spot.
The two British Indian races of this cat may be distinguished
as follows : —
a. Coat shorter, pattern at most faintly indi- [Vig., p. 260.
cated dorsally and laterally temminckn Horsf . &
b, Coat longer and fuller, a distinct pattern
of dorsal stripes and lateral rosettes tristis Edwards, p. 263.
PBOFELIS. 261
27 a. Profelis temmincMi temminckii (Horsfield & Vigors).
The Golden Cat or Temminck's Cat.
Felis temminckii, Horsfield & Vigors, Zool. Journ. iii, p. 451, 1828 ;
and of most subsequent authors, including Blanford and others.
Felis moormensis, Hodgson, Gleanings in Science, iii, p. 177, 1831 ;
id., Proc. Zool. Soc. 1832, p. 10 ; id., Journ. As. Soc. Bengal, x,
p. 908, 1841 (murmensis).
Felis aurata, Blyth, Proc. Zool. Soc. 1863, p. 185 (not of Temminck).
Felis nigrescens, Gray, in Hodgson's Cat. Mamm. Nepal in Brit.
Mus. ed. 2. p. 4, 1863.
Felis temminckii bainsei, Sowerby, China Journ. Sci. ii, p. 352,
1924.
Profelis temminekii temminckii, Pooock, Proc. Zool. Soc. 1932.
p. 754*.
Vernacular. — Hso-Hpai, Miao-Hton (N. Shan States) ; Kya
Min (Burmese).
Locality of the type, of temminckii, Sumatra ; of moormensis,
Nepal ; of nigrescens Sikkim ; of bainsei, Tengyueh, S.W.
Yunnan.
Distribution. — Nepal, Sutktm, Assam, and Btjrma ; also
Yunnan, Laos, Tong-king, the Malay Peninsula, and Sumatra.
Coat short and smooth in summer, moderately full in
winter. Colour variable, that of the upper side ranging from
very dark brown to red, ochreous-tawny or nearly grey,
the spinal area generally darker in all phases than the flanks
and outer side of the limbs, the hairs frequently speckled
with pale bands, which may be reddish or bleached whitish,
the grey phase resulting when the whitish speckling is extensive.
The underside ranges from brownish, with the chin buff in
dark skins, to whitish, with the chin white in paler skins.
Pattern, except on the head, where it is persistent, generally
inconspicuous or obsolete, at most consisting of some dark
stripes on the nape, faint stripes on the throat and inner side
of the fore leg, and of faint pale lines, sometimes forming
a network, on the shoulders, flanks, and thighs, and indicating
the spaces between large blotches like those of the Marbled Cat.
Although some skins show a combination of tints, the
dominant colour phases in British Indian skins in the British
Museum have the following geographical incidence : —
Dark brown, sometimes silvered on the flanks, from Nepal
and Sikkim (Hodgson and Mandelli) ; bright red, from Nepal,
* When more is known about this cat in Sumatra it is possible
that the Nepalese form moormensis may prove to be racially dis-
tinguishable from typical temminckii. But at present there is not enough
material from that island to separate the two. Sumatran specimens
are known to vary from " red " to nearly black, like those from Nepal.
Dark specimens from the latter country and Sikkim were called nigrescens.
The name bainsei was given to a trade skin by Sowerby. It was
described as brownish, with a tendency to greyish on the sides. All
the evidence is against Sowerby's view that the dark and red phases
of this cat are racially distinct.
262
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PLATE XXI.
MAMMALIA.
Temminck's Cat (Pnfelis temminckii temminckii).
(Drawn from grey skin from Upper Chindwin.)
Striped Temminck's Cat (Profelis temminckii tristh)
from Nam Tamai, Upper Burma.
PKOFEUS. 263
Sikkim, Sumbrabuni in Upper Burma (H. A. C. Stevenson),
and Teju in the Mishmi Hills, 3,000 ft. (H. W. Wells) ; ochreous
tawny in various shades from Sonapur, Kamrup (Sir D. Ezra),
Yekhum in the Naga Hills (J. P. Mills), Maymyo, Upper
Burma (G. C. Shortridge) ; grey, darker or lighter, from Myit-
kyina, Upper Burma (Capt. A. W. Kennion), and Upper
Chindwin (C. W. A. Bruce). There is also a grey skin from
Xien Quang Koo, Laos (Delaeour and Lowe).
Of the skins above referred to, one from Nepal and one
from Yekhum in the Naga Hills show grey stripes on the sides,
indicating the evanescent body-pattern, which in most skins
has disappeared altogether. Similar traces of the pattern
are visible in a skin from the Malay Peninsula.
The following are the only flesh-measured skins (in English
inches) in the British Museum : —
Head and Hind
Locality and sex. body. Tail. foot.
Myitkyina, Upper Burma ; ad. cj . 33 17 —
Tejo, Mishmi Hills ; ad. £ 32 18f- 7
Hue, Annam ; ad. <J 30 191 6i
The weight of the specimen from the Mishmi Hills was 32 lb.
In Fokien and other parts of Southern China this cat is
represented by a closely allied form, P. temminckii domini-
canorum Selater, which is distinguished by its fuller, longer
coat and bushier tail in the winter months.
•lib. Profelis temminckii tristis (Milne-Edwards).
Fetis tristis, A. Milne-Edwards, Rech. Mamm. p. 223, pi. xsxi,
1872 ; id., Congr. Internat. Zool. Nat. Mose. ii, p. 257, 1893 ;
Elliot, Mon. Felidas, pi. 22, 1883; B. Howell, Proe. U.S. Nat.
Mas. tsxv, p. 32, 1929.
Felis sem&novi, Satumn. Ann . Mus. Zool. Soc. St. Petersb. ix,
p. 524, 1904.
Profelis tenvminckii tristis, Pocock, Proo. Zool. Soe. 1932, p. 760.
Locality of the type of tristis, unknown (a trade skin from
Pekin) ; of semenovi, N".E. Szechwan.
Distribution. — Tibet, Szechwan, and % Uppbb Bttkma.
Distinguished from the typical race by its longer, fuller
winter coat, and by the retention of the pattern as usually
conspicuous blackish stripes on the head, nape, and back,
distinct blotches or rosettes sometimes joining to form oblique
chains on the flanks, solid spots on the limbs, and transverse
bands on the upper side of the tail. General colour typically
greyish, but frequently with an ochreous wash on the head,
nape, and shoulders. At Tsari, in Tibet, Kingdon Ward picked
up a very beautiful skin with the head, nape, spinal area,
tail, and underside rich rusty ochreous, the pattern everywhere
264= FELID.ffi.
black, the interspaces of the flanks silvery and the distal
fourth of the lower side of the bushy tail conspicuously
white.
No flesh-measured skins are available, but judging from
made-up skins in the British Museum from Szechwan and
Tibet the dimensions are approximately the same as in typical
temminckii. The entire skull is unknown, but the jaws and
teeth of the type ofsemenovi, a native skin from N.E. Szechwan,
seem to agree precisely with those of temminckii.
This race is included in the fauna of British India on the
evidence of a skin procured by Lord Cranbrook from a native
at Nam Tamai, 3,000 ft., in Upper Burma. The specimen
may have been locally killed or it may have come from over
the Tibetan border. The coat is not so full as in the Szechwan
skins, but, as likely as not, was procured in summer. The
general colour, too, is darker, the stripes on the back being
a good deal obscured by the blackish pigmentation of the
interspaces, and the hairs on the neck are speckled with
ochreous, on the flanks with grey.
Apparently all the skins of this cat that have come into the
hands of zoologists were procured from natives. It was
for many years regarded as a peculiar species, and from the
nature of its pattern Lydekker thought it was an Asiatic
representative of the American Ocelot. There is no doubt,
however, of its close kinship with Temminck's Cat.
Habits. — So seldom has Temminck's Cat been observed wild
by Europeans that there is very little trustworthy information
about its habits. By the natives of the Lushai Hills it is
stated to live amongst rocks. Those of the Mishmi Hills
report, on the contrary, that the female lies up in hollow trees
with her two cubs, which indicates a forest habitat. Probably
both accounts are true, the cat adapting its habits to its
surroundings. Since one was shot at Maymyo by Major
Stewart over a calf it had killed, and another was speared at
Victoria Point, Tenasserim, while feeding on a buffalo calf,
the species is clearly capable of preying upon comparatively
large game, and no doubt it subsists mainly on mammals up
to the size of small deer as well as upon such birds as pheasants
and wild fowl.
Of the habits of the spotted race tristis nothing is definitely
known, but there is no reason to suppose they differ from those
of its near ally.
PRIONAILTTRtrS. 265
Genus PRIONAILURUS Severtzow.
Prionailurus, Severtzow, Rev. Mag. Zool. (2) x, p. 387, 1S58 ;
Pocock, Ann. Nat. Mag. Hist. (8) xx, p. 338, 1917 ; id., Proc. Zool.
Soo. 1932, pp. 741-66.
Zibethailurus, Severtzow, Rev. Mag. Zool. (2) x, p. 387, 1858 ;
Pocock, Ann. Nat. Mag. Hist. (8) xx, p. 341, 1917.
Viverriceps, Gray, Proc. Zool. Soc. 1867, p. 268.
Type-species of Prionailurus, pardochrous (—bengalensis) ;
of Zibethailurus and Viverriceps, viverrinus.
Distribution. — The whole of south-eastern Asia in suitable
localities from Baluchistan and Kashmir in the west and
Manchuria and Korea in the north to Borneo and the Philip-
pine Islands in the south-east.
Tolerably large, medium or small-sized cats with the ear
low, rounded, and with a whitish patch on the back of it ;
tail a little over half the length of the head and body or less,
and a little over twice the length of the hind foot or less,
varying according to the species ; ground-colour variable,,
but pattern always distinct in the fresh coat and consisting
of four main stripes running back from the head on to the
shoulders, where they are typically broad and conspicuous,
but down the spine they are usually more or less broken up
into elongated spots ; sides of the body marked with spots,
which are frequently lanceolate, sometimes rosette-like,
occasionally tending to run into longitudinal chains, but never
fusing to form vertical stripes as in Felis ; fore legs typically
spotted externally to the paw, the hind at least to the hock ;
tail paler below than above, but the tints not sharply con-
trasted, and the tip never conspicuously white beneath ;
head with two cheek-stripes, a pair of white patches between
the eyes ; white upper lip.
The skull of Prionailurus differs from that of Felis (see below,
p. 285) in the sum of a number of characters. It is lower and
less vaulted ; the facial portion is shorter as compared with
the cranial, i. e., the distance between the occipital crest
and the tip of the postorbital process noticeably exceeds the
distance between that process and the end of the premaxilla ;
the postorbital processes are thinner, sharper, and less ligulate
before fusing with the process below them to complete the
orbital ring ; the nasal bones are not everted above the anterior
nares ; the floor of the orbit is longer ; the mesopterygoid
fossa is not widened anteriorly, and the median line of the
edge of the palate roofing it in front is typically notched,
not produced into a process ; the external pterygoid crest
is long and rises about the middle of the lateral wall of the
fossa, and the outer chamber of the bulla is much smaller as
compared with the inner.
266 FELIDJ3.
The skulls of the three species of this genus differ considerably.
That of P. bengalensis, the type of Prionailurus, is the most
generalized and most like the skull of Felis. The others
depart from it mainly in opposite directions. The skull
of rubiginosus is smaller and less developed muscularly,
resembling the skull of a young bengalensis, except in a few
specialized features mentioned below. The skull of viverrinus
is larger, with very marked muscular moulding. This species
was made the type of the genera Zibethailurus and Virerriceps
by Severtzow and Gray respectively*. In my paper on the
classification of the Felidse in 1917 I retained Zibethailurus
as a genus ; but I now think it better to regard it as a synonym
oi Prionailurus, to which, as I pointed out, it is unquestionably
nearly allied, differing from it in characters hardly greater
than those which separate the skulls of bengalensis and
rubiginosus.
Key to the Three Species based on External Characters.
a. Tail over half the length of the head and body-
arid over twice the length of the hind foot.
6. Larger, tail with distinct pattern ; body-
pattern bold, consisting of large black spots [p. 267.
or black and rusty rosettes on the flanks, bengalensis Kerr,
b'. Smaller, tail usually without pattern, at
most very indistinct ; body-pattern less
bold, consisting of small solid spots, but
two stripes on the shoulders always con- [p. 276.
spicuous rubiginosus Geoffroy,
a'. Tail less than half the length of the head and
body and less than twice the length of the [p. 281.
hind foot ; size, the largest of the genus . . . viverrinus Bennett,
Key to the Species based on Cranial Characters,
a. Skull of medium size, with nasals moderately
compressed and maxilla moderately ex-
panded above ; postorbital bar completed
later in life : muscular development of skull
moderate, at most a small sagittal crest and
postorbital area generally wider than the [p. 267.
muzzle in front bengalensis Kerr,
■a'. Nasals strongly compressed and maxilla
strongly expanded above ; postorbital bar
completed earlier.
b. Skull large, from 5 to 6 in. long, excessively
developed muscularly, with high sagittal
crest and postorbital area narrower than [p. 281.
muzzle in front, small upper pm retained . . viverrinus Bennett,
b'. Skull small, about 3 in. long, muscular
development weak, no sagittal crest, and
postorbital area always wider than muzzle,
which is vertical in front ; small upper [p. 276.
pm lost rubiginosus Geoffrey,
* These names were suggested by the specific name viverrinus, due to
Bennett's curious fancy that this cat is like a Oivet.
PEIOKAZLXTEUS.
26^
28. Prionailurus bengalensis Kerr. The Leopard-Cat.
(For bibliography of the British Indian races, see under the sub-
specific headings.)
Locality of the type, " Bengal."
Distribution. — As under the genus, but not found in Ceylon.
Size about the same as in the Domestic Cat, the tail typically
exceeding half the length of the head and body, and more
than twice the length of the hind foot. The coat soft and,
in the northern races, long and full. The ground-colour
variable, both individually and racially, from whitish, cream-
white, buff, ochreous-buff, to ashy-grey, with no bright tint.
Pattern similarly variable on the sides of the body, where the
spots may be large or small and solid, but usually the fore
part of the spots, especially when large, is invaded by an
Fig. 69. — Skull of Leopard-Cat (Prionailurus bengalerms), about
nat. size. (From Blanford.)
ochreous or rusty-ochreous tinge, so that they may become
nearly annuliform with dark centres ; and these " pantherine "
spots may show a tendency to coalesce in chains forming
longitudinal stripes.
The skull exhibits the mean in size and muscular develop-
ment in the genus. In the cranial part the temporal ridges
usually form a narrower or wider lyriform area, but occasionally
in the adult <J coalesce and give rise to a low sagittal crest ;
the postorbital area gradually expands from behind the
processes, where it is usually wider than the maxillary width,
sometimes about the same width, occasionally a little narrower ;
the postorbital bar is sometimes complete in skulls that have
just reached maturity, more often in older skulls, possibly
always in very old skulls, and the nasal bones are never strongly
constricted.
268 EELID-SE.
I regard as subspecies of P. bengalensis a large number of
described forms, including euptilura Milne-Edw. of Mancburia,
chinensis Gray of Southern and Central China, etc., which
many, even recent, authors cite as distinct species. They
have the same type of skull and teeth and are distinguished
merely by intergrading characters, like length of coat and
differences of tint and pattern. If granted the status of species,
the other British Indian cats, rubiginosus and viverrinus,
assigned to Prionailurus, must logically be given generic rank.
The distribution of the Leopard-Cat justifies as a working
hypothesis the view that, like the tiger and the panther, it is
a northern form that spread southwards over the Indo-
Malayan area before its disruption into continental islands,
and that it entered Hindostan by the route to the east of the
Tibetan plateau, but was too late to get a footing in Ceylon
after reaching the southern part of Peninsular India.
Owing to the individual variation in colour many names-
were given to British Indian specimens. Of these I have
selected two as connoting distinguishable subspecies, a northern
from the Himalayas, which has retained to a greater degree
the fuller winter coat of its still more northern ancestor,
and a southern, in which the coat has lost its fullness in warmer-
latitudes both to the west and east of the Bay of Bengal. This
appears to be a plausible explanation of the similarity between
skins from Lower Burma and Southern India ; and the race
to which I assign them, and to which the name bengalensis
has been restricted, is " environmental " and not " local."'
Skins from Assam are of doubtful allocation, since, as might
be expected, they are intermediate between Himalayan and
Burmese skins.
28 a. Prionailurus bengalensis bengalensis Kerr. Pennant's
Leopard-Cat.
Felis bengalensis, Kerr, Anim. Kingd. p. 151, 1792 ; and of most
subsequent authors, including Blanford, Mamm. Brit. Ind.
p. 78, 1888 (excl. most of the synonymy).
? Felis nipalensis, Horsfield & Vigors, Zool. Journ. iv, p. 382, 1829.
Ltopardus ellioti, Gray, Ann. Mag. Nat. Hist, x, p. 260, 1842.
Felis wagati and tenaeserimensis, Gray, Proc. Zool. Soo. 1867,
p. 400*.
* The typical example of this race was a stuffed specimen seen at
Hammersmith and described as the Bengal Cat by Pennant (Quadrup. i,
p. 272, 1781), who was told that the animal swam to a ship anchored off
the coast of Bengal. Although the story hardly bears the impress of
truth, and Pennant's description agrees better with rich-coloured
examples of the Sumatran race than with any Indian skins I have seen,
I adhere to the traditional acceptance of "Bengal" as the locality,
and restrict it to the coast of that province to the west of the Ganges.
The status of nipalensis is doubtful. The name was given to a specimen,
in the Zoological Gardens " said to have been brought from Nepal
PBIONAILUBUS.
269
Vernacular. — Chita Billi (Hindi) ; Ban Biral (Bengal) ;
Borka (Coorg) ; Wagati (Kanara and Dharwar) ; That-qyuk
(Burmese) ; Hen-wap (Shan States).
Locality of the type of bengalensis, Southern Bengal ; of
nipalensis, % Nepal ; of ellioti, the Bombay Presidency ;
■of wagati and tenasserimensis, Tenasserim.
Distribution. — Peninsulas India, Bitbma, Siam, Indo-
•China.
Coat comparatively short and thin and the tail not fluffy
in the winter months, as attested by specimens from S. Coorg
(Shortridge) in January and February and in the Palni Hills
<McCann) on April 25. Ground-colour and pattern alike
variable individually, the colour from ochreous-buff to buffish-
white on the flanks, but typically darker on the head and back,
" bright " and " dull " specimens occurring in the same or
adjoining localities, but the " bright " dominant. Spots on
Fig. 70.-
-The Bengal Cat (Prionailwrus bengalensis).
(From Blanford.)
the flanks typically large and well spaced, sometimes " solid "
with scarcely any brown dilution, more often " pantherine,"
and sometimes showing a tendency to run into chain-like
lines ; the stripes on the back also vary, particularly the
broad ones on tne shoulders.
Pennant described the type of bengalensis as " beautiful
pale yellowish-brown." The type of ellioti, on the other hand,
is dull buflish-white or grey, with the well-spaced spots on the
flanks mainly black. It is possibly faded ; but an adult $
•to Calcutta." Tlie type is tinlike any skin of this species I have seen
from Nepal. Its darkish-grey hue suggested to Blyth the possibility
of the specimen being a hybrid between a Domestic Cat and a Nepalese
example of bengalensis. But the ears and the pattern, as well as the
jaws and teeth left in the skin, are of the PrionaUurus-type. The skin,
indeed, differs only in its softer fur from grey, bold-patterned skins of
P. vwerrinus, and almost exactly matches a skin of the bengalensis-type
from the Malay Peninsula (Dr. Cantor).
270 HELIDiE.
from Virajpet, S. Coorg, 2,000-3,000 ft., closely resembles it.
Between this and the richest-coloured skin from the same
locality, which has the dorsal area from the head to the root
of the tail rich ochreous-buff, with the flanks paler and the
spots diluted in front or centrally with rusty-ochre, every
gradation exists. An example from Haleri, N. Corg, 3,555 ft. ,
is like the brightest from Virajpet, but has the spots more
diluted and hence more " pantherine." A skin from Ootaca-
mund in the Mlgiris, 7,000-8,000 ft. (Gosse), and two from the
same range (Phythian Adams) are richly tinted ; and one
from Tiger Shola in the Palni Hills, 5,700 ft. (McCann), is
intermediate in tint between the bright and dull types, and
has markedly pantherine rosettes.
A series of six, mostly rich-coloured, " native " skins from
the Chin Hills, 50 miles west of Kindat in Upper Burma
(Mackenzie), shows similar variation in tint. One is " dull-
coloured," another almost exactly matches the skin from
N. Coorg, and also closely resembles the type oftetiasserimensis,
mentioned below ; a third is a handsome dark skin, with the
back almost infuscate, the flanks ochreous-buff, the belly
buff, the pattern bold, with the neck and shoulder-stripes
broad, the rosettes large and sometimes completely surrounded
with black. A fourth skin, reddish all over, even on the belly
and backs of the ears, may be artificially stained. Another
with broad stripes and pantherine rosettes was captured at
Thayetmyo, Upper Burma (Mackenzie). A series from
Toungoo in Lower Burma (Mackenzie) closely matches the
Southern Indian skins, the rosettes in one coalescing to form
rusty-oehreous, black-bordered bands. The co-types of
Gray's Felis wagati from Moulmein are very like his type of
dlioti from Bombay. If faded, as is not unlikely, they
probably originally resembled the brighter-tinted skins from
Coorg. The type of tenasserimensis Gray is a little richer in
tint than the Moulmein skins and has the pattern much
bolder, the rosettes being large, few in number, and diluted
with ochxeous-brown, and the broad shoulder-stripes join
posteriorly to form a nearly symmetrical pattern of two Vs,
with oehreous-brown between the arms, Finally, two examples
from Mergui Town (Shortridge) differ from the Moulmein
skins, and still more from the type of tenass&rvmensis, in having
narrower, more broken stripes and smaller, more numerous
spots. Not surprisingly they approach in these respects
some skins from the Malay Peninsula and the Javan and
Sumatran races of bengalensis.
As stated above, Assamese skins are of doubtful racial
reference, but in coat they seem to agree better with typical
bengaknsis than with the Himalayan race. One from Tura
HRIONATLURUS. 271
in the Garo Hills, 1,450 ft., is dull buffy- white, like the typical
Himalayan form and several skins of bengalensis, and one
from Mokokchung in the Naga Hills, 4,500 ft., though not
so pale, also fits the Himalayan race ; a skin from Shulong,
5,200 ft., is rich-coloured, and one from Lakhimpur is similar,
but not so bright. Nevertheless the available skulls seem to
agree better in size with those of horsfieldi from the Himalayas
than with those from S. India assigned to bengalensis, although
there are very few of the latter to judge from (see table of
measurements, p. 275).
Of the distribution of this cat in India Blanford said that
it is " common in Lower Bengal " and " is also found in the
Syhadri Range or Western Ghats, Coorg, Wynaad, Travancore,
etc., and in some, perhaps all, of the other forest-regions of
the Peninsula, though not very abundantly. I have never
seen a specimen during several years' wandering in the Central
Provinces and the northern part of the Bombay Presidency.
There is, however, a skin said to be from the neighbourhood
of the Coromandel coast in the Calcutta Museum ; and a living
specimen from Jeypore, west of Vizagapatam , was quite recently
sent to the Zoological Gardens in London." So far as the
western and central portions of Peninsular India are concerned,
this account was confirmed by the collectors for the Bombay
Mammal Survey ; but no specimens were secured either in Lower
Bengal or in the Eastern Ghats. Clearly the race is now,
at all events, rare over the whole of the eastern portion of the
Peninsula.
Habits. — According to Shortridge it is fairly plentiful in
Coorg, more so than the Jungle-Cat, and is particularly
numerous around villages, destroying large numbers of fowls.
This confirms the account given by Jerdon, who adds that it
shelters in hollow trees and in May produces three or four
kittens in caves or beneath masses of rock. It is essentially
a forest animal, preying upon small mammals and birds.
28 b. Prionailurus bengalensis horsfieldi Gray. Horsfield's
Leopard-Cat.
Felis nipalensis, Hodgson, Jouxn. As. Soc. Bengal, i, p. 341, 1832
(not nipalensis, Horsf. & Vigors, 1829).
Leopardus horsfieldi, Gray, Ann. Mag. Nat. Hist, x, p. 260, 1842.
Felis pardochrous, Hodgson, Calc. Journ. Nat. Hist, iv, p. 286,
1844*.
* Hodgson originally identified and described Nepalese examples
of this oat as nipalensis Horsf. & Vig. ; but subsequently, discovering
his mistake, substituted pardochrous, which, being assigned to a described
form, is not a " nomen nudum," as Blanford stated. But in the mean-
time Gray had proposed horsfieldi for a skin believed to have come from
Bhutan, but possibly from Upper Assam (McClelland). Since the types
272 FELIDiE.
Vernacular. — Bandaris $, Biralu $ (Hindi).
Locality of the type, of horsfieldi, Bhotan ; of pardochrous,
Nepal.
Distribution. — Kashmir, Kumatxn, Nepal, Bhutan ; north-
ern and eastern range uncertain.
Distinguished from the typical race by its apparently larger
skull and by the more luxuriant coat and more bushy tail
in the winter months. There is some evidence too that on
■the average the ground-colour is paler and not so richly
■ochreous, but the colour and the pattern vary as in typical
bengalensis, and specimens belonging to the two may be
closely matched.
Excluding some old skins from Simla, Nepal, and the types
of horsfieldi and pardochrous from Bhutan and Nepal
respectively, the British Museum has several collected by the
Survey from Sikkim up to 8,800 ft. (Crump), from Satthar
Hill, Gorkha in Nepal (Baptista), January, from various
localities in Kumaun, including Naini Tal, 5,000 to 6,000 ft.
(Crump), and a rather bright-coloured, immature skin from
Sardalla, Kashmir, 8,700 ft. (Stockley), the most western
known locality for the race.
The possibility of the existence of another race in the
north-eastern area of British India is suggested by several
skins noticeably darker and more richly tinted than those
assigned to horsfieldi. Some of them are certainly native
skins, and may have been traded over the border and not
captured at the localities entered on their labels. One of these
was from Nani Tisang, 2,500 ft., near the Tibetan border
(Lord Cranbrook). It appears to resemble very closely the
type of Felis scripta Milne-Edwards (Nouv. Arch. Mus. vii,
Bull. p. 92, 1870, and Rech. Mamm. p. 341, pi. 57, 1872)
of horsfieldi and pardochrous are scarcely distinguishable, horsfieldi
comes in as the older name.
Two additional names given to Himalayan, or alleged Himalayan,
skins by Hodgson and Gray must here be referred to, since they were
cited by Blanford as synonyms of his " var. pardichroa " of bengalensis.
The first is Felis ogilbii, given by Hodgson (Calc. Journ. Nat. Hist, viii,
p. 44, 1847) to a skin from Sikkim. Blanford identified as the type
of ogilbii one of Hodgson's skins in the British Museum from Sikkim.
It is an artificially dyed native skin, with the tail less than half the
length of the head and body ; but it is not the type of ogilbii, since
the tail in the latter considerably exceeded two-thirds the length of
the head and body. No doubt it was a specimen of the Marbled Cat
(Pardofelis marmorata) (see p. 256). The second is Felis herschelii,
given by Gray (Cat. Cam. Brit. Mus. p. 28, 1869) to a skin labelled
" Gangootri " which is in Tehri Garhwal. But since this skin, both in
coat and colour, differs from the skins of horsfieldi from the Himalayas,
and is, on the contrary, absolutely indistinguishable from richly tinted
skins of the Sumatran and Bornean race (P. bengalensis sumatrana),
I have no doubt its alleged locality is erroneous, owing to substitution
•of labels.
PLATE XXII.
MAMMALIA.
Photo F. W. Bond.
Leopard-Cat (Prionailums bengaknsls).
Photo F. W. Bond.
Fishing-Cat (Prionailurus v/verrinus).
PRIONAILURUS. 2;5
from Szechwan, and has similar large, dark, normally separated
rosettes. It is also like the type of P. ingrami Bonhote-
{Aim. Mag. Nat. Hist. (7) xi, p. 474, 1903) from Kwei Chow,
China, a synonym, of scriptus, except that the coat is not so
thick. The others are from Nepal (Hodgson) and Sikkim
(Crump). One of Hodgson's is certainly a native skin and
the other may be, but the evidence of this has been destroyed
by the thoughtless " make-up " of the skin in the conventional
style in the British Museum. The pattern in both shows
a tendency to run into chains of rusty and black spots. The
Sikkim skin was picked up in Darjeeling, already dressed,
and, as Crump supposed, may not have been locally caught.
In any case it is much brighter than normal skins of horsfieldi.
Admittedly these skins may be exceptionally well-coloured
skins of horsfieldi. In that case this race will be represented
by " bright "- and '" dull "-tinted specimens, like bengalensis
in Southern India and Burma. If they belong to the more
northern race, then P. bengalensis scriptus Milne-Edw., with
ingrami and, I think, anastasim Satunin (Ann. Mus. Zool.
St. Petersb. ix, p. 528, 1904), from Kansu, as synonyms, which
appears to differ from horsfieldi by its shorter tail and brighter
colour, will come in as an element in the British Indian fauna.
The habits of this race are no doubt the same as those
of typical bengalensis. According to a note by Crump the
specimens he procured at Naini Tal in Kumaun were trapped
in dense jungles near a stream. It is probably fairly common,
he thinks, in the hills where there is heavy forest, but it is
not often seen owing to its liking for dense cover.
28 c. Prionailurus bengalensis trevelyani, subsp. nov. Tre-
velyan's Leopard-Cat.
Locality of the type, near Gilgit, 5,000 ft.
Distribution. — Northern Kashmir and the Uther Punjab
in the drainage area of the Indus and Jhelum, approximately
long. 74° E. ; also S. Baluchistan.
Distinguished from P. bengalensis horsfieldi by its rather
fuller longer coat, from 36 to 40 mm. long, and by the typically
paler, greyer, sometimes nearly silvery ground-colour of the
upper side, due to the absence of buff or at most to a faint
wash of that hue, but low down on the flanks and on the
outside of the limbs there is a faint buff tinge ; the " pantherine ! '
spots may be invaded by tawny or bumsh-tawny hairs in front.
All the known skins of this race were collected comparatively
recently, and their pale colour is not attributable to fading
from a long period of preservation. Three from Gilgit
(Col. W. R. F. Trevelyan) show slight variation in tint which
is probably seasonal, one collected in March being a trifle
brighter than a skin, bought in the bazaar, of a specimen
VOL. I. T
274 TELma:.
killed at Ishkoman in June, which, like the undated type,
is greyish-white in the ground-colour. An undated skin
from Chilas, a few miles nearly due south of Gilgit (Capt. L. W.
Wooldridge), and a <J skin from Kotli, Murree, 5,800 ft. (Wells),
June 17, with the moult in progress, are also whitish-grey.
Closely resembling these skins is one from the Las Belas
Territory, 80 miles north-west of Karachi, which is of interest
as the only representative of the Leopard-Cat ever recorded
from Baluchistan.
The dimensions and weights recorded below are about the
same as in the other races. The skull, too, is similar both in
size and shape, as shown in the table of measurements.
The occurrence of this Leopard-Cat at Gilgit and in south-
eastern Baluchistan suggests that it extends in suitable
localities over the intervening area to the west of the Indus.
In Kashmir it no doubt intergrades with Horsfield's Leopard-
Cat (P. b. horsfieldi), since Col. Stockley's specimen of the
latter from Sardallu attests its westward extension into that
country. But in the southern part of its range Trevelyan's
Leopard-Cat is widely separated from typical bengafansis,
which is unknown, apparently in Sind, Cuteh, and Bajputana.
Of its habits nothing has been recorded ; but possibly it is
less addicted to dense jungle and forest than the other British
Indian races. Its generally pale colour, recalling that of the
Snow-Leopard, suggests that its habitat may be rocky situa-
tions, and that it breeds and takes shelter in crevices or small
caves.
Comparatively few flesh-measurements of adult British
Indian skins of this species are available. The following indicate
general similarity in size of the three races described : —
Head and Hind
Name, locality, and sex. body. Tail. foot.
Murree ; ad. S 21| 12 4£
"horsfieldi.
Naini Tal, Kumatm ; ad. 3 21$ 12 4f
■Gorkha, Nepal ; ad. <? 21f 12$ 5
Gorkha, Nepal ; ad. $
•Gorkha, Nepal ; ad. § 19$ \\t 43
bengalensis.
Palni Hills : just ad. <J 20 9i 4*
S. Coorg : ad. 9 20$- 11 4$
Garo Hills, Assam ; ad. $ 20$ 11* 4*
Toungoo, Burma : ad. <J 24$ 121 4*
Toungoo, Burma j ad. S 21$ 11$ 44
Toungoo, Burma ; ad. ? 19f H} 4$
The weights of the <J from Murree, Gorkha, and the Palni
Hills was 6 lb., from the Garo Hills 6f lb., from Naini Tal 7 lb.,
and from Gorkha, the larger of the two, 8 lb. The ? from
S. Coorg was 6J lb. and from Gorkha 6 lb.
PBIONAILUETJS.
275
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276 FELIDiE.
When discussing the size of the " varieties " of the Leopard-
Cat Blanford quoted McMaster's statement that Burmese
specimens are smaller than South Indian specimens, and this
conclusion he accepted when he wrote of Gray's specimens
of wagati as representing a small race. But from the table
of flesh-measurements it is quite clear that specimens from
Burma are not smaller. One of the specimens from Toungoo,
indeed, is the largest Leopard-Cat recorded. Blanford's
opinion about the small size of wagati rested on his examination
of the skulls of Gray's specimens, which he apparently thought
were mature. They are obviously immature, as shown by
the skulls, and consequently undersized.
In the table of skull-measurements (p. 275), Assamese
skulls from the Garo and Naga Hills have been included
under horsfieldi because of their size, which comes between
the two $ skulls from Kumaun, the largest and smallest
respectively from that district. From the evidence it appears
that these northern skulls are bigger than those from S. India,
and on the average bigger than those from Burma.
The mastoid width in the skulls from Gilgit, Almora,
Kumaun, Garo Hills, and S. Coorg is respectively 37, 40, 39,
and 35 mm., noticeably less than half the condylobasal length.
In all skulls of the species the proportion is about the same.
29. Prionailurus ruljiginosus Geoffroy. The Rusty-spotted
Cat.
Felis nibiginosa, I. Geofiroy, in Belanger's Voy. Ind. Or. p. 140, pi.,
1834 ; and of nearly all subsequent -writers on the Indian fauna,
including Blyth, Jerdon, and Blanford.
Prionailwrue rubiginosus, Pocoek, Ann. Mag. Nat. Hist. (8) xx,
p. 339, 1917.
Locality of the type and only known species, Pondicherry,
S. India.
Distribution. — Southern India and Ceylon.
Distinguished from the British Indian races of P. bengalensis
by its smaller size, by the tail being uniformly coloured or
at most with very indistinct pattern, and by the pattern on
the body being less bold, consisting of small, tolerably widely
separated solid spots, often lanceolate, frequently brown on
the sides, and exhibiting a longitudinally lineal arrangement
on the back ; but on the head and shoulders it forms definite
stripes, a pair on the shoulders above being noticeably dis-
tinct ; the ground-colour varies according to the race.
Apart from its smaller size, the skull of P. rubiginosus differs
from that of P. bengalensis in several particulars, principally
in the shortening of the upper jaw, accompanied by the down-
ward curve of the end of the nasals and the nearly vertical
plane of the anterior nares and of the maxillo-premaxillary
PRI0NA1LURUS.
277
suture ; also the maxillae are greatly expanded above and the
nasals strongly constricted in their posterior portion ; the
orbits in the adult are completely surrounded, and there is
never a sagittal crest, the temporal ridges forming a lyriform
area ; in the teeth the inner lobe of the upper carnassial is
more reduced, and as an accompaniment of the shortening
of the muzzle the area behind the upper canine is also short
and has lost the normal small premolar in front.
Fig. 71. — Rusty-spotted Cat, Prionailurus rubiginosus.
(From Blanford.)
29 a. Prionailurus rubiginosus rubiginosus Geof&oy.
Felis rubiginosa, Geoffroy, in Belanger's Voy. Ind. Or. p. 140,
1834 ; and of Jerdon, Blanford, and others (in part).
Vernacular. — Namali pilli (Tamil, Madras) ; Ark-philU
(Waddars) ; Kiraba-bekhu (Kanarese).
Locality of the type, Pondicherry.
Distribution. — Southern India.
Colour above from the head to the tail- tip and on the outside
of the limbs grey, but darker and more drabby on the back,
and the limbs paler than the flanks, the hind limbs whitish
distally, and the pale areas on the head white ; the pattern
varying from black above and below, brownish-black on the
flanks to entirely brown, the spots on the flanks sometimes
quite faint.
This description is taken from three skins in the British
Museum : one from Madras (Jerdon) , one from Nellore (Bartlett) ,
and one from Khandala, between Bombay and Poona, 1,700 ft.
(P. Gosse), April 22. The last is an unfaded skin, with the
coat fresh and soft and the pattern dominantly black or
blackish. The others, undated, with the pattern brown
and much less distinct, are old skins and may be faded, but
278 JETELID-ffi.
they agree very closely in general appearance with Geoffroy's
original figure of the type shipped from Pondicherry.
The flesh-dimensions (in English inches) of the only skin
so measured are : — Khandala, adult $ : head and body 16jt ;
tail9£; hind foot 3-|.
29 b. Prionailurus rubiginosus phillipsi, subsp. nov.
Felis rubiginosa, Kelaart, Prod. Faun. Zeyl. p. 47, 1852 ; Phillips.
Man. Mamm. Ceylon, p. 158, 1935.
Vernacular. — Wal balala, Kula diya (Sinhalese) ; Kadu-
poona, Verewa puni (Tamil) ; Kardup-poonai (Tamil at Jaffra).
Locality of the type, Mousakanda, Gammaduwa, C.P.,
3,000 ft.
Distribution. — Ceylon.
Distinguished from the typical Indian race by being, on
the average at least, darker, richer, and less grey in tint.
The type, March 16 (W. W. A. Phillips), is warm brown
above from the head to the end of the tail, the tail being
Fig. 72. — Skull of type of Phillips's Rusty-spotted Cat, Prionailurus
rubiginosus phillipsi, from Gammaduwa.
obscurely marked with darker transverse stripes ; the flanks
are a little paler, but the white of the underside is bordered
by an ochreous wash ; the fore legs are rich ochreous externally,
the hind a little paler, and the pale areas on the face are tinted
with buff.
Another specimen from Gammaduwa, undated, is very
like the type; others from Kandy, 4,500 ft., Ambawela,
5,600 ft., March 24, Hakgalla in Uva, 5,000 ft., April 6 (E. W.
Mayor), and some labelled merely " Ceylon," although varying
somewhat in depth of hue, are all darker than the Indian
skins referred to above. One skin, however, from Weligatta
(1. W. Mayor), June 28, is greyer and scarcely distinguishable
from ike skin of typical rubiginosus from Khandala, except
that the head is darker, not so grey. Comparison between
FKIONAILTriUJS. 279
this skin, and others dated suggests seasonal change in colour
between the beginning of April and the end of June.
Flesh-measurements (in English inches) are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Phillips's largest ; ad. <J 18| 10 3§
Ambawela ; ad. <j 17 8 3f
Gammaduwa (type) ; ad. <J 16 9* 3£
Weligatta ; ad. <J 151 9J 3§
Phillips's average of 5 ; ad. <J 16i 8i 3&
Hakgalla, Uva ; ad. $ 16" 7g 3£
Phillips's largest ; ad. $ 16| 8f 3£
Phillips's average of 3 ; ad. $ 15 8+ 3 J
The weight of the <J from Ambawela was 3 lb., of those from
Gammaduwa and Weligatta 3£ lb., these agreeing with
Phillips's largest g. Phillips's largest $ was 2§ lb., and the
average of 3 $$ 2| lb.
The mastoid width in the two skulls from Gammaduwa
and the one from Weligatta is 34, 31, 33 mm., less than half
the condylobasal length. In all the skulls the proportion
is about the same.
Habits. — The two races of P. rubiginosus differ in habitat.
According to Jerdon's observations at Madras the Indian race
frequents grass in the dry beds of tanks, brushwood, and
occasionally drains in open country and near villages, and was
said to be not a denizen of the jungles. The specimen from
Khandala was trapped in a ravine below the cemetery, not
far, apparently, from the village.
Shortridge's remark that this cat is " apparently rare "
in Dharwar seems to apply to it everywhere in India, judging
from the few specimens preserved in the British Museum and
from the failure by the collectors of the Mammal Survey to
shoot or trap it, apart from two secured by Gosse and Short-
ridge. Its most northern recorded locality is Seoni in the
Central Provinces (Sterndale).
Phillips states that the Ceylon race '" occurs in the jungles-
throughout the whole island from the highest mountain-peaks
to the sea-shore, but is nowhere very common. The type
was killed in " heavy forest." It is mainly nocturnal, usually
lying up during the hours of sunshine in a hollow log, tree
or thicket in small woods of heavy timber or in thick scrub-
jungles, only rarely being found away from the jungles.
It is an excellent climber, often seen in trees, but probably
feeds mainly on the ground, preying upon small mammals-
and birds, possibly on lizards and frogs as well, and not
infrequently it breaks into hen-houses near the jungle and
slaughters the fowls. The litter consists of two or three
kittens born in a hollow log or small jungle-cave.
280
FELnxas.
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PRIONAILLTtUS. 281
30. Prionailurus viverrinus Bennett. The Fishing-Cat.
Felis viverrinus, Bennett, Proc. Zool. Soc. 1833, p. 68 ; and of
most recent authors, including Jerdon and Blanford.
Felis himalayanus, Jardine, Nat. Libr., Felinse, p. 230, pi., 1834.
Felis viverriceps, Hodgson, Journ. As. Soc. Beng. v, p. 232, 1836.
Yiverriceps bennettii, Gray, Proc. Zool. Soc. 1867, p. 268.
Locality of the type of viverrinus, " India," probably the
Malabar coast ; of himalayana the Himalayas ; of viverriceps,
Nepal ; of bennettii, " India."
Distribution. — India, precise range unknown ; Ceylon,
and east of the Bay of Bengal to Cochin China and Java.
Distinguished in external characters from P. bengalensis
by its much larger size, shorter tail — which is a good deal less
than half the length of the head and body — less completely
webbed feet, and incompletely sheathed claws, the points of
which normally project beyond, the hairs of the paws ; the
coat also is on the average harsher, and the pattern, although
exhibiting the same general arrangement, is subject to much
less variation. The marked differences in the skull between
the two species are described below.
The general colour varies from deep olivaceous-tawny
above and greyish on the flanks to tolerably uniformly yellower
tawny or nearly ashy-grey, without any bright hue. In the
pattern the stripes may be comparatively broad, especially
on the shoulders, or narrow, and the spots on the flanks may
be roundish, ovate or lineate. The tail is the same tint above
a.s the back, spotted at the base and banded distally ; the
limbs are about the same tint as the flanks, and the pattern
extends to the wrist and hock. The underside is white,
with heavy pattern forming two collars on the throat and
transverse bars behind the fore legs on the chest.
Despite the extensive geographical range of the species,
there appear to be no distinguishable local races. Skins
resembling each other occur in widely separated areas, and
skins from the same district may differ considerably from one
another. For instance the type, probably from the Western
Ghats *, is a dark skin, with bold pattern, and is indistinguish-
able from three skins collected near Batavia in Java. Equally
dark, but with rather finer pattern, is a skin from Barkot,
Dehra Dun, 5,000 ft. (Capt. F. S. Tuker), and one probably
from North India (Lord Ripon). But two skins from Nepal
(Hodgson) are paler and tawnier, one being paler and greyer
tawny than the other. One skin from Sehwan, Sind (Capt.
Watson), is decidedly tawny, rather richer than the tawnier
of the two from Nepal, but another from Mirpur, Sind
* This is inferred from its being received by Bennett with a specimen
of John's Langur from the same donor, both being recorded as from
" India."
282 FEiXD-ai.
(Commander J. J. Walker, R.N.), is pale olivaceous-grey,
with no tawny tint. Nearest to this comes a skin from
Kanthalai, E.P., Ceylon (W. W. A. Phillips), which has, however,
a slight tawny wash on the back. Another skin from Kandy
(Whyte) is still tawnier, very like the tawny skin from Sind.
Erom countries outside British Indian limits there are two
skins from Nhatrang, Annam (Dr. Vassall), one matching
the tawnier of the Nepalese skins, the other being tawnier
than the tawny Sind skin, but a skin from Cochin China is
dark olivaceous tawny, like the type and the Javan skins.
Of the above-mentioned skins two only are dated, the one
from Kanthalai, July 30, and the one from Mirpur, Sind,
Pec. 24. Both are pale, despite the difference in the time of
the year. Hence there are no data justifying the conclusion
that the differences in tint are seasonal.
It is noticeable that apart from the type, which, like the
Javanese skins, has a bold pattern, the skins from British
India have on the average a finer pattern of narrower stripes
dorsally and streak-like, lineate spots on the flanks.
The only measurements of this species certainly taken in
the flesh are those recorded by Phillips from Ceylonese speci-
mens. These, however, agree very closely with some dimen-
sions given by Jardine and Hodgson of unsexed Himalayan
specimens : —
Head and Hind
Locality and sex. body. Tail. foot.
Himalayas (Jardine) ; ad. S • 28* 94- —
Nepal (Hodgson) ; ad. <J ? 30" 10l —
Ceylon, Phillips's largest; ad. •£ 30& 12 J 7|
Ceylon, Phillips's average of 3 ; ad. <}. 28* 11 6}
Ceylon, Phillips's average of 3 ; ad. §. 26" 10 5$
In addition to its larger size, the skull of viverrinus differs
from that of hengdknsis mainly in characters resulting from
the greater development of the masticatory muscles, which
has produced a high sagittal crest and a long, narrow, post-
orbital " waist," always narrower than the width of the
muzzle above the canines. These differences are naturally
particularly well marked in the adult $, the skull of which
is considerably larger than in the adult $.
The mastoid width of the <J skulls from Sind, Ceylon, and
of the second ? skull from Nepal is respectively 60, 55, and
43 mm., much less than half the condylobasal length. A similar
proportion obtains in all the skulls of the species.
Several additional skulls labelled "India," and others
from Indo-China and Java agree, apart from minor details,
with those entered in the table, and, like the skins, bear out
the view that the species is not susceptible, on the available
evidence, of division into local races.
PRION AILXJRUS .
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284 FELiD^i.
Habits. — Although occurring occasionally at comparatively
high altitudes, e. g., 5,000 ft. at Dehra Dun, this cat is mainly
a lowland species partial to reed-beds and marshy districts.
Hodgson recorded its habitat in Nepal " as the open lowlands
of the lower regions," and Jerdon wrote of it as inhabiting
the " edge of swampy thickets in Purneah," " marshy regions
at the foot of the Himalayas," and " reed-beds " near Calcutta.
This habitat is not restricted to British India, two Javan
skins in the British Museum being labelled as killed in " swamps
bordering the coast near Batavia." The species does not,
however, appear to be common anywhere; its distribution
in India is very imperfectly known, and there appears to be
no record of its occurrence in Burma, although its existence
in that country may be inferred from its being found in Indo-
China and Java.
The popular name " Pishing Cat " reflects the testimony of
several observers that it feeds on fish. According to Buchanan-
Hamilton it even eats hard-shelled freshwater molluscs, a gas-
tropod (Ampullaria), and a mussel {Vnio). The name, never-
theless, is somewhat misleading *. The teeth are not especially
adapted in any way for catching and eating fish or for crushing
shells of molluscs ; they are fitted, as is the whole organization
of the species, for preying upon any terrestrial vertebrate
animals it can overcome, and there are recorded cases of calves,
sheep, dogs, and large snakes being killed by it. It has
also been known to carry off human babies.
According to Phillips this cat in Ceylon is found '' sparingly
in the jungles aE over the island except, perhaps, in the dry
northern zone, Tirripane, 365 ft., near Anaradhapura, N.C.P.,
being the most northern point whence it has been recorded.
It is usually found in or near the heavier or larger jungles, but
may be met with in scrub or in reed-beds and long grass
beside rivers and in swamps f. It may be seen at any hour
of the day, and although it can climb well it seldom takes
to trees, and feeds mainly on the ground. Phillips confirms
the reports of observers in India that the Fishing Cat puts up
a desperate and generally successful fight when attacked by
dogs.
Nothing seems to be known about the breeding habits, but
there is no reason to suppose they differ from those of other
wild cats.
* Specimens I observed in the Zoological Gardens, London, showed
no greater liking for fish than wild cats of other species.
f The specimen from Kanthalal, sent by Phillips to the British
Museum, had entered a tank from which it was unable to get out.
tblis. 285
Genus FELIS Linnaeus*.
Felis, Linnaeus, Syst. Nat. ed. 10, i, p. 41, 1758.
Chaus, Gray, List Spec. Mamm. Brit. Mus. pp. 44-5, 1843.
Type of Felis, catus Linn, (the Marbled or Blotched Tabby
Domestic Cat) ; of Qhaus, chaus, Guld.
Distribution. — Central and Southern Europe, Central and
Southern Asia as far east as Kansu and Annam ; the whole
of Africa except the Congo Forest area ; everywhere for the
most part in more or less open country — not in thick forest.
Distinguished from the other genera of Pelinae by a well-
marked combination of external and cranial characters.
The ears are high, triangular, narrowed at the summit, and
never have a white patch at the back; the pattern, when
retained in the adult, consists on the side of the body, at least
in front behind the shoulder, of vertical lines of spots or of
stripes due to the linear confluence of spots, and the paws
are narrow, with comparatively weakly developed claw-sheaths
and interdigital webs.
The skull has the cranial portion short as compared with
the facial, the frontal postorbital process, which is broad, ligu-
late, and blunt, being about in the middle of the total length ;
the nasal branch of the premaxilla is broad, especially on
a level with the tip of the nasal, then somewhat abruptly
narrowed to a point ; the mesopterygoid fossa is wide in front,
its anterior palatine edge typically has a median backwardly
projecting point, never a notch, the external pterygoid crest
is reduced to a triangular point just above the hamular process,
the posterior edge of the palate has a well-developed emargina-
tion inside the last upper tooth, and the partition of the bulla
at its summit is remote from the internal crest of the tympanic
bone, so that the outer chamber of the bulla is comparatively or
very large.
* This name is here used in the sense adopted in my paper on the
" Classification of the Felidas " (Ann. Mag. Nat. Hist. (8) xx, p. 333, 1917).
In addition to Chaus I there cited as synonyms of Felis : — Qatus Fitz.,
Gatolynx Severtz., both based on F. catus Linn., and Otailurus Severtz.,.
based on F. megalotis, a Feral Domestic Cat from Timor. Since then
several generic or subgenerie names have been introduced : — Poliailurus-
Lonnb. for a Central Asiatic Cat, pallida Biichn., Eremselwus Ognev
for a Transcaspian Desert Cat, ihinobius, and Mierofelis Roberts for the
little S. African species, nigri/pes, the distinctive characters of which
I described in 1907. I regard these names as merely connoting well-
defined species. The four species I have seen as living animals, namely,
the European Wild Cat (sttvestris) , the typical African Wild Cat of various
races (constantina), the Jungle-Cat (chaus), and the Black-footed Cat
(nigripes), exhibiting the extremes in size and other features, all have
an unmistakable, if undefinable, " facies," distinguishing them at once
from such species as the Leopard-Cat (PrionaMurus) and its allies ;
and Ognev's figure of the living Eremselurus clearly shows that it belongs
to the same group.
286 FELmaE.
The two British Indian species of this genus may be dis-
tinguished as follows : —
■a. Size as in the Domestic Cat, tail more than half
the length of the head and body and over
twice the length of the hind foot ; pattern
of dorsal stripes and of lateral spots con-
spicuous ; spinal crest and stripe hardly
noticeably differentiated ; skull shorter, with [Gray, p. 287.
broader zygomatic arches constantina ornata
■a'. Size larger, tail less than half the head and
body, and less than twice the length of the
foot ; pattern in adult obsolete dorsally,
at most faint on the flanks ; spinal crest and
stripe clearly differentiated ; skull longer,
with, narrower zygomatic arches chaus Guld., p. 290.
31. Felis constantina Forster*.
Felis constantina, Forster, TJebers. Buffon's Naturg. vierfuss. Tiere,
vi, p. 313, 1780.
Felis lybica, Forster, loc. ait. ; Meyer, Syst. Zool. Entol., etc.,
p. 101, 1793.
FeUs ocreata, Gmelin, Suppl. to Ouhn's Bruce, Reisen Abyss, ii,
p. 27, 1791 ; Schwann, Ann. Hag. Nat. Hist. (7) xiii, p. 421, 1904.
Locality of the type of constantina, Constantine in Algeria ;
■of lybica, Gafsa or Kafsa in Tunis ; of ocreata, Ras el Feel
in Abyssinia.
Distribution. — Southern, Eastern, and Northern Africa,
Sardinia, South-Western to Central Asia and Northern
India, agreeing closely with that of the Caracal and Hunting
Leopard.
Size typically about that of the Domestic Cat, some of the
African races smaller ; the tail more than half the length of
the head and body and more than twice the length of the
hind foot. Typically there is a small hair-tuft on the tip of
the ear and a crest of longer hair on the spine behind the
shoulder. General colour very variable racially, from more
•or less sandy to ashy or stone-grey, deep blackish-grey or
brown above, with the crest darker ; the underhair of the back
usually ochreous or buffy, sometimes without bright tint ;
lower side from white with a faint buff collar on the hind throat
* Formerly this cat was for the most part cited as Felis lybica,
Meyer being regarded as the author of the name, although he admitted
its adoption from Forster. But in 1904 Schwann showed that ocreata
Gmelin, based on Brace's " Booted Lynx," antedated Meyer's lybica ;
and since 1904 the species has been mostly known as ocreata. But
Matschie discovered (SB. Gea. Nat. Fr. Berl. 1912, no. 2 a, p. 59 and
1918, no. 3, p. 102) that in 1780 Forster gave the names constantina
and lybxn to two eats described by Buffon from Constantine in Algeria
and Gafsa in Tunis respectively. Both these names apply to the
common African Wild Cat, for which cafra and caligata are also familiar
terms. I adopt constantina because it has line-priority over h/bica
in Forster's work. J
FELIS. 287
to paler or darker buff almost throughout ; ears typically
ochreous or rusty-brown externally, sometimes blackish
at the tip. Except that the distal end of the tail is always
banded black and white or grey, with a black tip, the pattern
is very variable in distinctness, sometimes practically obsolete,
but usually when absent from the body represented at least
by the brachial bands on the fore legs, by faint stripes on the
hind legs above the hocks, and a few spots on the chest. At its
best in African races it consists of longitudinal stripes on the
head, nape, and back, vertical stripes on the fore flanks,
spots usually tending to run more or less into stripes on the
hind flanks, stronger, more numerous stripes on the legs and
spots below, and two stripes on the cheeks.
The skull is short and broad, with the zygomatic arches
strongly salient in the adult, especially in the orbital region,
the width across the middle of the orbits being much more
than half the total length of the skull. At least in all the
larger races, including the Indian, the upper carnassial has
a large inner lobe with a well-developed cusp.
31 a. Felis constantina ornata Gray. The Indian Desert-Cat.
Felis ornata. Gray, Illustr. Ind. Zool. i, pi. 2, 1832 ; and of most
subsequent authors, including Blanford.
Felis servalina, Jardine, Nat. Libr., Felinse, p. 232 (F. ornata on
pi. 25), 1834.
Felis torquata, Blyth, Proc. Zool. Soe. 1863, p. 185 (in part ;
not of Cuvier*).
Vernacular. — Jhang-Meno (Cutch).
Localities of the types of ornata and servalina, India.
Distribution. — The deserts of Cesttbal and Western India ;
range to the west of Sind unknown.
Distinguished from the African races of constantina, at least
on the average, by the more distinct pattern of spots, more
stony-white or grey superficial colour, less ochreous under-
hair, greyer, less ochreous backs to the ears, and less
differentiated spinal crest.
In the winter coat, which is soft and tolerably full, and
from about 35 to 40 mm. long, the general ground-colour
above is greyish, owing to a silvery band near the tip of the
contour hairs, with the back somewhat darker, but the creamy-
buff summit of the underwool, exposed when the coat is
disturbed, may give a cast of that tint to the pelage. There
is some richer buff on the upper cheek and on the muzzle
in front of the eyes, except on the lips below the rhiuarium,
which are white like the lower cheek ; there is also some white
below the eye and a good deal on its inner side in front ;
* For F. torquata see p. 305, under " Feral Domesticated Cats of
India."
288 FELIDiE.
the limbs and the tail become gradually white distally, and the
whole of the underside, nearly to the tip of the tail, and the
inner sides of the legs are white. The pattern is pronounced,
consisting on the body of solid black spots, which on the side
behind the fore leg tend to form a couple of vertical stripes ;
the forehead is generally spotted, but the crown and nape
have narrow dark stripes ; there are two stripes on the cheek
and on the base of the limbs, and on the tail the spots tend to
run into transverse stripes, which at the distal end of the tail
form complete rings and a black tip. Below there are some
spots on the chest behind the fore legs. The soles of the feet
are black, and on the hind foot the black sometimes extends
nearly to the hock.
When the moult is in progress the contour hairs become
harsh, lose their brightness, and, being less plentiful, expose
the underlying hair, which, even when faded, gives a sandier
hue to the pelage ; the pattern at the same time becomes
faint, and the spots may appear lanceolate.
Flesh-measurements (in English inches) are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Khairpur, Sind ; ad. (J 23£ 121 5
Sambhar, Bajputana ; ad. <J 22 10 4J
Rhoda Motha, Outoh ; ad. 3 19| 11 5-
Sambhar, Rajputana ; ad. <J 18| 10J 5
Sambhar, Rajputana ; just ad. 9 . . . . 18 10 44
The average length of the ear is about 2 J in.
The weight of the adult <$ from Cutch was 7|- lb.
Hitherto this cat has been regarded as a distinct species
restricted to south-western Asia, but its close kinship with the
typically African form Constantino, is beyond doubt. Its
generally paler hues are adaptations to desert life, precisely
comparable to those of the Persian race of the leopard. The ears
may, however, have a decidedly ochreous tinge. The spinal crest
is usually not so noticeable as in many African skins, but it is
very pronounced in the skin of an adult $ collected in June
at Persepolis by Sir J. E. B. Hotson. This skin is in moult,
has only a little underwool and the pattern somewhat obscured,
but the ears are ochreous-tinted. I am doubtful if this
skin is racially distinguishable from typical ornata, but
provisionally it may be given that rank and identified as
F. ocreata nesterovi Birula (Ann. Mus. Zool. St. Petersb. xxi,
Suppl. pp. i-ii, 1916) from Lower Mesopotamia. Birula
assigned this race to F. ornata. In the Persepolis skin the head
and body are 19| in., the tail 12 in., and the foot 5 in.
A closely related northern race, distinguished by its thicker
and longer winter coat, from about 45 to 50 mm. long, and
perhaps by its slightly larger size, is F. Constantino, caudata Gray
FEUS.
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(Proo. Zool. Soc. p. 31, 1874). In addition to the type from
IDiokand, Ferghana, the British Museum has specimens from
Transcaspia, Djarkent, Meshed in N.E. Persia, and Bala
JVturghab in Afghanistan. Possibly this race may extend into
Baluchistan.
In the retention of the spotted pattern and its average
less differentiation into transverse stripes, a specialized
arrangement in all species of Pelidae, where it occurs, ornate
and caudata are more primitive than typical constantina. Prom
this it may be inferred that Asia was the " home " of constantina,
which, like F. chaus, migrated thence into Africa.
According to Blanford this cat ranges from Sind and the
Punjab to Saugor and Nagpur, about 80° E. long., but is very
rare south of the Narbada. I have seen specimens from
Jacobabad (Prater) and Thar Parker (Priestley) in Sind ;
Oambat in Khairpur (Prater) ; Rhoda Motha in Cutch, 100 ft.
(Crump) ; Kishimgala and Sambhar in Rajputana (Adam) ;
Rohtas in the Salt Range, 1,100 ft. (Wells) ; and Sehore in
Bhopal, O.I., 1,600 ft. (Whitehead).
Habits. — Very little is known. Crump remarked that it is
probably commoner in Cutch than is usually supposed, judging
from the numerous tracks observed in quite open country
round the burrows of gerbils. One he shot was feeding on the
fresh carcase of a sheep, but it was not suggested that the cat
killed it. At Rajkot in Kathiawar the cat was also not
often observed. One shot in a patch of grass near a tank
had the remains of a rat and some vegetable matter in its
stomach. No doubt the cat feeds on small desert mammals,
especially gerbils, and on birds. Since the specimen from
Sehore was killed by dogs in scrub-jungle, the habitat
appears to overlap that of the Jungle-Cat.
32. Felis chaus Guldenstadt. The Jungle-Cat.
Felis chaus, Guldenstadt, Nov. Com. Acad. Petrop. xx, p. 483, pis.,
1776 ; Schreber, S&ug. iii, p. 414, pi., 1777 ; and of subsequent
authors, including Blanford, Mamm. Brit. Ind. p. 86, 1881.
Felis catotynx, Pallas, Zoogr. Ross. As. i, p. 23, pi., 1811.
Felis shawicma, Blanford, Journ. As. Soc. Beng. xlv, p. 49, 1876*.
* Felis shawiana, described by Blanford as distinguished from
F. ornata by its apparently larger size, judging from the skull, and by its
shorter tail, judging from two native skins, has been accepted as a valid
species by all students of the Asiatic Felidse except W. L. Sclater,
who examined the specimens in Calcutta and came to the conclusion that
the tails were imperfect and that shavnana is a synonym of ornata (Cat.
Mamm. Ind. Mus. Calc). I am indebted to Col. R. B. Seymour Sewell,
F.R.S.,_for the chance to examine Blanford's material. The species is
composite. The skins are those of young F. constantina caudata (see
above, p. 288) with the tails imperfect, as Sclater detected, and the
skull is the skull of Felis chaus j and since the skull is marked " type,"
I have added shawiana to the synonymy of this species.
MAMMALIA.
PLATE XXIII.
Indian Desert-Cat (Felis constantina omata).
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Jungle-Cat (Felis chaus).
FELIS.
291
Locality of the type of chaus and of catolynx (here selected),
Terek River, north of the Caucasus ; of shawkma, Yarkand.
Distribution. — From the Caucasian and Caspian areas,
Turkestan, and Mesopotamia, thence westwards through Syria
to N. Africa and eastwards through Persia into India, CBXiiON,
Bubma, and Indo-China.
The largest of the existing species of Felis, with the tail
comparatively short, almost invariably less than half the length
of the head and body and less than twice the length of the
hind foot.
A distinct spinal crest and a small tuft on the ears. General
colour above varyingfrom clear grey to tawny or deep brownish,
dependent on the tint of the speckling of the pelage, which is
black and white or buff or ochreous ; spinal crest richer and
darker ; backs of the ears varying from nearly uniformly
ochreous with at most a small black tip, including the "pencil,"
or extensively black at the tip and base and ochreous in the
middle ; the underside sometimes almost entirely white,
Fig. 73. — Skull of adult <J Jungle-Cat {Felis chaus prateri)
from Sind. x J.
except for a broad buffish collar on the hind throat, but
usually the chest behind the fore legs is invaded by buff,
and sometimes most of the underside is rich ochreous-buff
with only the fore throat and chin, the axillae, and the inguinal
region whitish ; cheeks, muzzle, and outside of limbs greyish
or ochreous ; tail typically paler than the body. Pattern
on the head and back absent, on the flanks at most faintly
traceable ; some spots on the chest below and an indistinct
stripe on the throat-collar ; two brachial stripes, varying in
distinctness, on the fore leg. Similar stripes on the hind leg
above the hock ; tail with an irregular blackish stripe above
in its proximal portion, its distal end conspicuously striped
black and white, with tip black. Kitten with pronounced
pattern.
The only external character by which skins of this species can
always be distinguished from F. constantina is the comparative
u2
292 mELiDJS.
shortness of the tail. The skull, too, is similar in all its essential
features, but when fully mature may be distinguished by being a
little more elongated in the muzzle and by being narrower across
the orbital portion of the zygomata, with the cheek lower,
both features due to the less developed lower rim of the orbit.
There is little need, in my opinion, for separating this species
from Felis as a distinct genus, Chans, as has recently been done
by Ognev.
32 a. Felis ehaus chaus Giildenstadt.
Synonymy as above, under the species, -with the addition of FeUs
ehaus typica, De Winton, Ann. Mag. Nat. Hist. (7) ii, p. 291,
1898.
Locality of the type, Terek River, north of the Caucasus.
Distribution. — Turkestan, western and southern shores of
the Caspian Sea, Persia, and Baltjchistak.
Coat in winter thick and long, the contour hairs of the
flanks up to 47 mm., of the crest 69 mm., the average of
four specimens being 44 and 66 mm. Colour of the upper
side varying from pale grey when the speckling is black
and white to tawny or greyish- tawny when the pale speckling
is ochreous-buff or greyish-buff ; the underhair of the back
buff or grey ; the spinal stripe darker, dull or brightish
ochreous.
The winter coat in this race is rather thicker and longer
than in the Himalayan race, affinis, and the skull differs from
that of the four Indian races recorded below in having on the
average larger teeth. The two large upper premolars (pm s
and pm 4 ) measure from 21 to 23| mm., and the three lower
cheek-teeth {pm z , pm^ m^ from 26 to 29 mm., as opposed to
an average of about 20 and 25 mm. respectively in the Indian
races. But the difference is only an average one, some
individuals of the Indian races having teeth nearly or quite
as large as in smaller-toothed forms of the typical race.
Guldenstadt recorded this race from the Persian provinces
of Gilan and Masanderan. In the British Museum there is
a nearly topotypieal example from the Caucasus (Warsaw
Museum) ; four from Persia, one labelled " Persia " (K. Loftus),
without further particulars, and one from Seistan (Indian
Museum, Calcutta) agree tolerably closely in colour with the
skin from the Caucasus, and the skull and teeth of Loftus's
specimen almost exactly match those of the Caucasus specimen.
These three skins have a decided tawny tone, but one from
9 miles south of Shiraz (Sir J. E. B. Hotson) is much greyer.
Still greyer and paler is one from Gursala, Jimfb, 2,000 ft.
(H. R. Sykes). An almost exact match of the last is a clear
ashy-grey " native " skin procured by P. M. Sykes 100 miles
east of Bampur in Baluchistan, 1,400 ft. It is on the evidence
FEUS.
293
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294 FELIDiE.
of this skin, which has no skull, that the typical race of
F. chaus is included in the fauna of British India.
Only two of the above-mentioned skins were measured in
the flesh : —
Head and Hind
Locality and sex. body. Tail. foot.
S. of Shiraz ; ad. <J 27$ 11* 6£
Gursala ; yg. ad. ? 24 10J 5$
To the west of Persia this eat is represented in the British
Museum by specimens from Baghdad, Ramadi, and Mosul,
in which the two carnassial teeth (pm 4 -, m t ) are 16 and 11 \
mm. respectively. These connect typical chaus with furax
de Winton from Jericho, in which these two teeth are respec-
tively 18 mm. and 12 mm. The skull from Ramadi, an adult <J,
with the total, condylobasal, and mandibular lengths 137, 122,
and 93 mm. respectively, is the largest skull of F. chaus
I have measured, and slightly exceeds the skull of the adult <J
from Seistan entered in the table above.
32 h. Felis chaus afflnis Gray.
Velis affinis, Gray, Illustr. Ind. Zool. i, pi. 3, 1830.
Lyndhus erythrotus, Hodgson, Journ. As. Soc. Beng. v, p. 233,
1836.
Felia jacquemonti, GeoSroy, Jaequemont'sVoy. iv, p. 58, pis. 2 &3,
1844.
Locality of the type of affinis, Gangootri in Tehri Garhwal ;
of erythrotus, Nepal ; of jacquemonti, Kursali, 8,500 ft., near
Dehra Dun.
Distribution. — Himalayas, from Kashmir at least to Stkktm.
Distinguished from typical F. chaus by the slightly shorter
and less woolly winter coat and by the skull and teeth being
on the average smaller sex for sex.
In the winter coat the hairs of the flanks range from about
36 to 42 mm., of the crest from 58 to 60 mm., the average
in six skins from Kumaun, September to March, and six from
Gorkha, Nepal, December to March, being 39 and 59 mm.
respectively. There are two extreme, but intergrading,
colour-phases, the " grey " and the " tawny." In the former
the grey of the flanks is paler or darker, in accordance with
the dominance of the black or white speckling in the pelage,
the spinal stripe is speckled black and dull ochreous or buff ;
the cheeks, muzzle, and legs externally are greyish, and the
underside, except for the pale buff on the collar and generaEy
some on the chest, is dominantly white. In the tawny phase
the speckling of the flanks is black and buffy or pale ochreous,
of the spinal stripe black and rusty ; the ears, cheeks, muzzle,
and sometimes the whole of the underside, except the chin,
are brighter or duller ochreous.
FELIS. 295
A handsome blackish variety of this cat, represented by
three skins, was collected by St. G. Burke in the United
Provinces. They differ a little individually, but the finest
of them has the muzzle, crown, backs of the ears, and the
spinal area behind the shoulders jet-black, and the legs and.
tail mostly black ; but the neck and sides of the body are
thickly spangled with silvery-white, and the underhair is
mostly ashy-grey, smoke-grey on the crest. They resemble
the similar variety from Karachi described below (p. 300).
In addition to the co-types of affinis from Gangootri and
Hodgson's specimens of vryihrotus from Nepal, old and soiled
skins, the British Museum has many specimens, recently
collected, mainly by the Mammal Survey, from the following-
localities : —
Kashmir, Arapul, 6,300 ft., Trol, 6,000 ft. (Col. Stockley),
two exceptionally dark, blackish-grey skins ; DharmsaJa,
4,000 ft. (H. Whistler) ; Kangra, 2,000 to 5,000 ft. (Wells) ;
Kumaun, Ramnagar, 1,140 to 1,500 ft., Almora, 5,600 ft.,
Khati, 7,600 ft. (Crump) ; Dehra Dun (Capt. Tuker), nearly
a topotype of jacquemonti, but very pale grey, and further
differing from it in having black ear-tips ; Nepal, many
localities, mostly in Gorkha (Baptista), Darjeeling, 4,500 ft.
(Crump), and Kuivi in the Naga Hills, 6,000 ft. (J. P. Mills),,
a July skin probably belonging to this race.
Flesh-measurements (in English inches) of F. chaus affinis
are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Kashmir, Trol. ; ad. $ 27J llf 6£ 19
Kangra, Damtal ; ad. <J 26f 8f 6 —
Kangra, Gopalpur ; ad. cJ 24f 9f 5 12
Kumaun, Almora ; ad. <J 25f llf 6| 12£
Kumaun, Ramnagar ; ad. (J . 25 llf 6£ 16
Nepal, Gorkha : ad. <J 28f 13J 6* 16
Nepal, Gorkha ; ad. <? 26 llf 6| 12
Kashmir, Hxaful ; ad. 2 25f 10£ 6J 14
Kangra, Gopalpur ; ad. 2 24 9 5f —
Kumaun, Almora ; ad. 2 23£ 11J 5| 11J
Nepal, Gorkha ; ad. 2 24f 10 6 10J
Nepal, Hathiban ; ad. 2 22§ 11£ 5| 11
Darjeeling ; ad. 2 25§ 10| 6 —
The average length of the ear is a trifle under 3 in.
The following table of skull-measurements shows considerable
individual variation in size both in the skull and the teeth,
especially in the $. As suggested by the large skull, without
a skin, from Kashmir (Radclifie), the Jungle-Cat of that area
may prove to be on the average larger than those occurring
farther east in the Himalayas ; but the difference between the
two skulls from Garhwal (B. B. Osmaston), which also have
no skins, is only a trifle less than the difference between the
296
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pelis. 297
larger and the smaller Kashmir skulls ; and the $ skull from
Khatmandu (Dr. Oldfield) is about the same size as the skull
from Arapul, Kashmir (Stockley), and is much larger than the
2 from the Bhaber Tract, Nepal (Dr. T Longstaff).
32 c. Felis chaus kutas Pearson.
Felts kutas, Pearson, Journ. As. Soc. Beng. i, p. 75, 1832.
Vernacular. — Berka (Rajmahal Hill tribe) ; Kliatas, Jangli
billi (Hind.) ; Bariberal (Beng.) ; Mungra (Kathiawar) ;
Jhang Meno (Cutch).
Locality of the type, Midnapore in Bengal, about 70 miles
west of Calcutta.
Distribution. — The northern part of Peninsulas India
from Cutch to Bengal.
Distinguished from the Himalayan race, affinis, by being
a little smaller, shorter in the tail, lighter in weight, and by
its shorter and less luxuriant winter coat, but the seasonal
difference in the coat in those respects is well marked. In
six adult examples from Hoshangabad in full winter coat the
average length of the hair on the flanks is 37 mm., on the
spinal stripe 50 mm.
The type of kutas was a dark- coloured Jungle-Cat ; and
a topotype of it from Salbani, Midnapore, 200 ft. (Crump),
September 19, in new coat, is also very dark, blackish, speckled
with buffy-grey above, with the spinal stripe rusty, the ears
oohreous-brown, with black at the tip and base, the under-
side reddish-buff, with the fore throat, axillae, and groins
white ; the stripes on the legs and spots below are blackish.
A skin from Chaibasa in Singbum, 800 ft., August 11, and one
from Jagodib, Hazaribagh, 600 ft., April 20, are rather paler
than the Midnapore specimen, but like each other ; but
one from Nimia Ghat, Hazaribagh, 1,000 ft., June 26, is not
so grey owing to the shedding of many of the contour hairs
with the moult. A series of seven skins from Bahgownie in
Darbhanga, 1,500 ft., north of the Ganges, July to September,
is inseparable from skins from Lower Bengal, two only being
tawnier. All are short-coated with little under,wool, but
possibly the winter coat may show them to be transitional
to affinis. Many from various spots near Hoshangabad, in
Central India, on the same parallel of latitude as Midnapore,
*md collected in January, February, March, and April, are in
full winter coat, and this is appreciably less luxuriant than in
winter skins of affinis . It is needless to mention all the localities
in the central districts of northern Peninsular India where
this cat was collected by the Survey. But the extension
of the race as far west as Cutch is attested by skins collected
by Crump at Chitrod, 150 ft., Charwa, 200 ft., Bhuj, 350 ft.
298 FELIDiE.
and Nokania, 450 ft., in July, August, and September. Apart
from the Nokania skin, July 7, which is in poor coat and tawnier,
the others closely match the skin from Chaibasa in- their dark
grey hue. Four skins from Rajputana are interesting as
transitional between this race and the next, described from
Sind. One from Kishangarh (Adam), December 26, although
grey in colour, is larger and heavier than any of the Bengal
specimens of kutas. A second from Sambhar (Adam),
February 2, although not measured in the flesh, has a larger
skull than the last, and would possibly have been longer in
the head and body, and its colour is tawny as in the Sind race.
A skin from Bhoria (Adams), October 3, of normal size, is
also tawny ; and a fourth from Mt. Abu (Crump), May 22,
although comparatively small and grey in tint, is a little
heavier than the first described from Rajputana.
Mesh-measurements (in English inches) and weights (in lb.)
of F. cfiaus kutas are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Midnapore ; youngish g 22§ 10£ 6 8£
Ohaibasa ; ad. S 22f 9f 5* 11
Hazaribagh ; ad. <J 23* 10 J 5f 9
Dain, Hoshangabad ; ad. <J . . 25$ 10£ 6 8£
Pachm&rhi, Hoshangabad; ad.
<S 23i 10* 5* —
Nokania, Cuteh ; ad. $ 25| 9§ 6 11£
Kishangarh, Eajputana ; ad. <J 28 11 6£ 12£
Bhoria, Kajputana ; ad. $ • • • 24 10 6 —
Mt. Abu, Rajputana; ad. <J.. 23|- 10 6 13
Hazaribagh ; ad. ? 2l| 9f 5f 6i
Bori, Hoshangabad ; ad. ? . . . 22$ 9£ 5£ 7£
Bhuj, (Dutch ; ad. 9 23f 10 5| &
Charwa, Gutoh ; ad. $ 22 10£ — 5J
The average length of the head and body and the weights
are decidedly less than in affinis ; but the length of the ear,
up to 3 in., is about the same.
32 d. Felis chaus prateri, subsp. nov.
Locality of the type, Jacobabad, on the Upper Sind
Frontier.
Distribution. — Sind, between the Indus and the Kirthar
Range.
Distinguished from typical F. chaus kutas by its more
uniformly tawny or sandy hue at all seasons of the year and
by its larger size, at least of the <J.
This race is based upon a series of six skins from Sind;
three adult <?£ from Jacobabad, February 22-28, an adult $
and a young <J from Naundero, Larkana, May 7 and 8 (S. H.
Prater), and an adult $ from Gholam, October 20 (C. McCann).
FELIS.
299
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Despite the difference of the season, they are all of the general
sandy tint indicated above, although differing to a slight
extent individually, but none of them shows the black and
white speckling, resulting in the darker or lighter grey hue
dominant in unfaded skins of the other Indian races. The
ears on the average also are more uniformly ochreous, with
the black at the tip and base less in evidence.
The coat is not nearly so full and soft as in the Himalayan
race affinis in winter, but is about the same as in the Central
Indian race kutas, the hairs of the flanks in adult skins from
Jacobabad and Naundero ranging from 30 to 34 mm. and of
the crest from 50 to 55 Trim. In the skin from Gholam,
October, the coat is very nearly the same ; but in the young $
specimen from Naundero, May 8, the coat is in full moult,
and in the adult ? from that locality, May 7, it is thinner and
coarser than in the Jacobabad series.
Eepresenting an interesting variety, but no doubt assignable
to this race, is a very beautiful silver and black skin from
Karachi, without further history. It is unmeasured and
undated, but is obviously in full winter coat, the hairs on the
flanks being 40 mm., on the crest 52, and there is abundance
of underwool. The backs of the ears, the crown, part of the
spinal area, most of the tail, the feet, and the backs of the legs
are jet-black, but the cheeks, forehead, neck, shoulders,
flanks, and lower side are conspicuously speckled with silver,
and there is some silvery speckling on the front of the fore leg
and on the root of the tail ; the chin is whitish and the under-
hair is everywhere clear whitish-grey. Apart from this last
character the skin recalls that of a " silver " or " silver-tip "
fox of the furriers.
The flesh-measurements (in English inches) of the adult
specimens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Jacobabad (type) ; ad. (J 29 10 6
Jacobabad ; ad. <J 27J 12£ 6
Jacobabad ; ad. (J 27£ 10| 5£
Naundero, Larkana ; ad. ? 23 10J 5£
Gholam ; ad. 9 23 9$ 5f
The weight of the largest $ was 20 lb., the heaviest, so far
as I am aware, recorded for Felis ckaus. The third on the
list was 14 lb.
32 e. Felis eaaus kelaarti, subsp. nov
Felis chaus, Kelaarfc, Prodr. Faun. Z
Felis affinis, Phillips, Man. Mamm.
Gray).
Vernacvlar .— Kadubekku, Mantbekku, Adaribehku (Kanarese)
Felis ckaus, Kelaarfc, Prodr. Faun. Zeyl. p. 48, 1852.
Fetus affinis, Phillips, Man. Mamm. Ceylon, p. 160, 1935 (not of
Gray).
FELIS. 301
Kadabek, Bellabek (Wadari) ; Jungli Mdmar (Marathi) ;
Baul, Bhdoga (Mahr.) ; Burakatchki (Haran Shikaris) ;
Kebbali (Coorg) ; Jungli-billi (Dekhani) ; Junkapilli (Telegu) :
Walabalala, Hindun diviya (Sinhalese) ; Kadu poona and
Kardup-poonai (Tamil).
Locality of the type, Cheddikulam, N.P., Ceylon.
Distribution,. — Ceylon and Southern India.
Distinguished from the more northern Peninsular Indian
race, F. chaus kutas, by the coat, at its best, being shorter
and less luxuriant and exhibiting very little seasonal difference
in those respects, and by the speckling of the hairs, as a result
of the shorter coat, being finer. In a series of seven adult skins
collected between December and March the average length
of the flank-hairs is 26 mm. of the crest 40 mm., both being
about 10 mm. shorter than the average of kutas in the
same months of the year.
Many skins representing this race and collected by the
Mammal Survey — on the western side of Southern India mainly
by G. C. Shortridge and on the eastern by N. A. Baptista —
are very uniformly coloured dorsally. The colour of the upper
side is dominantly grey, black, and white-speckled, with the
spinal stripe oehreous, the backs of the ears rusty or paler
ochreous, with a very variable amount of black at the tip and
base, the legs typically rich oehreous, the underside with a vary-
ing amount of buff on the chest behind the fore legs. The
pattern is just traceable high up on the flanks, stronger lower
down, there are blackish or buff spots on the chest, and the
upper portions of the legs are more or less distinctly
striped.
The skins were obtained at the following localities and
dates: — Dharwar, 2,500 ft., October, December, January,
April (Shortridge) ; N". Coorg, 3,555 ft., January (Graham) ;
Nilgiri Hills (Phythian Adams) ; Travancore (Pillay) ; Palrri
Hills, 5,000 ft., March (McCann) ; Vontimitta Range, 325 ft.
August, Kurnool, April, May (Baptista).
Of these one skin only, from Dharwar, October, exhibits
buff and black speckling on the flanks, and two from Kurnool,
April and May, have the legs greyish-buff. The darkest grey
of all, with most black on the ears, is a skin from Haleri,
North Coorg. The skin from Travancore, collected at
Bheemanagari, is a kitten exhibiting pronounced pattern on the
flanks.
The type, a subadult <J from Cheddikulam, N.P., Ceylon
(E. W. Mayor), is indistinguishable from examples from
Dharwar and elsewhere in Southern India. Ceylon has been
chosen as the typical locality for the race as being the most
southern country where F. chaus occurs.
302
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KELIS. 303
Flesh-measurements (in English inches) are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Kumool ; ad. <? 25J 10£ 6£
Vontimitta Range ; subad. <J 24£ lOf 6
Dharwar ; subad. <J 24f llj 6
Cheddikulam (type) ; yg. 6* 23 7£ 5f
N. Coorg ; ad. $ 25 9 —
Palni Hills ; ad. 9 24£ 10 5f
Dharwar ; ad. 9 24 10J 5£
The weights of the $ specimens from Kurnool and the
Vontimitta Range were 12 and 11 lb. respectively, of the
young $ from Cheddikulam and the $ from the Palni Hills
81b.
32/. ? Felis ehaus fulvidina Thomas.
Felis affinis fulvidina, Thomas, Proc. Zool. Soc. 1928, p. 834.
Vernacular. — Kyaung-ba (Burmese) ; Kyung tset-hun
(Arakanese).
Locality of the type, Kampong Tomb, Annam.
Distribution. — Annam, Siam, Bubma.
Specimens of the Jungle-Cat from Burma are of doubtful
identity. A good many skins from Lower Chindwin (District
Commissioner), Mt. Popa (Shortridge), and Toungoo (Mac-
kenzie), collected between the middle of August and the end of
October, are short-coated and on the average tawnier in hue
than kutas, none exhibiting the grey tint observable in so many
skins of that race. The teeth, too, are on the average a little
larger. There are also a couple of skins from Siam. I suspect
they will prove to be identical with F. chaus fulvidina Thomas,
based on a single unmeasured skin, without skull, from
Kampong Tomb, Annam (Delacour and Lowe), and dis-
tinguished from Indian specimens by the richer oehreous
hue of the upper side. It is also richer-tinted than the
Burmese and Siamese skins, but this may be only an individual
peculiarity.
The flesh-measurements (in English inches) and weights
(in lb.) of some specimens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Mt. Popa ; ad. <J 29i 10£ 6J 13
Toungoo j ad. J 26£ 10f 6f —
Mt. Popa; ad. 9 23| ll| 5f 8
Toungoo ; ad. 9 23£ 10£ 5J —
, The skull of the large <J from Mt. Popa is mislaid ; but the
eondylobasal length of the $ from Toungoo is 108 mm., of the
adult $ 105 mm., and of the $ from Mt. Popa 102 mm. The
304 FELIDJE.
upper carnassial (pm i ) in the $ from Toungoo is 15 mm.,
and in a skull from Lower Chindwin (14±) ; in £ skulls-it is
from 13 to 13J mm., and the lower carnassial (m x ) is from
10 to 10£ mm. Hence these teeth are as large as in typical
F. chaus chaw. But in a specimen from Koh Lak, Siam
(Malcolm Smith), these same teeth are 12 and 10 mm.
Habits. — The habits of the Jungle-Cat seem everywhere to be
the same. It is the most familiar of the wild eats of India, and
is found in bush-jungle, long grass, and reed- beds near rivers,
lakes, and marshes, mostly in comparatively dry zones,
but in Kashmir, according to Col. Ward, it also occurs among
rocks and in old buildings, nearly every old building about
Srinigar, where it is plentiful, being occupied by it. The
Mammal Survey collectors recorded it as plentiful in suitable
localities all over the country, and testified to the frequency
with which it was to be seen hunting by daylight. According
to Shortridge its long legs give it a very distinctive appearance,
its movements in the open recalling those of a small panther.
He added that it is very swift and exceedingly strong for its
size, and is probably capable of pulling down quite large game.
It seems, however, to prey mostly upon smallish mammals
and on birds up to the size of pea-fowl, and may be a great
pest to_ poultry owners. The discovery of quills in the feet
of specimens in Kumaun led Crump to believe that it may
prey upon porcupines ; at all events it seems that it may
attempt to do so. But it also eats frogs when pressed by
hunger, according to Ward, and Pallas long ago stated that it
is a fish-eater.
There are no good reasons for doubting Blanford's statement
that the Jungle-Cat breeds twice a year, but his authority
is unknown.
In Ceylon, according to Phillips, it is restricted to the dry
zone of the northern part of the island, but is comparativelv
rare *.
The Feral Domesticated Gats of British India.
Under the genus Felts notice must be taken of the domesti-
cated cats which have run wild in all parts of British India,
have established themselves in some localities as obtrusive
elements in the fauna, and have been described as representing
valid species.
™t ^l 6 . 18 ' h0 Y? ver > a skin in the British Museum, collected by
Whyte between Handy and Colombo. It was always regarded as
£. chaus, but from its dark hue and unusually distinct pattern I think
it is probably a hybrid between chaus and a Domestic Cat. But, even
so, if the locality is correct, an example of chaus was probably there
EBLIS. 305
The dominant type is the eat I called " the Striped Tabby,"
which is a common house- cat in England and has run wild
in nearly all the warmer parts of the world (Proc. Zool. Soc.
1907, p. 143). It was described from Nepal as F. forquata
by Cuvier (Hist. Nat. Mamm. pi. 126, 1826), as F. inconsjoicua,
also probably from Nepal, by Gray (Charlesw. Mag. Nat. Hist,
i, p. 577, 1837), as huttoni, from Kandahar, by Blyth (Journ.
As. Soc. Beng. xv, p. 169, 1846) *, and Blanford treated it
provisionally as a valid species under the name torquata.
The colour is typically darker or lighter grey, sometimes with
a tawny, more rarely a '' bluish " tinge, often with white
patches on the paws, and the pattern is usually distinct,
consisting of narrow longitudinal dorsal stripes and of vertical
lateral stripes commonly running into spots posteriorly, with
bands on the legs and the end of the tail. These are obviously
similar to the Desert-Cat, but not so pale. Col. Ward wrote
of them, under the name torquata, as plentiful in Kashmir and
unmistakable for any other species. The British Museum
has skins from Srinagar in Kashmir, Kangra, 5,000 ft., Gan-
gootri in the United Provinces, Nepal, Sambhar in. Rajputana,
Karachi, Junagadh in Kathiawar, N. Coorg, S. Mysore,
Ambawela, Ceylon, 5,090 ft., and Maymyo, 3,500 ft., the
N. Shan States, 2,795 ft., and Toungoo, 100 ft., in Burma.
A red cat with the same style of pattern also occurs, as
attested by skins from Sind, the United Provinces, and
Pilibhit, 800 ft. There are also red skins showing scarcely
any body-pattern, except very small faint spots on the flanks,
from Nepal, Kumaun, and Mogul Sarai.
A different style of feral cat, called the " Chaus-type "
by Blyth, who believed it to be a hybrid between tame cats
and Jungle-Cats, represents what English cat-fanciers call
the "Abyssinian breed." It is " pepper and salt " in colour,
without trace of pattern except stronger or weaker stripes on
the base of the legs and tail-tip. It recalls F. chaus in colour,
except that the spinal stripe is not so differentiated, but it is
smaller, and has the tail much longer. It may sometimes
be seen in the streets of London and elsewhere in England,
and since the skulls of the Indian specimens I have seen are
those of typical tame cats, the evidence that the coloration
is due to an infusion ofF. chaus amounts to very little. There
are skins of this type in the British Museum from Darjeeling,
Bhutan, Hazaribagh, the Vontimitta Range, the Mishmi Hills,
2,250 ft., and Toungoo, 500 ft.
Finally, a black feral cat from Victoria Point, Tenasserim,
has a very short tail like many Domestic Cats from Malaya.
* Blyth later thought huttoni %ras the same as ornata (Cat. Mamm.
As. Soc. 1863, p. 63).
VOL. I. X
306 felid^;.
Genus CARACAL Gray.
Caracal, Gray, List Mamm.Brit. Mus. p. 46, 1843 ; Proc. Zool. Soc.
1867, p. 27 ; Fitzinger, SB. Akad. Wiss. Wien, lx, p. 24, 1869.
(Felis, Lynx or Caracal of other authors.)
Type of the genus, melanotis* Gray (=Felis caracal of
earlier authors).
Dtsirifarficm..' — The whole of Africa, except the forested area,
from Morocco to the Cape, S.W. Asia, and the northern part
of Peninsulas India, coinciding very closely with that of the
Lion.
Distinguished from Felis externally by the presence of
a long tuft on the tip of the ear, typically exceeding in length
half the height of the ear, by the shallower interdigital webs,
especially of the hind feet, which are very similar to those of
the Hunting Leopard or Cheetah, except for the presence of
the claw-sheaths. Also there is no trace of pattern, except
a few spots on the underside and inside the fore legs, even
in newly-born kittens. In the skull the nasal branch of the
premaxilla is narrower, longer, more attenuated, and passes
further between the nasal and the maxilla, so that the junction
between those two bones is much shorter ; also there is no
abrupt notch on the posterior edge of the palate on the inner
side of the last upper molar, the postcanine space in the jaws
is relatively shorter, and only very occasionally is the minute
upper premolar retained.
33. Caracal caracal Muller. The Caracal.
Felis caracal, P. L. S. Muller, Syst. Nat., Suppl. p. 30, 1776 ;
Guldenstadt, Nov. Comm. Acad. Petrop. xx, pp. 499-500,
1776 ; Sohreber, Saugth. iii, pi. 110, 1776 ; p. 413, 1778 f-
General form like that of Felis, but the hindquarters better
developed and the tail shorter, though only a little, than in
F. chaus, reaching to the hocks, and about one-third the length
of the head and body or a little more. Coat and colour varying
greatly in accordance with distribution and habitat, and also
seasonally, in the same locality. The coat sometimes smooth
* It seems expedient to follow Matsehie, 1912, and Thomas, 1926,
in considering that Gray substituted melanotis for the older specific
name caracal because he disliked identity between generic and specific
names, and did not intend to restrict the name melanotis to the specimen
described by Sohreber, which came from the Cape of Good Hope.
Schreber also did not restrict the name caracal to the S. African race
which in 1926 Thomas described as colonite.
t Although Muller, Giildenstadt, and Schreber published this name
in the same year, I adopt Matschie's decision to give priority to Muller,
who cited Arabia as the type-locality two years before Sohreber described
Hie animal from the Cape of Good Hope. Giildenstadt mentioned no
locality, and some authors question the " binominal " status of his text.
CARACAL. 30T
and sleek, with no wool, sometimes long, very furry, with,
abundance of wool. Colour varying from pale sandy fawn
to reddish, deep brown or dark grey, but always tolerably
uniform above and on the outside of the limbs and on the
tail, except that the tail-tip may be darker than the rest.
The colouring of the head is characteristic, the ears are
typically black, frosted with white externally, with white
hairs internally, and the " pencil " or tuft is composed of
black and grey hairs ; but the lower eyelid is white, and there
is a whitish patch on the inner side of the eye in front ; the
chin and the upper lip below the rhinarium are white, but there-
is some black above the corner of the mouth where the mystacial
vibrissse rise. The underside, except for a buff collar on the
hind throat, and the inside of the limbs are whitish, but the
chest at least is marked with spots and the inside of the fore legs
with stripes which may be pale buff in light skins or deep brown,
nearly black, in dark skins.
33 a. Caracal caracal earacal Muller.
Felis caracal, P. L. S. Muller, Syst. Nat., Suppl. p. 30, 1776.
Felis caracal bengalensis, Fischer, Syn. Mamm. 1829, p. 210
(nom. preocc. in Felis).
Caracal caracal schmitzi, Matsehie, SB. Ges. Nat. Fr. Berlin, 1912,
p. 64.
Caracal caracal aharonii, Matsehie, torn. cit. p. 66 *.
Vernacular. — Siyeh Gush (Pers. and Hindi) ; ? Tsogde (Little
Tibet) ; ? Ech (Ladak).
Locality of the type of caracal, Arabia ; of bengalensis,
Bengal ; of schmitzi, the Dead Sea ; of aharonii, mouth of the
Chabur River on the Upper Euphrates.
Distribution. — Central India, the Punjab, Sind, Balu-
chistan, and thence westwards at least to Arabia and
Palestine.
As native names for the Caracal in Little Tibet and Ladakh,
respectively, Blanford cited the two queried above. These
were taken from Blyth (Joura,. As. Soc. Beng. xi, p. 759,
1842) on information supplied by Vigne, who had a good
drawing of a trained Caracal he had seen in Little Tibet.
Blanford suggested that Vigne's specimens were tamed
animals, presumably meaning that they were imported.
In my opinion that supposition was correct. But Col. Ward
(Journ. Bomb. Nat. Hist. Soc. xxix, p. 28, 1923) said Vigne
* The names schmitzi and aharonii, given to Caracals from the Dead
Sea and "Upper Euphrates respectively, are entered as synonyms of
typical caracal because I am unable to distinguish racially skins and
skulls of British Indian specimens from Persian, Sinaitie, and Arabian
specimens, Arabia being the type-locality of caracal Muller.
x2
308 FEIIDJB.
was most likely right in saying that the Caracal was found
on the Upper Indus in Baltistan, adding that he knew of a skin
in Srinagar, alleged to have come from Ladakh, which the
Balti men called the " Bch." On the other hand, Col. C. H
Stockley wrote to me : — " I am quite sure that the Caracal
does not exist in Kashmir proper. I have been on the look-out
for a skin for over thirty years, and have never seen one in the
Srinagar skin-shops. It is essentially an animal of hot, dry
country, and probably exists in the low ground round Jammu,
which is under the same rule as Kashmir proper. I have
seen and shot it in the Salt Range west of Jhelum." If the
Caracal occurs in Ladakh, it will surely prove to be a race
distinct from the Caracal of the plains of India. But it must
be remembered that the Lynx in its short, reddish, summer
coat (see p. 312) is decidedly Caracal-like, and that its native
name " Ec " (Ward) is apparently a variant of " Ech."
Coat in winter with a considerable amount of underwool
and tolerably long, from 30 to 35 mm. ; in summer short and
sleek, about 10 mm. long, and with little, if any, wool. General
colour variable seasonally. An undated winter skin from Sind,
apparently in fresh coat, is rich reddish-fawn above, the
hairs paler at the base, but with the extreme tip blackish,
and a whitish subterminal band, giving a slightly grey cast to
the back ; ear with hardly any white frosting ; outer side of
limbs and upper side of tail about the same colour as the body ;
the underside and inner side of the limbs whitish except for the
pale fawn hind throat; a few pale spots on the chest and some
similar marks on the fore leg, one representing the normal stripe ;
head like the back, with the typical dark and light patches, the
•cheeks blending with the dark crown and light throat. A skin
from 100 miles east of Bampur, Baluchistan (P. M. Sykes),
March 10, in faded winter coat, is much paler sandy fawn
above, with the underwool of the middle of the back slightly
darker than the surface colour. A $ skin in summer coat
fromDhonsa, Cutch, 200 ft. (Crump), July 20, is pinkish-grey
at a little distance, darker down the middle line of the back,
the grey cast being much more evident owing to the pale band
of the flat-lying coat concealing to a greater extent the fawn
of the hair, and the ear is extensively frosted with white.
A skin from Dizful, Persia (K. Loftus) is like the Baluchistan
skin, but a shade browner and less red. One from the moun-
tains north of Tor, Sinai (Anderson), and one from El Arish,
N. Sinai (Col. Stirling), differ in almost precisely the same way
as the skins from Sind and Cutch, and are merely trivially
distinguished from them. One collected near Mecca (H.
Philby) and one from Aden (Percival and Dodson), September
skins, are like the paler, greyish sandy skin from El Arish,
Sinai, the Aden skin being a trifle the darker of the two.
PLATE XXIV.
MAMMALIA.
Photo F. W. Bond
Caracal (Caracal caracal).
Phoco F. W. Bond.
Lynx (Lynx lynx isabellinus).
CARACAL. 309
The following are the flesh-measurements (in English
inches) of some specimens assigned to this race : —
Head and Hind
Locality and sex. body. Tail. foot. Ear.
Punjab (Col. Ward) ; <J 29 9 — —
Western Dun, TJ.P. (Col. Ward) ; <J 27* 9 — —
Cuteh (Crump) ; ad. £ 27£ lOf 7i 3 i
Mirzapore (G. O. Allen) ; ? sex 27 7 — —
Near Mecca (Philby) ; subad. ? . . . 25£ 9 6-J 3
The Mirzapore specimen is interesting for the shortness
of the tail, which disproves the contention of Fischer and
Matschie that the Indian race is characterized by its long tail *.
The measurements of the just adult skull, the only one
available from British India, are entered in the table (p. 314).
They are very nearly the same as those of an adult <J from
El Arish in Sinai.
Habits. — Since Crump, who was collecting for the Mammal
Survey in many localities in the plains of Northern and
North-Western India, where the Caracal might be expected
to occur, secured only one specimen, the species is clearly
very rare at the present time. This is borne out by the
testimony of others, who report the killing of a Caracal as
an unusual event. Although there seems to be no evidence
that it was ever plentiful in the past, it will in all pro-
bability follow the fate of the Hunting Leopard or Cheetah
and cease to be a member of the British Indian fauna
unless strict measures can be enforced for its protection.
It is essentially an inhabitant of more or less open or bush-
country, and preys upon any mammals or birds it can overcome
and capture. It is one of the most active of the wild cats
both in running and leaping, and, although a savage and
desperate fighter when cornered, is capable of being tamed if
captured sufficiently young. Taking advantage of these
qualities, Indian potentates formerly trained the Caracal
for hunting small game of all kinds in much the same way that
they make use of the Hunting Leopard for chasing antelope.
It must not, however, be supposed that the speed of the
Caracal, although surpassing that of most cats of its size,
is comparable to that of the Hunting Leopard, or that it
pursues running prey for a long distance. Its methods of
capturing prey are essentially the same as those of ordinary
cats, and it is possibly not more active than the Jungle-Cat,
making allowance for its superior strength.
* Bufion's " Caracal of Bengal," upon -which Fischer's F. caracal
bengalensis is supposed to have been based, was described from a drawing
sent to the French naturalist by Edwards. The ears depicted are those
of a Caracal, but the tail, "reaching the ground" (Desmarest), is as-
long as that of the Indian Desert-Cat.
310 FELIDiE.
The cubs, two or three, possibly occasionally more in
number, resemble their parents in colour. The spots on the
underside are typically a little more distinct, but there is no
trace of pattern on the body as there is in the kittens of the
Jungle-Cat, which may be almost " self-coloured " when adult,
and in the cubs of lions and pumas.
Genus LYNX Kerr.
Lynx, Kerr, Anim. Kingd. Cat., between pp. 32-3, and p. 155, 1792 ;
Miller, Cat. Mamm. West. Europe, p. 470, 1912 (with several
generic synonyms).
Type of the genus, vulgaris Kerr (=Felis lynx Linn.).
Distribution. — Europe, and Northern Asia to the limit of
tree-growth, Central Asia as far south as about lat. 30° N. ;
JN. America as far south as Mexico *.
Fig. 74. — Skull of adult <J of the Tibetan Lynx {Lynx lynx isahelMwus)
from Gilgit. X J.
Distinguished from Caracal by its much shorter tail, about
one-seventh the length of the head and body, by the presence
of a bushy ruff or fringe passing over the cheek from the ear
to the throat and by the retention of the white patch on the
back of the ear and typically of the pattern on some part of
the upper surface, legs or tail, the pattern, when well denned,
consisting of narrow stripes on the head and back, rounded or
lanceolate spots on the flanks, spots on the legs, and stripes
on the terminal part of the tail. The skull and teeth are
essentially the same as in Caracal, but the external pterygoid
crest, which maybe strong in well-developed skulls of Caracal,
is apparently only represented by an inconspicuous ridge.
* There are two well-defined species in the Old World, L. lynx of
Northern and Central Europe and Asia and L. pardeUus of Spain,
Sardinia, and ? Asia Minor, and two in the New World, a northern,
L. canadensis, and a southern, L. rufus. But L. canadensis is probably
only a subspecies of L. lynx.
LYNX. 311
In the shortness of the tail Lynx is more specialized than
Caracal, but in the retention of the pattern and the colour of
the outside of the ear it departs less from the normal feline
type.
34. Lynx lynx Linnaeus.
Felis lynx, Linn., Syst. Nat. 10th ed. i, p. 43 ; Miller, Cat. Mamm.
West. Europe, p. 471, 1912.
Locality of the type, near Upsala, Sweden.
Distribution. — Northern and Central Europe and Asia,
Asia Minor.
As may be seen from the synonymy cited by Miller, a large
number of names, based to a great extent upon seasonal
differences in colour, have been given to the Lynx of Northern
and Central Europe. In the typical race, Lynx lynx lynx,
from Scandinavia, the colour varies from greyish-white,
either with distinct black spots or hardly any pattern, to
brownish with only a few white-tipped contour hairs, and the
pattern definite or obscure. The colour of the underhair
is deep rusty-ochraceous or reddish-brown, darker on the
back than on the sides, and in winter the hairs on the soles
of the feet are long and completely conceal the pads.
34 a. Lynx lynx isabellinus Blyth. The Tibetan Lynx.
Felis isabeUina, Blyth, Journ. As. Soc. Beng. xvi, p. 1178. 1847 ;
id., Proc. Zool. Soc. 1863, p. 186.
Felis lynx, Scully, Proc. Zool. Soe. 1881, p. 201 ; Blanford,
Mamm. Brit. Ind. p. 89, 1888.
Lynx lynx wardi, Lydekker, The Field, civ, p. 576, 1904.
L/ynckus isabellinus Jcamensis, Satunin, Aim. Mus. Zool. Acad.
Sci. St. Petersb. ix, p. 13, 1904*.
Vernacular. — Patsalan (Kashmir) ; Phiauku (Lahul) ; Ec
(Baltistan) ; Jungle BUM (Dun).
Locality of the type of isabellinus, Tibet ; of wardi, Altai
Mts. ; of Jcamensis, Kam, S.E. Tibet.
Distribution. — Tibet ; Kashmir ; Kuen Lun Mts. ; Turkestan,
Thian Shan, and Altai Mts.
* Scully and, following him, Blanford quite correctly considered
the characters upon which Blyth separated this Lynx from the typical
Scandinavian Lynx to be not of specific importance, as Blyth thought.
But since they occur in the skins I have examined they are worth
distinguishing as of subspecifie significance. The type of wardi, a skin
with no skull, in the British Museum, is indistinguishable from the skin
of isabeUirms from Ishkoman, Gilgit, both being copiously silvered,
with bright bufi or oehreous-buS wool, except that the obscure pattern
is brownish or fuscous instead of blackish. This is a difference of no
importance. Hence Rasewig (' Semja Ochoton,' Moscow, no. 5, p. 106,
1908) correctly recorded isabellinus from the Altai. I earn find nothing
in the description of Jcamensis to justify regarding it as racially distinct
from isabellinus.
312 FELIBiE.
Distinguished from the typical Scandinavian race by its
paler, more " isabelline " underhair on the average, by the
exposure of the pads in the winter coat, and apparently by its
rather smaller skull.
The colour is variable seasonally and independently of the
season. In several specimens observed in the Zoological
Gardens, two of them captured in the Dharmo Valley, 10,000 ft. ,
the short summer coat was rufous-tawny, with the back rather
darker than the flanks, whereas in the longer, fuller winter
coat the exposed portions of the hairs were extensively white,
giving a decidedly grey aspect to the pelage, especially on the
flanks. The distinctness of the pattern varied individually
at all seasons *.
The individual variation in the colour and pattern of the
winter coat is well illustrated by four skins received from
Col. W. R. F. Treveiyan from the Gilgit district. One
from Rostan, 10,500 ft., November, and one from Nagar,
January, have the upper side mostly silvery-grey, varied by
buffy patches, where the hairs are parted, and the pattern,
consisting of black stripes dorsally and of black spots on the
flanks and legs, stands out in bold relief. Another, from
Ishkoman, December, is like the preceding in its grey hue,
but the pattern is not black, is scarcely traceable dorsally,
and is fuscous or brown on the legs. A young specimen,
Gilgit, 6,500 ft., December; is not nearly so silvery as the last
three, tawnier on the back aDd the fore quarters, with the head
tawny and the pattern everywhere obscure. Another, from
Gilgit, 5,000 ft. (Major Biddulph), December, is very like
the last in the tawny hue of the back and silvered flanks and
limbs, but the pattern is everywhere distinct, though not so
distinct as in the skins from Rostan and Nagar. An undated
skin from Yassin (Dr. Scully) is speckled grey and black and
is similar to the skin from Ishkoman, Gilgit, but is more
ochreous everywhere owing to the exposure of the wool
by the moulting of the contour hairs, and the pattern is
nowhere appreciable. Major D. G. Lowndes (Jotirn. Bomb.
Nat. Hist. Soc. xxxiv, p. 234, 1931) saw at Lahul two adult
specimens, in summer coat (August), which he described as
" bright orange-red " and three half -grown cubs which were
grey-brown, one showing signs of turning red.
Two skins from Tibet (Hodgson) vary similarly. One
is brownishabove, due to a blend of the black and buff of the
contour hairs, with some silvering only low down on the
* These observations bear out Blyth's description of the differences
between, the summer and winter coats. His skins, said to have come
from Tibet, were sent to him by a Mr. Lushington, who lived at Almora
in Kumaun. Possibly, therefore, they were from " Little Tibet,"
which is part of Kashmir.
LYNX. 313
flanks ; the pattern is black, but rather confused and less
conspicuous than in the skin from Rostan, Gilgit, owing to the
darker ground-colour. The other is greyer, but has no pattern.
A skin from Kuen Lun Mts. (Capt. Wellby), August, in
short summer coat, is tolerably uniformly tawny-red, without
pattern, and differs from the silvery spotted Gilgit skins exactly
as the reddish European skins differ from the silvery spotted
skins.
The seasonal variation in the length of the coat (in mm.)
in three skins is as follows : —
Cheek- Ear-
Loeality and season. Back. Belly. fringe. tuft.
Rostan, Gilgit (Nov.) 40 76 75 73
Tibet (winter) 40 89 65 50
Kuen Lun (summer) .... 30 42 50 45
None of the skins examined was measured in the flesh,
and I am not aware that any recorded dimensions were so
taken. Blanford gave the head and body as 33 in. and the
tail as 7§ in., and Col. Ward recorded a <$ from Changehemno
with the head and body 34 in., the tail 8 in. ; but according
to him the length of the head and body may be 46 in. This
dimension must, I think, have been taken from a stripped and
stretched skin. The measurements (in English inches) of
three skins in the British Museum are as follows : —
Head and
Locality. body. Tail.
Bostan, Gilgit 36 5J
Ishkoman, Gilgit 42 6|
Tibet 40 7 (nearly)
In Blanford's and Ward's records the tail is exceptionally
long for the head and body, and probably included the terminal
hairs. The three skins in the British Museum are all probably
stretched more or less in the head and body. Col. Ward's
other specimen, if measured in the flesh, was a huge Lynx,
as long as some adult Panthers. If taken from a stripped skin
the record has little value. I have seen many living adult
examples of the Lynx ; but none was nearly so long as a
normal Leopard.
The weight, according to Blanford, may be 60 lb.
The immature skull of the specimen, no doubt a 9> from
Nagar, Gilgit, has smaller teeth than those measured, pm A
being 17 and m x 12§ mm. Two supposedly <J skulls of typical
Lynx lynx from Sweden, recorded by Miller, had a condylo-
basal length of 143 and 144 mm. and a mandibular length of
109 and 108J mm. respectively.
Habits. — Of the habits of this Lynx very little has been
recorded except that it has been stated to inhabit more open
314
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OTOCOLOBUS. 315
and rocky districts than the typical form from northern
Europe. To this Blanford attributed the more naked soles of
its feet. Col. C. H. Stockley, however, informs me (in litt.) that
it is " nearly always found in country where cover of dense
vegetation is available, such as the willow-scrub in the Shyok
Valley and patches of Tibetan gorse and grass in Bupshu.
I have seen it at 9,000 ft. in June and at 18,000 ft. in July and
August, so its range in elevation is great. It was said by the
local people not to exist in the Mahomedan country forty
miles below the junction of the Nubra with the Shyok, where
the river begins to flow through a big and steep gorge, and I do
not think it is anywhere partial to rocks and steep ground."
It no doubt preys upon all the mammals it can overcome,
from wild goats and sheep, especially the females and young,
to hares and mouse-hares, as well as upon pheasants,
francolins, and other birds.
Lowndes's report that he saw three half-grown cubs in August
suggests that the litter consists of from about two to four,
and. that the young are born early in the year. Stockley also
saw a half-grown litter in early August.
Genus OTOCOLOBUS Brandt.
Otocolobiis, Brandt, Bull. Acad. Sei. St. Petersb. ix, pp. 37-9, 1841
(1842) ; Severtzow, Rev. Mag. Zool. (2) x, pp. 286-90, 1858 ;
Pocoek, Proc. Zool. Soc. 1907, p. 299, figs. ; Birula, Ann. Mus.
Zool. St. Petersb. xxi, p. 155, 1916 ; Ognev, Mamm. of USSR,
etc., iii, p. 174, 1935.
Trkhselurus, Satunin, Ann. Mus. Zool. St. Petersb. ix, p. 1, 1905 ;
Pocoek, Ann. Mag. Nat. Hist. (8) xx, pp. 335 & 349, 1917 *.
Type and only species of the genus, Felis manul Pallas.
' Distribution. — Central Asia from Transcaspia to Mongolia
and Kansu and southwards to Afghanistan, N. Persia,
Baltjohistak, Kashmib, and Tibet.
Distinguished from other genera of the Felinse in external
characters by having the short, slightly-angled ears set very
wide apart and low down on the sides of the head, so that their
inner rim hardly rises above the transverse plain of the fore-
head, which is exceptionally wide ; and also by the pattern
on the body, when visible, being represented by a few narrow-
transverse stripes restricted to the loins. The skull also has
many peculiarities. It is very wide in its cranial portion
and has the muzzle very short and steeply sloped, so that the
* Satunia introduced this name under the impression that Otocolobics
■was preoccupied by Brandt for a Ground- Squirrel (CitelVus). He was
misled by Palmer (Index Gen. Mamm. i, p. 487, 1904). Brandt, in
1844, accidentally substituted Otocolobiis for Colobotis, which he had
applied to the Rodent in question.
316
FELIDiE.
upper margin, of the anterior nares is about on a level with the
inferior edge of the expanded orbits ; there is no noticeable
excrescence adjoining the infraorbital foramen, and the
temporal ridges are widely separated ; the outer chamber
of the bulla is exceptionally large, as capacious as the inner,
the line of the partition running from the stylomastoid
Fig. '5.— A. Upper view of skull of Pallas's Cat (Otocolobus manul) from
Ladakh. B. Left upper camassial (pm*) of the same, showing
reduction in size of the inner lobe. C. The same of the Indian
Desert-Oat (Felis Constantino, ornata), showing the normal size of
the inner lobe in Felis. A, nat size ; B and C, enlarged.
foramen across the summit of the bulla to the occipital suture*.
A peculiarity of the dentition is the reduction of the inner
* For the structure of the bulla in this and other genera of Feline
see my paper in Ann. Mag. Nat. Hist. (8) xviii, p. 328, 1916.
OTOCOLOBTTS.
317
lobe of the upper carnassial, hardly more than its root per-
sisting ; also the normal small upper first premolar is absent.
Prom the general form and detailed structure of the skull
it may be inferred that Otocolobus is a very specialized form
of Felis exemplified by the European Wild Cat (F. sylvestris)
and by its near ally F. constantina, of which ornata is the
British Indian representative.
35. Otocolobus manul (Pallas). Pallas's Cat.
Felis mamd, Pallas, Reise Buss. Reichs, iii, p. 692, 1776, id., Acs.
Acad. Sci. Petrop. pt. 1, p. 296, 1781 ; and of most subsequent
authors till 1905, including Blanford, Mamm. Brit. India, p. 83,
1888.
Otocolobus or Trickeelurus manul of most authors since 1905 or
1907.
Locality of the type, Dschida River, S. of Lake Baikal.
Distribution. — As recorded under the genus.
Size that of a small Domestic Cat, with the tail longish,
nearly half the length of the head and body ; the coat long,
Fig. 76.-
-Side view of skull of Pallas's Cat (0. manul) from Laclakh.
Nat. size.
full, and soft, and the tail bushy, hardly tapering at the point ;
general colour above varying from silvery-grey to whitish-
tawny or reddish, below with the chin, fore throat, and belly
white, the hind throat and chest dusky or brownish ; tail,
apart from its stripes, rather paler than the back : the legs
like the flanks above, but becoming buffy to ochreous on the
paws, which are either black or buff beneath ; the pattern is
very variable in its distinctness, at its best consisting of
crowded black spots on the crown and a few, mostly short,
318 ffELIDJE.
vertical stripes on the hind back and loins, the longest and
most conspicuous being the first, set mid-way between the
shoulders and the root of the tail ; there may be some indistinct
spots on the thighs, but the stripe on the fore leg below the
elbow is always conspicuous ; the tail may have about six
stripes ; three in its proximal half are narrow and widely spaced
above, but expand below, and three in its distal half are broad
and closer together and expand and generally unite below,
the last fusing with the black tip, so that the underside of the
terminal part is mainly black ; the backs of the ears are
greyish or fawn-grey, there is some buff on the muzzle, white
round the eyes, surrounded by a narrow black rim, giving
a " spectacled " appearance to the face, and two black stripes
on the cheeks, with a whitish area between them, run back
to fuse with the dusky patch on the fore throat. The pattern
of the face is very clearly disruptive.
The typical northern race, according to Birula, has the
winter coat " flavescent " white, with very little black in it,
the wool " rufescent," the head greyish-white above and not
densely spotted with black, the hind back marked with some
more or less conspicuous or almost obsolete stripes, and the
tail obsoletely striped. The only skin I have seen, from the
Kirghiz Steppes (Brandt), has the head distinctly spotted,
but there are hardly any stripes on the hind back.
Satunin admitted two species of Tricheelurus, a northern,
T. manul, ranging from Tartary through Siberia to the east
of Lake Baikal, and a southern, T. nigripectus, found in Tibet
and Kashmir. In 1907 I suggested that the latter was at
most a subspecies of the former : and that view was adopted
in 1916 by Birula, who mentioned some particulars in the colour
and pattern of the winter coat and some cranial characters
by which the two might be distinguished. It does not seem
that he had many specimens, if more than one, of nigripectus
to judge from ; and the material of this cat at my disposal
shows that the colour and pattern of nigripectus are more
variable than he supposed, and that the alleged cranial
differences, on which he and, following him, Ognev relied,
do not exist*. Since, however, it is highly probable that the
Pallas's Cats of two such widely separated areas are racially
distinct, I have adopted that opinion.
* He claimed that in typical manul the nasal bones are gradually
narrowed from front to back, and that the presphenoid is expanded in
front, i. e., not overlapped by the palatine bones forming the adjacent
floor of the mesopterygoid fossa, whereas in nigripectus the nasals are
strongly constricted a little behind the anterior nares and the pre-
sphenoid is narrow in front. In an adult skull of nigripectus in the
British Museum the nasals are only slightly constricted and the pre
sphenoid is expanded in front.
OTOCOLOBUS. 319
35 a. Otocolobus manul nigripectus (Hodgson).
Veils nigripectus, Hodgson, Journ. As. Soc. Beng. xi, p. 276,
pi., 1842.
Otocolobus nigripectus, Satunin, Ann. Mus. Zool. St. Petersb. ix,
P . 9, 1905.
Trich&lurus manul nigripectus, Pocock, Proc. Zool. Soc. 1907,
p. 302 ; Birula, Ann. Mus. Zool. St. Petersb. xxi, p. 133, 1916.
Locality of the type, Tibet.
Distribution. — Tibet and Kasemeb.
Distinguished, according to Birula, from the typical northern
race by the winter coat being silvery-grey, with more black in
it, the wool paler, the head thickly spotted with black, and
the stripes on the hind back and tail more conspicuous.
The few skins in the British Museum show individual
variation. One from the Yamdok Lake, Tibet, 15,000 ft.
(Col. Waddell), September, is silvery-grey, the contour hairs
being white distally, with a small black tip ; the under-
hair is ochreous and shows beneath the contours, especially
on the flanks ; the hind back has a few widely spaced stripes,
and the tail, which is not so silvery as the back, has six or seven
black, annuliform stripes and a black tip ; the crown is darker
than the back and covered with black spots ; the fore leg
has the paw ochreous above, black below ; the hind leg is
cream in front, ochreous behind below the hock, with the
paws black below, and there are some indistinct stripes on the
outside of the thigh.
One of Hodgson's Tibetan skins, with the head missing,
is like the last, but the soles of the paws are buff. Another
differs in having much less silvery speckling, owing to the
moult of most of the contour hairs ; the nape and the top of
the head are largely black and the wool is all darkish grey,
with no bright colour. The differences are probably seasonal.
A fourth skin, from Ladak (Capt, Strachey), apparently the
only example of the race hitherto recorded from British Indian
territory, differs from the skin from the Yamdok Lake
in its fuller coat, deeper ochreous wool, absence of stripes,
on the loins, and the stripes on the tail not so strong, except
at the end It clearly comes nearer the typical form as described
by Birula, and in its richer-tinted wool connects typical
manul Math the next described race.
35 b. Otocolobus manul ferrugineus Ognev.
Felis manul, Eadde & Walter, Zool. Jahrb. iv, p. 1013, 1889.
Otocolobus manul ferrugineus, Ognev, C. B. Acad. Sci. XJBSS»
p. 308, 1928 ; id., Mamm. of USSR, etc., iii, p. 186, 1935.
Locality of the type, Transcaspia.
320 FBLIDJ5E.
Distribution. — Transcaspia (Askhabad ; Serachs according
to Radde), N. Persia, Afghanistan, and BAUTcmSTAsr.
Distinguished from the more northern races of the species
and from O. manul nigripectus by the rusty-reddish hue of the
pelage.
This "' red " form of 0. manul has been known to me for
many years from trade skins for which no definite localities
were known. I regarded them as " erythristic mutants"
of the normal form ; but since the phase appears to have
a definite distribution, it may be regarded as a subspecies,
as Ognev claimed. There are two skins of it in the British
Museum. One from Meshed in N. Persia (Major P. M. Sykes)
has the wool bright red, only sparsely silvered by the contour
hairs, and the pattern is everywhere obsolete, being red where
it is faintly detectable. The other, from the Toba Plateau,
on the southern Afghan-Baluchi border (Col. Wilson), has the
wool not so red as the last, but much more rusty-ochreous
than in nigripectus and less concealed by the silvery contours.
It also has distinct blackish spots on the head and stripes
on the cheeks, but those on the loins and tail are very faint.
This skin, is intermediate between the skin from Meshed and
the skin of nigripectus from Ladakh. The evidence that
ferrugineus comes into the British Indian fauna is supplied
by a very red skin preserved in the McMahon Museum at
Quetta and secured by Sir Henry McMahon, as he told me in
litt., in "the mountainous wooded tract between Ziarat and
Hindubagh in the Zhob district of Baluchistan, 100 miles east
of Quetta."
The following dimensions (in English inches) of the two
races of T. manul here recorded are taken from skins, no
flesh-measurements being available : —
Head and
Name, locality, and sex. body. Tail.
Tibet (flat skin, Zool. Soe.) ; ad. <J 20 9
Yam. Lak, Tibet (made-up skin) ; imm. $ 20 9J
Tibet (Hodgson) ; ! ad. $ 18J 8J
ferrugineus.
Toba Plateau (flat skin) ; ? <J 26 114-
:\Teshed ; § 24 12
The measurements suggest that ferrugineus may be a larger
form ; but taken from preserved skins they are quite unreliable.
The weight of typical manul, according to Pallas, is from
6 to n lb.
OTOCOLOBTTS.
321
The following are the measurements (in mm.) of two skulls
of nigripectus : —
Locality and sex.
.2 e>
•2— 3
■3§B
ostorbital
width.
iterorbital
width
axillary
width
=31?
1*
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93
85
71
39 19+
26
63
12
9
85
75
69
41
19
23
56
10
8*
"f
The measurements of the skull from Tibet, which is no
longer available, are taken from my paper published in 1907.
Apart from its smaller size, it differs from the Ladakh skull
in having the postorbital bar mesially incomplete and the
forehead between the eyes not quite so flat, both, no doubt,
a question of age.
Habits. — Col. C. H. Stoekley tells me (in litt.) that he saw
a fine specimen of this cat amongst the sand-hills at the south
end of the Tso Moriri Lake in August 1911. It was basking-
on a little patch of sand just below an overhanging bush,
and he watched it at close range for some minutes.
Nothing else, however, has been recorded of the habits of
the two races described above, but they probably differ in no
important respects from those of the northern race, which
inhabits rocky districts of Central Asia and preys on small
mammals, especially picas or mouse-hares (Ochotona) and birds.
Its pallid colour matches its surroundings, and the high-set
eyes, short, low-set ears, and flat forehead are adaptations for
peering at prey over the edge of a sheltering rock with the
smallest possible area of the head exposed. A specimen kept
in the Zoological Gardens carried its tail low, with the end,
which was continually jerked up and down, upcurled, displaying
the black of its underside. Its " spit " was a short, sharp
" ts, ts, ts," projected through closed lips, and its sexual
call was a combination of the bark of a small dog and the
'" hoot " of an owl.
VOL. 1.
322
FELTD2E.
Subfamily ACINONYCHINiE.
Resembling the Felinse in having a normal hyoidean
apparatus, but differing in the structure of the feet, which
have lost all trace of the cutaneous lobes constituting the
projecting sheaths of the claws*. The claws of the four main
Fig. 77. — A. Lower side of fore paw of Hunting Leopard (Acinonyx
jubaUis), stripped of hair to show the absence of the cutaneous
claw-sheaths, the pointed digital pads, the ridged plantar pad, and
shallow webs. B. The same of the hind paw.
(For comparison with the paws of the Leopard, p. 195.)
* There has been a good deal of misapprehension about the retracta-
bility of the claws of the Cheetah ; but J. Gt. Wood was not far wide
of the truth when, about 70 years ago, he stated, after examining the
paw of a living specimen, that "the claws were as retractile as those
■of a eat."
-a
c
o
c
a
o
3
I
ACINO^YX.
323
■digits of the front and hind paws are comparatively blunt and
only slightly curved, hut on the first digit, the " dew-claw,"
■of the fore paw the claw is large, sharp, and strongly curved.
The digits themselves can be considerably spread owing to
the shallowness of the webs, the pads are hard, the digital
pads being compressed at the tips, the plantar pads have
a pair of low ridges, and the carpal pad is conical and pointed
at the end. Other characters of the single representative
of this subfamily, Acinonyx jubatus, are mentioned under the
generic and specific descriptions.
Genus ACINONYX Brookes.
Acinonyx, Brookes, Cat. Mus. Joshua Brookes, p. 16, 1928 * ; Palmer,
Index Gen. Mamm. p. 74, 1904 ; Hollister, Proe. Biol. Soe. Wash.
xxiv, p. 226, 1911 .
Gynailurus, Wagler, Syst. Amphib. p. 30, 1S30 (Oynxlurus of many
subsequent authors).
Guepardus, Duvernoy, L'Inst. Paris, ii, p. 145, 1834.
Cynofelis, Lesson, Nouv. Tabl. Regne Anim. p. 49, 1842.
Type-species of Acinonyx, venator (=venatica) ; of Gynailurus,
jubatus ; of Gtiepardus, guttatus ; of Cynofelis, jubatu*
Fig. 78. — Side view of adult ? skull of Hunting Leopard (Acinonyx
jubatus) from Nigeria, showing especially the domed crown and
small post-canine space, x §.
Distribution. — As under the species.
The best generic characters are supplied by the skull, which
differs from that of other Felidse in a combination of features,
of which a few may be mentioned. It is short and dome-
* I am compelled to take this reference on trust, there being no
available copy of this Catalogue of Brookes's collection to consult.
Y2
324 FJELID^E.
shaped, the facial and cranial portions sloping tolerably steeply
and evenly from its highest point a little in front of the very
short postorbital processes, which are situated near the middle
of its length ; and, in conformity with the downward slope of
the crown, the basieranial axis is inclined upwards, not in the
same plane as the basifacial axis. There is no trace of sagittal
crest, and the suborbital portion of the zygomatic arch is
hardly at all salient, and is in about the same plane as the outer
surface of the maxilla, which has a very small, low-set, often
divided infraorbital foramen. The nares, especially the
posterior, are large. The mandible is weak, with a straightish
inferior edge and a nearly vertical chin ; its anterior dental
area is not raised, the sockets of the front teeth being in line
with those of the cheek-teeth behind. On account of this
feature, combined with the close proximity of the upper and
lower premolars to the small canines and the absence of the
cusp on the small inner lobe of the upper carnassial permitting
the anterior blade of the lower carnassial almost to reach
the palate, there is no trace of post-canine space when the
jaws are closed. The skull, as a whole, shows very little
evidence of muscular development, and its bone is thin and
light. In its general shape and the closure of the jaws it is
the very opposite of the skull of the Clouded Leopard (see
p. 247).
36. Aeinonyx jubatus (Schreber). The Hunting Leopard or
Cheetah.
Felisjubata, Schreber, Saug. iii, pi. 105, 1776, text p. 392, 1777.
Gynailurus or Cynselurus jubatus of most authors, including
Blanford, from 1830 to 19U.
Aeinonyx jubatus, Hollister, Proe. Biol. Soc. Wash, xxiv, p. 226,
1911.
Locality of the type, Cape of Good Hope.
Distribution. — The northern part of Peninsular India,
South-west Asia, and the greater part of Africa, except the
central forested area.
Bodily size and length of tail very much as in the Panther
(P. pardus), but the body not so robust, the head, with rounded
ears, relatively considerably smaller and shorter, and the
legs much longer and thinner, with small compact paws,
adapted for swift running. The coat is short, with or without
underwool, according to the season, and in the winter at least,
usually at all seasons, is developed into a mat of longish hair
up to 3 in. or more in length, the so-called " mane," which
extends down the nape and over the summit of the shoulders.
The mystacial vibrissse are short, the others poorly developed,
if present, but there is a row of rigid lashes on the upper eye-
ACINONYX. 325
lid. The general ground-colour of the upper side varies from
tawny to pale buff or greyish- white, but the distal end of the
tail is white above ; there is some white about the eyes and on
the muzzle below the rhinarium.and the whole of the underside
is white from the ehin to the tail- tip. A constant feature
of the pattern is a conspicuous black stripe extending from
the eye to the mouth on each side. For the rest the pattern
consists normally* of tolerably closely-set solid spots, not
" rosettes," showing lineal arrangement at most only on
the head and nape, where they are smaller than on the body ;
small spots extend to the paws and large spots along the
proximal two-thirds of the tail, but on the terminal third
they form transverse black stripes, the tip being normally
white.
The " mane " on the nape and shoulders of the adult,
above described, is the remnant of an extensive mat-bike
growth of long hair which in cubs covers the whole of the
upper side from the crown to the root of the tail. This
gradually disappears as growth advances. In the cubs of all
the other species of 3?elidse the hair is of tolerably uniform
length all over the body.
The first two specific names unquestionably given to
Hunting Leopards were fortunately applied to specimens
from districts at the extremes of the geographical range of
the genus, namely, South Africa and India. These appear to
be at most racially distinguishable. The South African form,
A. jubata jubata, has a thicker, more woolly coat, at least
in winter, a feature which suggested the name laneus given
to a red-spotted variety of it from Beaufort West by P. L.
Sclater. Other names have been used for this animal, some
of doubtful application, but they are for the most part of later
date than the name here taken for the Indian race, and it is
probable that the Indian form extends into North Africa.
There is no evidence, for example, that the Cheetah of Somali-
land is distinct from it.
36 a. Aeinonyx jubatus venatieus (Griffith).
Felis venatica, Griffith, Vert. Anim., Carnivora, p. 93, 1821 ;
Gray (nee H. Smith), Griffith's Anim. Kingd. v, p. 166, 1827.
Aeinonyx venator, Brookes, Cat. Mus. Joshua Brookes, p. 16,
1828.
* Except in the Rhodesian form that I named Aeinonyx rex, in 'which
the pattern consists of longitudinal bands down the back and large
stripe-like blotches, often fusing, on the flanks. This is no doubt
only a " mutant " of the ordinary African Cheetah, not a distinct
species. In some abnormal Leopards a simUai variation of pattern
occurs.
326 FELIDiE.
Oynselurus or Cynailurus jubatus, Blyth, Blanford, and other
writers on the fauna of India *.
Locality of the type of veTiatica, and venator also according
to Hollister, India.
Distribution. — Formerly Nobtheen India south of the
Ganges from Bengal to Rajputana, the Punjab, and Shad;
also Cbntbal India and the northern part of the Deccan,
but now almost, if not quite, extinct in Hindustan. Probably
the Persian and Mesopotamian Cheetah, also almost extinct,
belongs to this race.
Distinguished from typical jiihatus from Cape Colony by
its thinner, less woolly winter coat and by the absence of
the mane, according to Griffiths, probably in the summer
coat.
Griffith described this Cheetah as " the maneless Hunting
Leopard," and published a coloured plate of the example
he saw, showing a perfectly smooth nape and a short, sleek,
no doubt summer coat. The only Indian example I have seen f
has a fairly long, soft coat, with a little underwool, but there
is a decided mane, running down the nape and over the shoulder.
It is probably in winter coat, the hair on the back and sides
being about 25 mm., on the belly 90 mm., on the mane of the
fore nape 55 mm., and over the shoulder 80 mm. The colour
is buffish-tawny or sandy-fawn down the back, becoming
gradually paler on the sides, the tint of which blends with the
white of the belly. The entire underside from the chin to
the tip of the tail and the inner sides of the legs are white, and
there is some white on the front of the muzzle, below the eye,
and above it in front, involving and emphasizing the upper
end of the black stripe curving down from the eye to the mouth.
The small spots on the head and nape are lineally arranged,
but elsewhere on the body and legs they are irregularly
scattered and black, extending to the paws and over the
greater part of the tail, but the terminal 9 or 10 inches of the
tail have transverse black stripes ; the ear is tawny at the
summit externally, black at the base, the black extending
higher behind than in front. The pattern differs in no respect
from that of typical African Cheetahs. No flesh-measurements
* Blanford's omission of venaticus from the synonymy of the Indian
Hunting Leopard is curious. Hollister, who revived the name for
the Indian form, to -which he gave fall specific rank, presumably solely
because of its distribution, was unacquainted with the first description
of 1821 and assigned it to Hamilton-Smith : but Gray wrote the accounts
of all the Mammals, except the Ruminants, in vol. v of Griffith's 'Animal
Kingdom ' (Saunders, ' Gray's List of Memoirs,' etc., p. 6, 187S).
t This skin, from Jubbulpore, C.P., was kindly sent to me for the
British Museum by Mr. J. A. Clough in response to my appeal for an
Indian Hunting Leopard, which was not at the time represented in
the National Collection.
ACINONYX.
327
are available, but the dressed skin, allowing for the loss of
about half an inch at the tip of the tail, is : — Head and body
50 in. ; tail 27 in. ; hind foot 9J- in.
The skull of the Indian Hunting Leopard is unknown, but
the measurements (in mm.) of two African skulls are as
follows : —
Locality and sex.
EH
Zygomatic
Postorbital
width.
Interorbital
width.
s
is
8
5
1 Brit. Somaliland ; just. !
ad. $ 150
i Zoutpansberg, S. Afr. ; '
ad. o i 193
i
139
172
i
107 50 34 ! 43
131 54 42 i 53
105! 22
i
133 24
17
19
These are the largest and smallest $ skulls available for
measurement. They probably represent approximately the
extremes in size for that sex. The one from S. Africa is
typical jubatus. Hence, if the suggestion that the Indian
Hunting Leopard is the same as the one inhabiting Somaliland
is correct, the difference in dimensions between the two skulls
probably indicates that venaticus is a smaller race than
jubatus.
The other Asiatic race, the Transcaspian Hunting Leopard,
A. jubatus raddei Hilzheimer (SB. Ges. Nat. Er. Berlin, no. 5,
p. 291, 1913), apparently differs from typical venaticus by its
fuller winter coat. Hilzheimer divided the Hunting Leopards
of Africa and Asia into several " species," A. raddei being
one of them. Ognev (Mamm. USSR, iii, p. 309, 1935) cited
raddei as a race of venaticus, accepting, apparently, Hollister's
specific separation of the Asiatic from the African forms.
In my opinion the differences between all Hunting Leopards
are too trivial to be given more than subspecific status.
Habits. — The names Gynailurus and Oynofelis (Dog-like
Cat) reflect the opinion, still apparently held by some sports-
men, that the Hunting Leopard serves to connect in a
measure the Cat and Dog families. The resemblance to
the Dog, however, is purely functional and superficial.
The Hunting Leopard is in reality a highly specialized cat.
Its structural peculiarities have been secondarily acquired
in adaptation to its method of hunting. Instead of leaping
on its prey from a point of vantage, at comparatively
close quarters, like the majority of cats, this species, as
a general rule at all events, runs down antelopes in the
open by sheer speed of foot, after creeping as near its quarry
328 BBLma:.
as possible under cover of low vegetation All its more
obvious external characteristics are clearly modifications for
insuring success in the enterprise. Its sandy hue, broken
up by the black pattern, must favour concealment during the
stealthy stalk and during the initial rush when cover is quitted.
Similarly the small, light head, narrow body, powerful hind-
quarters, long, sinewy legs, and paws with spreading digits,
free claws, and hard pads, are all clearly fitted for traversing
hard, open ground at topmost speed ; and the enlarged nasal
passages are a provision for a plentiful supply of oxygen
to the lungs in the event of the continuance of the chase
over a distance of a quarter of a mile or more. Also the high
dome-shaped skull, with uplifted orbits, enables the Cheetah
to peer over the top of low bushes or other cover with the
least possible exposure of the head when the quarry is being
stalked.
The Cheetah, like the Lion, Hysena, and some other species
common to Africa and India, entered India by way of Persia
and Baluchistan. It formerly had a wide distribution in
Western and Central India south of the Ganges, and extended
through the Deccan at least as far south as Coimbatore
(R. C. Morris), its range agreeing tolerably closely with
that of the Blackbuck. But it is now to all intents and
purposes a thing of the past so far as the fauna of India is
concerned. Being mainly diurnal, and frequenting compara-
tively open country, and defenceless as well, it was easily
found and killed by native and English shikaris, the latter
enjoying the sport of killing it on horseback with a spear,
the greater endurance of the horse enabling it to overtake
the quarry in the end.
The habitat was the open plains or the low hills bordering
them, any kind of cover or rock-shelter being used to lie up in.
Cheetahs climb when young, but very seldom do so when adult,
unless treed by dogs.
At an early date Indian potentates took advantage of the
Cheetah's quiet disposition, docility, and peculiar method of
hunting to tame and employ it 'for coursing Blackbuck*.
The Cheetahs, it is alleged, had. to be caught full grown, after
learning how to hunt from accompanying their parents.
If taken as cubs and reared in captivity, training them was
a tedious unsatisfactory task.
They were commonly caught in nooses set about trees
which it was known they were in the habit of visiting to
" sharpen " their claws. Taming them was not a long process.
When taken out for the chase the Cheetah was hooded, after
the manner of a hawk, and tethered on the top of a lowbullock-
* Since the Indian Cheetah became rare, specimens have been
regularly imported from Africa to replace it foi this aport.
acinonyx. 329
cart. The cart was then driven as near as possible to the
antelope, which paid little heed to it. The Cheetah's hood
was then taken off and the animal, on sighting the antelope,
slipped from the cart and either went headlong at the herd,
if near enough, or stalked it, making use of any cover nearby,
until within rushing distance, perhaps of 100 yards or so.
Traversing the ground with such speed that the species has
been described as the fleetest of mammals, the Cheetah often
overtakes the rearmost of the herd, which gets under way
the moment it perceives the Cheetah approaching- The
victim, perhaps one of the slower bucks, is usually apparently
struck over by a blow of the Cheetah's fore paw, is then seized
by the throat, to which the Cheetah holds on until enticed
to let go by the offer of a ladle of blood taken by one of the
men from the antelope's arteries. The hood is then slipped
on again and the beast put back on the cart. If, however,
the herd has sufficient start, the Cheetah, which is short-
winded in comparison, and can only keep going at full speed
for at most about 600 yards, may fail to get up to it and
abandons the chase. So far as I am aware Dunbar Brander
is the only authority to maintain that the speed of the Black-
buck, once fully under way, is faster even that the initial
speed of the Cheetah ; and he makes the suggestion that the
" incredible " speed of the Blackbuck and of the Chinkara,
surpassing he says, but I know not on what authority, that of
any African antelope, has been acquired as a means of escape
from the Indian Cheetah.
His view that the speed of these antelopes exceeds that of
the Cheetah is probably correct ; but it must be remembered
that such African antelopes as the Springbuck, Grant's
Gazelle, the Impala, and others have been chased for genera-
tions by Cheetahs, and have the same need of speed to escape
them. They are probably as fleet as the Indian species.
As regards the actual speed of the Cheetah, Mr. Gandar Dower,
who recently imported trained specimens from E. Africa
and tested them against a motor-car, found that they travelled
at the rate of 45 miles per hour when apparently going at full
speed, that is to say, they can cover a quarter of a mile, nearly
the full distance they can keep it up, in 20 seconds, and 100 yards
in about 4-| seconds, faster than the best greyhound and twice
the speed of the fastest human " sprinter."
It seems clear that this exceptional speed for a carnivorous
mammal was acquired by the Cheetah, both in Africa and
elsewhere, for the capture of fleet-footed antelopes, represented
in India by the Blackbuck and Chinkara mentioned above.
But they preyed upon other game as well. They have been
known to attack Nylghaie and to take domesticated goats
and sheep ; and no doubt they fed upon smaller mammals,
like hares, and upon birds as well. It may be assumed,
330 VIVBEEID^E.
moreover, that they frequently hunted in couples, as has
been observed in the case of the African races.
As above stated, Cheetahs are timid animals, never known,
apparently, to attack man unprovoked, and only rarely to
charge when wounded.
The sounds they make are typically " feline." They
growl, snarl, spit, mew, and purr like a domestic cat, sure
evidence of close kinship, but in addition they have a whistling
note like a bird's.
The period of gestation in an African Cheetah was stated
to be about thirteen weeks (90 days), and the young, two or
four to the litter, are born, with the eyes closed, in any con-
venient shelter.
Family VIVERRID^.
Distinguished in external characters from the Felidse by
the hind foot being five -toed owing to the invariable presence
of the first digit, by the retention of the interramal tuft
of facial vibrissae, and typically by the longer muzzle
and shorter limbs. The skull differs by the position of
the post-palatine foramina on the maxilla, almost always
well in advance of the maxillo-palatine suture *, and usually
about the level of the second premolar ; by the distinct external
division of the auditory bulla into its two elements either
by a definite groove or, when rarely this is obliterated, by the
depression of the tympanic bone in front of the swollen
entotympanic. The dental formula is, typically, i. |, c. = >
pm. | , to. | = 40, but the number may be reduced, although
never to the same extent as in the Felidse ; and in all the
British Indian representatives of the family the lower carnassial
(mi) differs from that of the Felidse by retaining the inner
cusp (metaconid) of the anterior part of the crown and by the
presence of a definite " heel."
This family, which is clearly less specialized than the
Felidse, contains a great number of highly diversified genera,
and is susceptible of division into several subfamilies,
based mainly on the structure of the feet and of some highly
specialized scent-glands, derived from the skin, which are
present in most of the species and are situated in the region
of the external generative organs. The feet vary in accordance
with the terrestrial or arboreal habits of the animals. In
those that live mainly on the ground, like the Civets, the feet
* In one skull of the Binturong (Arctiatis), from Borneo, aberrant
in this respect, the foramina are only about 1 mm. from the suture
as far back as the upper carnassial (pm l ).
V1VEEBID.E. 331
may resemble very closely those of the Felidee, even to the
possession of lobes of skin to protect the points of the retractile
claws ; but in the arboreal species like the Tree-Civets and
Binturong the feet are very different, especially in the large
size of the carpal and metatarsal pads, although the claws
are usually sharp and curved and retractile to a certain
extent. The scent-glands consist of an hypertrophied mass
of sebaceous glands which, when the organ is highly
developed, pour their secretion into an extensive pouch formed
of two thick flaps of skin, the edges of which are capable of
being tightly pressed together, closing the pouch and storing
the secretion. These glands are usually found tolerably
equally well developed in both sexes ; and from observation
on captive animals which have been seen to rub the secretion
on the walls, bars or other parts of their cages, it seems that
the perfume is used to scent a locality and enable the animals
to find each other.
The Viverridee are the most primitive of all the families-
of ^Eluroid Carnivora. They are found all over the Oriental
Region and even beyond it across " Wallace's line,'' all over
Africa, whence they pass into southern Europe, and they aTe
the only indigenous Carnivora inhabiting Madagascar, unless
the Fossa. (Cryptopr octet*) be granted family rank. Their
occurrence in Madagascar and in Celebes, as well as some of
the adjoining islands, shows them to be ancient inhabitants
of the tropics of the Old World.
In his volume on British Indian Mammals, Blanford,
following the custom of his day, included in the Viverridae
the Mongooses as a special subfamily, Herpestinse. They
are here regarded as a distinct family, the Viverridse of this
volume being equivalent to Blanford's Viverrinse.
Key to the British Indian Subfamilies, based mainly on
External Characters.
a. No scent-glands, perinaeum short, prepuce close
to scrotum and vulva to anus ; feet terrestrial,
digitigrade ; m 1 small, to 2 absent, m t with small [p. 332 .
heel Prionodontinae,
a'. Seent-glands present, at least in $, perimeum
longer, prepuce always far in advance of scrotum
in ,J ; in 1 large, m 2 present, iiij with large heel.
6. Seent-glands present in both sexes, in <J
perinseal, between scrotum and prepuce, in
$ behind the vulva or encircling it.
c. Feet terrestrial, digitigrade, carpal pad
remote from plantar pad, single and conical ;
metatarsal pads absent, hmd foot hairy
from heel downwards ; scent-glands opening [p. 342.
into highly specialized pouches Viverrinae,
* This genus has by some zoologists been classified with the Felidse.
It has nothing whatever to do with them.
332 vivebeid^;
a'. Feet soansorial, semiplantigrade, carpal and
metatarsal pads double, touching, and as
•wide as the plantar pads inferiorly ; glandu-
lar pouches less specialized.
(I. Scent-glands large, in $ surrounding vulva,
in $ extending to prepuce, penis not pen-
dulous ; carpal pads not narrowed above,
the whole or greater part of the hind [p. 376.
foot naked from the heel Paradoxurinae,
d'. Scent-glands comparatively small, not
extending to vulva in ? or to prepuce
in $ ; distal end of penis pendulous ;
carpal and metatarsal pads narrowed
above, greater part of hind foot hairy [p. 450.
from heel Hemigalinae,
b'. Scent-gland absent in 6", present in $!, but
mainly in front of vulva; feet as in Para- [p. 439.
doxurina? Aretogalidiinae,
Subfamily PRIONODONTDWE.
Prionodonthix* Gray, Proc. Zool. Soc. 1864, p. 519 (in part) ;
Pocoek, Proc. Zool. Soc. 1933, p. 970 (sensu striata).
Linsanglnss, Pocoek, Ann. Mag. Nat. Hist. (8) xvi, p. 350, 1915
(in part) *.
Distinguished from the other British Indian subfamilies
of Viverridae by the absence in both sexes of the perinseal
scent-glands, coupled with the shortness of the perineeum
itself, the vulva being close to the anus and the short penis
to the scrotum as in the Felidse. The teeth also are more
highly specialized, and show an approach to those of the
Felidse, although more primitive. The dental formula is : —
i. | , c. * , pm. * , m. I = 38. The incisors form a transverse, not
a curved, line ; the first three upper and the four lower pre-
molars are compressed and trenchant, with a high, sharp,
median cusp and small subsidiary cusps in front and behind it ;
the upper carnassial {pm 4 ) has a small inner lobe (protoeone)
set far forwards, a small cusp (parastyle) in front of the main
compressed, high, pointed cusp (paracone), and a compressed,
blade-like posterior cusp (metacone) ; the upper molar (m 1 )
is much smaller than the upper carnassial, triangular, trans-
versely set, and much wider than long, so that the upper
carnassial is nearly at the posterior end of the upper cheek-
teeth as in the Felidae. The second molar, present in the other
* Following Gray, I originally affiliated the West African genus
Poiana with JPrionodon, but evidence since found on made-up skins
of the existence of scent-glands in Poiana induces me provisionally
to regard the latter as a specialized form of Genetta, its likeness to
Prwnodon being possibly adaptive. In the structure of the feet, as
I showed in 1915, Poiana is more like Genetta, and the paroccipital
bone does not form a thickened prominence as in Prionodon.
PBIONODONTINiE.
333
subfamilies, is undeveloped. In the lower jaw the fore part
of the oarnassial has two compressed blade-like cusps as in
the Felidse, but there is a small cuspidate metaconid on the
inner side, and the heel, although small, is as wide as the fore
part of the crown and about one-fourth of its length.
Iff. !, i f\
D
A
Fig. 79.
Left fore-paw of Prionodon linsang,
showing the claw-sheaths,
the four-lobed plantar pad, and the bilobed carpal pad (c).
B. Left hind paw of the same, showing reduced claw-sheaths and the
three-lobed plantar pad, with its inner or hallueal lobe small and
remote.
C. Rhinarium of the same from the front.
D. Anal and genital area of female of the same, showing the vulva (6)
close to the anus (a), with no gland on the narrow perinseal region
between them.
E. The same of male, showing the area between the penis (p) and the
anus (a) occupied by the scrotum.
In the very small size of the first upper molar, the loss of
the second, and the greatly reduced heel of the compressed
lower carnassial this subfamily differs from the rest found in
British India.
334
VIVERKIDiE.
Genus PRIONODON Horsfield.
Linsangs.
Prionodon, Horsfield, Zool. Res. Java, no. 5 (under Mangusta
jamnica), 1822 (Sherborn) ; and of several subsequent authors,
including Blanford ; Pocook, Proc. Zool. Soe. 1933, p. 970*.
Prionodantes, Lesson, Now. Tabl. R. Anim. p. 60, 1842.
Linsang, S. Muller, Verh. Nat. Gesch. Nederl. i, p. 28, 1839, and of
some subsequent authors ; Poeock, Ann. Mag. Nat. Hist. (8)
xvi, p. 341, xvi, 1915.
Linsanga, Lydekker, Geogr. Hist. Mamm. p. 20, 1896 ; id., Cats,
Civets, etc., in Lloyd's Nat. Hist. p. 221, 1896.
Pardictis, Thomas, Proc. Zool. Soe. 1925, p. 499.
Type of Prionodon and Prionodantes, gracilis Horsf. (both
as subgenus of Felis) ; of Linsang and Linsanga, gracilis
Horsf. ; of Pardictis, pardicolor Hodgs.
^7 H
Kg. 80.
A. Left half of palate and teeth of Prionodon linsang from Tenasserim
pm 1 and pm* first and fourth premolars ; to 1 small and transversely
set first molar ; ppf, posterior palatine foramen.
B. Right half of dental part of mandible of the same, with fourth
premolar {pm t ) and first molar m v the latter with tricuspid anterior
portion and small " heel."
Right upper carnassial (pm 4 ) from outer side.
Right lower carnassial (m,j) from outer side.
Left auditory bulla of the same, showing small anterior or tympanic
The same of a specimen of P. pardicolor from Nepal, with larger
anterior and smaller posterior or entotympanic chamber.
C.
B.
E.
F.
* Owing to its assumed preoccupation by Priodontes, given to the
•Giant Armadillo, Prumodon was for many years rejected, and Linsana
emended to Linsanga by Lydekker, used in its place.
PRIONODON. 335
Distribution. — Eastern Himalayas (Nepal, Sikkini), Upper
Burma, Tenasserim, Laos, Tong-king, Siam, Malay Peninsula,
Sumatra, Java, and Borneo.
Size small ; tail long, nearly as long as the head and body,
about five or six times as long as the hind foot. Pattern
distinct, consisting of large spots, sometimes coalescing into
broad bands on the sides of the body and of transverse bands
on the tail. Head elongated with narrow muzzle, rhinarram
evenly convex above, with wide internarial septum, shallow
infranarial portion, and philtrum narrow and grooved, the
groove extending only about to the level of the lower edge of
the nostrils. The ridges of the basal portion of the ear are
complicated. The antero -internal ridge has a valvular flap
overhanging the anterior portion of the supratragus, on the
postero-external ridge there is a deep groove surmounted
by a crest running back below the bursa, and the bursa is
well developed, its posterior semicircular flap rising behind
the edge of the pinna, and its anterior flap is deeply emarginate.
The paws have the claws completely retractile. In the
fore paw there is a pair of claw-sheaths on each of the four
main digits ; the plantar pad consists of four lobes denned
by deep grooves and arranged in a strongly curved line, the
external carpal pad is large and oval, and the internal is
a small lobe attached to it ; the area round the pads is covered
with hair. In the hind paw there is at most a small protective
lobe of skin on the outer side of the claws of the four main
digits, the plantar pad consists of three lobes separated by
deep clefts, the internal or hallucal lobe being small and
set above the inner of the three main lobes. As in the front
foot, the area round the pads is hairy and the rest of the under-
side of the foot is hairy from the heel to the plantar pad,
there being no trace of metatarsal pads *. The paws, although
resembling those of the typical Pelidse in the retractile claws
and general hairiness of the underside, differ in the shape
and sharp definition of the lobes of the plantar pads, in the
presence on the fore paw of the internal lobe of this pad and
of a small inner carpal pad, and on the hind foot of the first
digit and of the reduced internal lobe of the plantar pad.
There are two pairs of mammse.
The delicate skull is long, low, and narrow, with the muzzle
narrow and elongated, its general shape being very different
from that of the Felidse. The occipital area is well defined
and has a strong crest, but there is no complete sagittal crest,
* The external characters of this genus were figured and described
by Hodgson (Oale. J. Nat. ii, pi. i, 1842, and viii, pi. i, 1847), by Mivart
(Proc. Zool. Soe. 1882, p. 158), and by myself in 1915 in the paper cited
in the synonymy.
336
VIVBKEIDiE.
the temporal ridges not meeting, and usually defining a wider
or narrower lyrate area ; the postorbital area is constricted,
about the same width as the interorbital area or a little narrower,
and a little wider than the maxilla above the canines ; the
postorbital processes are prominent, but bluntly angular, and
the mesopterygoid fossa is roofed in its anterior half by the
posterior extension of the palate.
Fig. 81. — Side view of skull of Prionodon linswng (skull of
type of P. maculosus Bl.).
The two well-marked species of this genus may be dis-
tinguished as follows : —
a. Pattern on sides of body consisting of large
spots not coalescing to form broad transverse
bands. Skull with the tympanic bone of bulla
relatively large and swollen, the bullate portion
(entotympanic) relatively small ; external [p. 337.
pterygoid crest less well developed pardicolor Hodgs.,
a'. Pattern on sides of body consisting of broad
transverse bands due to the coalescence of
spots ; tympanic bone of bulla relatively
small and but little swollen, the bullate
portion (entotympanic) correspondingly larger ; [p. 339.
external pterygoid crest better developed . linsang Hardw.,
The skull-characters mentioned under the diagnosis of
'pardicolor, associated with a shorter and more parallel-sided
mesopterygoid fossa, were regarded by Thomas as of generic
importance and formed the basis of his genus Pardictis, quoted
in the synonymy above. The proportions of the two component
bones of the auditory bulla are tolerably constant, but the
mesopterygoid fossa and the external pterygoid crest are
respectively inconstant in shape and development ; and all
three features are too variable in the family Viverridse to
be given, in my opinion, generic value.
PRIONODON.
337
37. Prionodon pardicolor Hodgson.
Prionodon pardicolor, Hodgson, Cale. Journ. Nat. Hist, ii, p. 51,
pi. 1, figs. 3-6, 1842, and viii, p. 40, pi. i, 1847 ; and of subsequent
writers on the Indian fauna, including Blanford, Mamm. Brit.
Ind. p. 103, 188S.
Pardictis pardicolor pardicolor, Thomas, Proe. Zool. Soe. 1925,
P. 499.
Prionodon pardicolor pardicolor, Poeock, Proe. Zool. Soe. 1933,
p. 972.
Viverra perdicator, Sehinz, Syn. Mamm. i, p. 366, 1844 (errore for
pardicolor).
Vernacular. — Zik-cMtm (Bhotia) ; SMiyA, and Silu (Lepoha).
Locality of the type, Nepal.
Distribution. — Nepal, Sikkxm, Assam, Upper Burma,
Yunnan, Laos, and Tong-king.
Ground-colour varying from brownish on the dorsal surface
and nape, paler buffy-olivaceous on the flanks, to bright,
.j^S'H v/yj
Fig. S2. — The Spotted Linsang (Prionodon pardicolor), from a drawing
by Hodgson of a specimen from Nepal.
almost orange-buff. The dark pattern consists of two long
nuchal stripes extending from the occiput to the shoulders,
and of a stripe on each side of the neck ; on the body there are
two rows of small spots on the spine coalescing posteriorly
to form a single stripe ; externally and below these are three
or four longitudinal rows of spots extending from the back
to low down on the flanks, shoulders, and thighs ; these
spots decrease in size below, and those of the two uppermost
rows are continuations respectively of the stripes on the neck ;
vol. i. z
338 yivebeid^;.
the fore leg is spotted to the paw, the hind leg to the hock,,
and the tail has eight or nine complete broad dark rings
separated by narrow white rings.
The British Indian examples of this species supplying the
material for the above-given description came from Nepal
(Hodgson), from Dikehu, 2,000 ft., Singhik, 4,600 ft., Chuntang
5,350 ft. (Crump), and Latohung in the Jellap Pass, 13,000 ft.
(P. Morris) in Sikkim, from Dening in the Mishmi Hills,
Upper Assam, 2,250 ft. (Wells), from Myitkyina (Capt. Abbay),
and the Chin Hills, 50 miles west of Kindat, 5,000 ft. (J. M. D.
Mackenzie) in Upper Burma.
The specimens from the Mishmi Hills and Upper Burma
connect the typical form, P. pardicolor pardicolor, from
Nepal and Assam with the race P. p. presina, described by
Thomas in 1925 from Tong-king, the typical locality being
Ngai-Tio, 4,800 ft. (Delacour and Lowe). Other localities
for it are Bao-Ha, Tong-king, 2,500 ft., Backan, Tong-king,
500 ft., and Xien Quang Koo, Laos. Thomas restricted the
name presina to specimens from high altitudes in Tong-king,
and identified the lowland forms from the same country as
typical pardicolor. But the characters on which he based
the race are unreliable, as was detected by Osgood (Field
Mus. Nat. Hist., Zool. xviii. p. 256, 1932), who identified the
Indo-Chinese specimens as Prionodon (Pardictis) pardicolor.
The type of presina was collected in June, when the moult
was imminent, and its hue is naturally paler and more washed
out than in the well-coloured winter skins with which Thomas
compared it. But on the whole it appears to me that presina
may be retained as a racial name for the Indo-Chinese specimens,
which appear on the average to be lighter or brighter in tint
and to have smaller, more oval, less symmetrically and less
lineally arranged spots than in the Himalayan race.
The adult $ specimen from the Mishmi Hills is the only
British Indian example of this race measured in the flesh.
The examples of presina from Tong-king were measured by
Willoughby Lowe.
Head and Hind
Locality and sex. body. Tail. foot.
Bao Ha, Tong-king ; ad. <J 15§ 14f 2£
Ngai Tio, Tong-king (presina type) ;
ad. <J 15J I* 2J-
Backan, Tong-king ; ad. <J 14§ 13£ 2§
Backan, Tong-king ; ad. § 13§ 12£ 2£
Mishmi Hills, Assam ; ad. £ 14f 12 2f
The weight of the specimen from the Mishmi Hills was 1J lb.
Skull-measurements are entered on p. 341.
Habits. — This species is apparently nowhere common.
The collectors employed by the Mammal Survey of British
India secured only a few. Those sent by Crump from
PRIONODON. 339
Sikkim and from the Chin Hills by Mackenzie were
evidently obtained from natives. The only available
specimens from Nepal came from Hodgson, who reported
the species as '" equally at home on trees and on the
ground : it breeds and dwells in the hollows of decayed
trees. It is not gregarious at all, and preys chiefly on small
birds, which it is wont to pounce upon from the cover of the
grass. The times of breeding are said to be February and
August, and the Utter to consist of two young, there being
two Utters each year " A tame specimen he had was " wonder-
fully docile and tractable, very sensitive to cold, and very
fond of being petted." It was fed upon raw meat, but refused
fish, eggs, and fruits. It was perfectly free from all odour
and was never heard to utter a sound.
Crump, who collected in Sikkim, said that the species is
reported to be nocturnal ; but his opinion that the animal
is a vegetable feeder because he failed to trap it with flesh-
bait is certainly erroneous, since its teeth are essentially
adapted for a carnivorous diet.
38. Prionodon linsang Hardwieke.
Viverra ? linsang, Hardwieke, Tr. Linn. Soc. xiii, p. 236, pi. 24,
1821.
Prionodon maculosus, Blanford, Proc. As. Soc. Beng. 1878, p. 71 ;
id., Journ. As. Soc. Beng. xlvii, pt. 2, p. 152, pis. 6 & 7, 1878 ;
Mamm. Brit. Ind. p. 104, 1888.
Linsang linsang, Lyon, Proc. TJ.S. Nat. Mus. xxxiv, p. 657, 1908.
Prionodon linsang, Robinson. & KIoss, Journ. Fed. Mai. St. Mus.
vii, p. 262, 1919 ; Pocoek, Proe. Zool. Soe. 1933, p. 974.
Vernacular. — Musang Blang (Malay).
LocaUty of the type of linsang, Malacca; of maculosus,
Bankachon, S. Tenasserim.
Distribution. — Irom Sumatra and the Malay Peninsula
northwards to Mottlmeik and, according to Mtiller, Siam.
Ground-colour paler and whiter than in P. pardicolor,
throwing the black or deep brown pattern into greater reUef.
The pattern is fundamentally the same in the two species,
but in P. linsang the nuchal stripes are much broader and the
spots of the two upper rows on each side fuse with each other
and with the median spinal spots to form five wide transverse
bands across the back, the first just behind the shoulders,
the last on the root of the tail and generally coalescing with the
basal caudal stripe : the two nuchal stripes blend with the
lower edges of at least the first of these bands. Low down on
the flanks there is a row of elongated spots or short stripes in
line with the spots on the side of the neck ; still lower there
are a few more spots, as well as some on the fore leg to the
paw and on the hind leg to the hock.
z2
340
vtvekkim;.
The co-type of maculosus does not differ in any characters
of importance from specimens from the Malay Peninsula,
whence the type of linsang came. Blanford was misled into
describing it as representing a distinct species by Horsfield's
statement that linsang was a synonym of the smaller Javanese
race, gracilis.
The ground-colour of linsang varies individually. In the
co-type of maculosus, an adult <J, it is whitish, with the dorsal
surface and mid-line of the nape slightly darkened by the
infuscate tips of the hairs ; the crown and muzzle are brownish,
the forehead brownish huffy-grey, the cheeks, throat, and
underside creamy- white, and the pale bands on the tail become
Fig. 83. — The Banded Linsang (Prionodon linsang), drawn from a skin
from Bankachon, a eo-type of Blanford's maoulosus.
gradually buff posteriorly. Two skins from Malacca, including
Hardwicke's type of linsang, and four from Perak (A. S.
Vernay and Robinson and Kloss) are generally similar to it,
but individually variable in the bumness of the dorsal inter-
spaces and of the underside and cheeks and of the pale rings
on the tail.
Blanford assigned to maculosus a specimen collected by
Mr. lamborg east of Moulmein in N. Tenasserim, and Lyon
recorded two adult $ examples from the Siak River and Sungei
Mandan in Sumatra, which from their flesh- and skull-measure-
ments appear to belong to this form, and Robinson and Kloss
secured one at Bencoolen.
PEIONODCWff.
341
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342 VIVEEBIDJE.
The flesh-measurements (in English inches) of some speci-
mens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Bankachon {maculosus co-type) ; ad. $. 16$ 16 24
Siak River, Sumatra (Lyon) ; ad. <J . . . 16J 141 2f
Bencoolen, Sumatra (Robinson, and
Kloss) ; ad. $ 16f 14 2
Taiping, Perak ; ad. 2 15£ 15 2*
Ulu Ijok, Perak ; ad. ? 14* 13f 2|
The weight of the adult $ from Siak River was 1^ lb.
The skull-measurements of an adult <J, the co-type of
maculosus from Bankachon, and of an adult $ from Perak
in the Malay Peninsula are entered on p. 341. The skull of
the first is the largest of the species I have seen. That of the
<J from Bencoolen, Sumatra (Robinson and Kloss), was
75 mm. in total length and 35|- in zygomatic width, and the
two c? Sumatran skulls recorded by Lyon were respectively
a little over and a little under 73£ mm. in total length, the
zygomatic width of the larger being 36 J mm.
The Linsang above described represents the typical race
of the species P. linsang linsang. In Java and Borneo it is
replaced by P. linsang gracilis, described by Horsfield first
as Felis gracilis and a year later as Prionodon gracilis (Zool.
Res. Java, no. 1, 1821, and no. 5, 1822) from Blambangan,
E. Java, and as Viverra hardwickii by Lesson (Mon. Mamm.
p. 172, 1827), also from Java ; Robinson and Kloss in 1919
made it a subspeciss of P. linsang. Its colour and pattern
are the same as in the Malayan and Tenasserim race, from which
it only differs in its average smaller size and less well-developed
skuU.
Subfamily VIVERRDOS.
In this subfamily, containing the typical Civets or Civet-
Cats, which are mainly or wholly terrestrial, the feet, adapted
for movement on the ground, are digitigrade, the cushion -
like indistinctly subdivided plantar pad and the pads of
digits 2 to 4 being alone applied to the ground, the first digit
being small, set well above the plantar pad, and constituting
a practically functionless " dew-claw " ; the outer element
-of the carpal pads is typically (in all British Indian forms)
alone retained, and forms a conical excrescence high above the
plantar-pad and separated from it in the middle line by
a hairy area continuous externally with the hairy skin on the
back of the fore leg above ; the hind foot is similarly con-
tinuously hairy behind from the heel to the plantar pad.
the metatarsal pads being at most represented by small
areas of naked skin ; the claws are short, sharp, and sometimes
MAMMALIA.
PLATE XXVI.
Photo W. S. Berridfe.
Pallas's Cat (Otccolobus manul).
Photo W. S. Berndge.
Large Indian Civet (Viverra z'ibetha).
VIVEREIN^!.
343
protected by claw-sheaths and as retractile as in the Felidse.
The scent-glands are wholly periaaeal and highly specialized ;
they are visible externally as a tolerably capacious pouch,
with hairy, tumid labia by which the orifice can be opened
or closed.
The skull is long, rather narrow, moderately well moulded
muscularly, but varies in details according to the genus.
The dental formula is: i. |, c. ~ , pm. |, m. f. The
teeth are well developed, and both cutting and crushing
in function ; the upper carnassial (pm i ) has the inner lobe
(protocone) set anteriorly and much shorter than the outer
part of the crown, which has the main cusp (paracone) high
Fig. 84.
A. Anal and genital area of Large Indian Civet (Viverra zibetha) g,
showing the anus (a) surrounded by a raised rim of integument,'
the divided scrotum, the prepuce (p), and the orifice of the glandular
sack (gl) partially open.
B. The same of the Little Civet (Vivemcula indica) $, showing the
anus (a), the vulva (v), and the sack of the gland fully distended.
Vertical transverse section of the gland of 6* Viverra zibetha, showing
the hair -lined cavity of the gland (gl), the thick secreting area of
cells, and the penis (p) below.
and pointed, and the hind cusp (metacone) long and blade-
like ; the outer edges of the two upper molars are turned
obliquely inwards, forming an angle of about 100° with that
of the upper carnassial ; the lower carnassial (m x ) has the
neel large, not much shorter than the fore part of the tooth,
which has its three cusps normally developed.
344 VIVERJRIDJE.
This subfamily is found throughout the Oriental Region,
where it contains the three genera described below. In
Africa it is represented by the African Civet (Givettictis civetta),
which, although formerly assigned to Viverra, differs from that
genus and from other Oriental forms in some structural details
in the feet and skull. The African Genets (Genetta) are also
most nearly related to the Viverrinae, but should perhaps
form a separate subfamily.
The three genera occurring in British India may be dis-
tinguished as follows : —
Key to the External Characters.
a. Anterior edges of the ears widely separated
by the broad forehead ; a dorsal crest of
hairs extending posteriorly at least from
the shoulders.
b. At least the 3rd and 4th digits of the fore
foot with well-developed skin-sheaths
protecting the claws ; feet thickly hairy [p. 344.
between the pads Viverra Linn.,
6'. Digits without skin-lobes protecting the
claws ; feet nearly naked between the [Pocock, p. 354.
pads Moschothesa
a'. Anterior edges of the ears set close together,
the forehead at this point narrow ; no dorsal
crest of long hairs ; claws, as in Mosohothera, [Hodgs., p. 362.
unprotected by sheaths of skin Vivekriouxa
Key to the principal Cranial Characters,
a. Suture between the malar bone and maxilla
comparatively short, much shorter than the
median length of the nasals and than half
the length of the cheek-teeth ; muzzle long
and stout ; the combined auditory bones
short, shorter than the width across the
occipital condyles.
6. Postorbital processes developed and situated
in front of the middle point of the total [p. 344.
length of the skull Vtvekra Linn.,
b'. Postorbital processes suppressed or nearly
so, their position behind the middle point [Pocock, p. 354.
of the total length of the skull Moschotheba
a'. The cheek-suture, above described, very long,
about as long as the median length of the
nasals and more than half the length of the
cheek-teeth ; muzzle short and slender ;
auditory bones large, their length exceeding [Hodgs., p. 362.
the width across the condyles Vivekeictdxa
Genus VIVERRA Linnseus.
Viverra, Linnseus, Syst. Nat. ed. 10, p. 44, 1758 ; and of subsequent
authors (part) ; Pocock. Journ. Bomb. Nat. Hist. Soc. xxxvi,
pp. 423-4, 1933 (sensu atrioto).
Type of the genus, V. zibetha Linn.
VIVEKEA.
345
Distribution. — Noetheen India (Nepal, Bhutan, Assam,
etc.), Bxtema, S. China, Indo-China, Siam, and from the
Malay Peninsula to the Philippines and Moluccas*.
Distinguished externally from the other genera of the
Viverrinse by the structure of the fore feet, of which the
Fig. 85.
A. Lower side of left fore paw of Large Indian Civet ( Viverra zibetha),
showing the claw-sheaths on the third and fourth digits.
B. The same of the left hind paw, which has no protecting elaw-sheaths.
C & D. Same aspect of the fore and hind paws of the Little Civet
(Viverricula indica). In some specimens there is more hair on the
* The smaller Oriental species (Viverra tangahmga), which ranges
eastwards from the Malay Peninsula and does not come into the fauna
of British India, has probably been introduced by man into the Moluccas
and elsewhere for the sake of its perfume.
346 VIVEEEIDJ3.
3rd and 4th digits are provided with lobes of skin which,
like those of the typical Felidse, act as protective sheaths
for the retractile claws. On the 3rd digit there is a pair of
these lobes, on the 4th a single lobe on the outer side. These
protect the claws of the four principal toes, when unspread,
the claw of the 2nd abutting against the inner lobe of the
3rd, the claw of the 5th against the outer lobe of the 4th, and
the claw of the 4th against the outer lobe of the 3rd. The
soles of the feet are hairy all round the pads, except for narrow
streaks of naked skin along the middle line of the digits and
•a similar streak running from the plantar to the carpal pad.
The carpal pad consists of a single external lobe, and the
metatarsal pads are suppressed.
The skull is long and. narrow, with narrow, nearly parallel-
sided, not strongly constricted waist, and the postorbital
processes developed, but small and a little in front of the
middle point between the tip of the premaxillsa in front and
of the occipital crest behind ; the sagittal crest is moderately
strong in the adult ; the infraorbital foramen is approxi-
mately above the junction point of pm z andjpm 4 , and the sub-
orbital portion of the cheek is comparatively short, the suture
between the anterior or malar bone of the zygomatic arch
and the maxilla being much shorter than the median length
•of the nasals, than half the length of the cheek-teeth, and than
the width across the occipital condyles, this width exceeding
the length of the compound auditory bulla.
39. Viverra zibetha Linnaeus. The Large Indian Civet.
(Bibliographical references are contained under the subspeeific
headings.)
Distribution. — Nobthekn India and Southern China over
South-East Asia to the Malay Peninsula.
The general colour is grey or tawny, with the pattern of
the flanks variable in distinctness, but certain features of
the pattern are very constant. There is always a black
spinal stripe running from behind the shoulders to the root
of the tail, but not continued uninterruptedly along the
upper side of that organ, which exhibits a variable number
of complete black and white rings, the black much broader
than the white ; the front of the muzzle on each side has
a whitish patch emphasized by blackish behind, and the chin
and fore throat are blackish ; the sides and lower surface
of the neck are conspicuously banded with black stripes set
off by white interspaces ; one of these stripes, starting behind
the ear, runs backwards nearly to the shoulder, where it turns
vertically downwards and passes on to the lower surface of
VIVBBEA 347
the hind throat to join its fellow of the opposite side ; a
second, starting below the ear, similarly runs backwards and
downwards to form a much broader black band on the middle
of the throat below ; a third, below the latter in its origin,
passes on to the throat in front of it, between it and the
black area of the fore throat. Hence the sides of the neck
and throat are ornamented with three black and two white
collars, of which the most conspicuous and constant are the
two white collars with the broad black collar between them.
The lower portion of the legs at least is black. There are
usually three, sometimes two, pairs of teats.
This species is considerably larger than V. tangalunga,
the only other species of the genus, and is further distinguished
by the white bands on the tail forming complete rings and the
pattern of the flanks, when distinct, showing a decided
tendency to run into vertical wavy stripes. In V. tangalunga,
which is about one-fourth smaller, the black dorsal stripe
is continued to the end of the tail, and the body-pattern is
composed of solid black spots, usually numerous, small, and
close set, and never showing a tendency to run into wavy
'" mackerel-like " stripes.
In F. zibetha, especially in the northern races, the coat,
colour, and pattern vary considerably in accordance with the
season. In winter the coat is long and thick; in summer
short and sleek. The body-pattern is strongly pronounced
in summer, indistinct or even obliterated in winter ; and
the ground-colour varies individually, even irrespective of
season, from tawny to clear, almost silvery-grey, the contrast
between the pattern and the ground-colour being more
emphatic in grey than in tawny specimens. There appears
to be but one moult, which takes place in May, June or July,
the date varying no doubt with latitude.
Males and females are approximately alike in size and in
cranial and dental characters.
The differences above alluded to in colour and pattern,
now known to be individual and to a great extent seasonal,
account for the number of names applied to most of the local
races of this Civet.
39 a. Viverra zibetha zibetha Linnseus.
Viverra zibetha, Linnaeus. Syst. Nat. ed. 10, p. 44, 1758.
Viverra undulata, Gray, Spic. Zool. p. 9. pi. 8, 1830.
Viverra orientalis or melanurus and V. civettoides, Hodgson,
Calc. Jom-n. Nat. Hist, ii, pp. 47-50 and p. 62, 1842.
Viverra zibetha zibetha, Pocook, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 428, 1933.
Vernacular. — Khatas (Hindi, but not restricted to this
348
VIVERKIM!.
species); Mach-bhondar,Bdgda$,Pudo-ganda(Beng.); Bhran
(Nepal Terai) ; Nit-birahi (Nepal) ; Kung (Bhotia) ; 8aphiong
(Lepoha) ; NingalicMtua (Pahari).
Fig. 86.
A & B. Outer view of left auditory region of two skulls of Viverra
zibelha from Gorkha, Nepal, to show individual variation in the
size and shape of the bulla.
C & D. Hinder end of palate of two examples of the same species, showing-
individual differences of no systematic importance.
E. Fore part of skull of V. zibetha from the North Shan States.
F. Palate of skull of same from Bhutan Duars.
(All figures J nat. size.)
VIVERRA. 349
Locality of the type of zibetha, Bengal ; of undulata, Nepal ;
of orientalis and melanwus, Nepal ; of civettoides, Darjeeling.
Distribution. — Nepal*, Sikkim, Bhutan, Upper Bengal,
and apparently S. Kamrup in Assam.
Winter coat thickened with underwool and the contour
hairs long, those of the dorsal crest at their greatest length
in February or March, about 60 to 70 mm., and of the flanks
about 35 to 40 mm. Summer coat with little or no underwool
and the contour hairs of the crest up to about 40 mm., of the
flanks about 20 mm. Skins from Sevoke, 500 ft., in Upper
Bengal (Crump), and from Bhutan Duars, 600 ft. (Baptista),
resemble in the length of the contour hairs skins from Sikkim
at various altitudes from 2,000 ft. to 6,000 ft. (Crump and
Baptista). General colour varying from deep tawny through
all shades of that tint to grey, specimens from the same locality
and killed on the same day sometimes differing considerably
in hue. Pattern equally variable. At its best in the new
short summer coat it may consist of conspicuous blackish
transverse stripes on the fore quarters and of loops and
rosettes on the hind quarters ; but at the same locality
and date it may be brown and much less conspicuous. In the
long winter coat it is not well defined, hardly traceable on the
fore quarters, and represented by obscure mottling and
brindhng on the hind quarters. Sometimes it is altogether
evanescent.
* I have seen no skins of this Civet from the Himalayas west of
Nepal, and feel sure Col. A. P. Ward was mistaken in recording it from
Kashmir (Journ. Bomb. Nat. Hist. Soe. xxxi, p. 8, 1927). He said it
is very rare, but occasionally found towards Chamba. His flesh-
measurements, head and body 30 in., tail 17 in., weight 22 lb., are
correct enough; but his remark that Blanford's skull-measurements
are far larger than anything in the western Himalayas shows that the
skull he had did not belong to this species. His further statement that
the animal is found " often living under thatched roofs " suggests
confusion with the Kashmir Toddy-Cat (Paradoxurus), although he
cited the latter under a separate heading. I am not acquainted with
any other record of V. zibetha in Kashmir ; and Col. Stockley tells
me he never came across it in that country or in Kumaon, although
all collectors agree that it is one of the easiest mammals to trap.
According to Blanford this Civet is found in Orissa and Chutia Nagpur,
and Dunbar Brander tells me (in litt.) that his dogs killed a large Civet,
which he took for zibetha, in "that very wild country "Uprora." He
also saw one at Pachmarhi, as recorded in his book. This was " working
along a ledge on the face of a cliff, within gunshot range," but was not
worth shooting because of an intervening chasm it would have taken
12 hours or more to get round. It would be interesting to know to
what species these " large Civets " belonged. Whatever it may be,
it must be very rare in Peninsular India, since no specimen was secured
anywhere south-west of the Ganges by the collectors of the Bombay
Mammal Survey.
350 VIVBBEIDiE.
The flesh-measurements (in English inches) and weights
(in lb.) of some specimens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Banso Bahari, Nepal ; ad. J. 32f 18$ 5£ 21
Gorkha, Nepal ; ad. <J 294- 18 5 20
Gorkha, Nepal ; ad. $ 32f 14|- 5$ 20
Narbong, Darjeeling ; ad. 6" 31 16 5 14
Paahok, Darjeeling ; ad. <$ . 30$ 15$ 5$ —
Rongli, Sikkiin ; ad. $ 29$ 16$ 5$ 20
Sevoke, Bengal ; ad. ? 32 — 5f —
Sevoke, Bengal ; ad. ? 30f 16$ 5$ —
Rajapara, S. Kamrup ; ad. <J 32$ 16$ 6$ —
Rajapara, S. Kamrup ; ad. $ 31f 16$ 6f —
There do not appear to be any appreciable differences in
size between the males and females. The greater length of
the feet in the pair from S. Kamrup (Wells) is probably due
in part to the inclusion of the claws.
39 b. Viverra zibetha pieta Wroughton.
Viverra zibetha picta, Wroughton, Journ. Bomb. Nat. Hist. Soe.
xxiv, p. 64, 1915, and xxvi, p. 46, 1918; Poeock, Journ. Bomb.
Nat. Hist. Soc. xxxvi, p. 432, 1933.
? Viverra zibetha surdaster, Thomas, Proc. Zool. Soo. 1927, p. 46,
and 1928, p. 145 ; Poeock, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 433, 1933.
Vernacular. — Ghoung Divin Se (Burmese, Upper Chindwin) ;
Kyoung-myeng (Burmese).
Locality of the type of picta, H'Kamti, 500 ft., on the Upper
Chindwin ; of surdaster, Xien Khouang, Laos.
Distribution. — Upper Bubma. and Assam, east of the Brahma-
putra apparently as far south as the North Shan States ;
also Laos, Tong-king, and Annam probably.
Resembling typical zibetha in the marked difference in
length and luxuriance of coat and distinctness of pattern
between summer and winter skins, but differing on the average
in having the pattern a little less obliterated in the winter
and showing more distinct brindling or spotting on the blackish-
grey hue of the shoulders and fore quarters.
This race is of doubtful status. It was founded by Wroughton
on a single specimen from H'Kamti, 500 ft., on the Upper
Chindwin River in Upper Burma, which was killed in July,
and exhibits a bold brown pattern on the tawny ground-colour
of its short coat. Wroughton contrasted it with winter
skins of zibetha, overlooking its close similarity to the Nepalese
example of the latter, which Hodgson named civettoides. He
subsequently withdrew the name, relegating it to the synonymy
of zibetha on discovering, as he thought, that the distinctness of
VIVBERA. 35 L
the pattern is not a racial character. But the examination
of a long series of skins shows that the race is admissible
on the character I have mentioned, although summer skins
are not distinguishable from those of typical zibetha.
In addition to the type, numbers of skins have been examined
from the following localities : — Golaghat, 300-400 ft., the
Garo Hills, 3,000-4,000 ft., the Jaintia Hills, 4,000 ft., and the
Naga Hills, 3,500-4,500 ft. As in typical zibetha, the length
of the winter coat does not appear to be affected by altitude,
the contour hairs being as long in January skins from Golaghat
as in March skins from the Garo Hills.
The probable extension of this race as far south as the
North Shan States is attested by skins from Gokteik and
Pyaunggaung (Shortridge), which, although in full moult,
are long-coated as in picta and zibetha, and thus differ from the
following race, pruinosa, to which Wroughton assigned them.
Mesh-measurements (in English inches) and weights (in lb.),
of V. %. picta and pruinosa are as follows : —
Head and Hind
Name, locality, and sex. body. Tail. foot. Weight.
picta.
Sadiya, Assam ; ad. <J 34 17f of —
Golaghat, Assam ; ad. $ ... 30 17} 4f U{
Jaintia Hills, Assam ; ad. $ . 32* 16f 5l 16
Garo Hills, Assam ; ad. $ . . 29-J 144 4* 18|-
H'Kamti, Upper Chindwin
(type); ad. <J 32 17 5 17|
Pyaunggaung, 3SJ. Shan
States ; ad. <? 33| 183 5| 20
Pyaunggaung, N. Shan
States; ad. <J 33| 17i 5J 18
pruinosa.
Bankachon, Tenasserim ;
ad. (J 31| 17 5 21
Thaget, Tenasserim (type) ; 21
ad. 30f 171 5|- 19
The skull and teeth of picta do not differ from those of
zibetha*.
* The race described by Thomas as surdaster was based upon a youngish
specimen from Laos and was distinguished by the small size of the
auditory bulla, measuring 15 mm. But the bulla of a fully adult
skull of zibetha from Nepal is the same length and 5 mm. shorter than
the longest bulla in skulls of zibetha. Skins assigned to surdaster have
been recorded also from Tong-king, Annam, and Cochin-Ohina.
Additional material may show that the race is valid ; but for the present
its status is doubtful. To the north no doubt both picta and surdaster
intergrade with the southern Chinese race, ashtoni ; but the latter is
longer in the coat, a January skin from Fukien having the crest and
flank hairs about 90 and 50 mm. long respectively.
354 vrvEKKnxE.
occurs from the base of the foothills up to 7,000 ft. or more,
being most abundant at an elevation of about 3,000 ft. It is
purely nocturnal, is a great thief among the village chickens,
-and can be easily lured to a trap baited 'with meat. It appears,
nevertheless, to be equally addicted to a vegetable diet,
feeding to a great extent on berries, and having a special
liking for eardamons. The stomachs of specimens examined
by Hodgson contained the remains of fowls, clearly taken
from a refuse-heap near a kitchen, of rats, shrews, and frogs.
The omnivorous diet of the animal, noted by several
•observers, was summarized by Blanford's statement that it
kills any birds or small mammals it can capture, and also
feeds on eggs, snakes, frogs, insects, as well as upon fruits
and some roots. It lies up in woods, under bushes, in thick
grass or in holes, probably dug by some other animal. There
are said to be three or four young to the litter, born in May
or June ; but Hodgson's suggestion that the eyes are open
.at birth is probably untrue. The newly-born young are black,
with white on the upper lip, the inner surface of the ear,
some rings on the tail, stronger below than above, and sometimes
at least with the two white collars on the throat faintly showing.
On the "Upper Chindwin this Civet, represented by picta,
is, according to Mackenzie, fairly common and a great fowl
thief. He shot a pair in the jungle fishing in a small pool
by a stream. Their stomachs contained respectively four and
five practically undigested fishes, about 3 in. long, with the
heads bitten off. No trace of the heads was to be found.
This seems to be the first record of V. zibetha being a fish-eater.
Shortridge recorded the race as plentiful at Pyaunggaung in
the North Shan States, where, like Viverricula, it appears to
attach itself to the vicinity of villages and is probably a
scavenger and a poultry-thief. He here confirms the experience
•of all collectors that V. zibetha is very easily trapped.
Nothing has been recorded of the habits of the southern
race, pruinosa, which, according to Shortridge, is plentiful
at Bankachon and, like all the Civets, very easily trapped.
Genus MOSCHOTHERA Pocock.
Moschothera, Poeoek, Journ. Bomb. Nat. Hist. Soo. xxxvi, p. 4A1,
tart-figs. 3 &. i, 1933.
Type of the genus, Viverra civettina Blyth.
Distribution. — Travancoke and from Loweb Btoma to
Siam, Indo-China, and the Malay Peninsula.
Distinguished from "Viverra by the complete absence of
skin-lobes acting as claw-sheaths on the 3rd and 4th digits
of the fore foot and by the scanty hair-growth on the fore and
hind feet between the plantar and digital pads. The skull
MOSCHOTHERA.
355
differs in the suppression of definite frontal postorbital
processes, which are represented merely by ridges marking
the anterior attachment of the temporal muscles ; the sagittal
Fig. 87.
A. Lower side of left fore paw of the Malabar Civet (Moschoihera
civettina) .
B. The same of left hind paw, showing the greatly reduced, disc-lite
remnants of the two metatarsal pads.
<3 & D. Lower side of left fore and left hind paws of the Large-spotted
Civet (Moschothera megaspila).
crest is a low ridge and the nasals and whole facial portion
of the skull are longer, the distance from the position of the
2a2
356 viveeeid^;.
suppressed postorbital processes to the tip of the premaxilla
exceeding the distance from the former point to the tip of the
occipital crest.
The two species assigned to this genus have long been
known, but not intimately, there being no information
derived from fresh specimens regarding the structure of the
scent-pouch or of the feet, the recorded characters of the feet
being supplied by relaxed skins.
The two are widely separated geographically, and by most
authors who have discussed them, including Blyth, Blanford,
and Lindsay, have been regarded as distinct species ; but
Robinson and Kloss, who detected their close kinship, con-
sidered them to represent merely local races of a single species.
Their wide geographical severance precludes, however, the
probability of the discovery of intermediate forms, and the
evidence is in favour of the earlier view of their kinship.
They may be briefly contrasted, as follows : —
a. Soles of the feet round the plantar pads com-
paratively well clothed with hairs ; no trace
of metatarsal pads. Skull with very 'weak
temporal crests on the frontal bones, lower
edge of mandible less emarginate behind and [p. 356.
teeth smaller megaspila Blyth,
a'. Soles of the feet comparatively naked ; rem-
nants of the metatarsal pads persistent.
Skull with stronger temporal crests on the
frontal bones, lower edge of mandible notice- [p. 358
ably emarginate behind ; teeth larger eivettina Blyth,
From the small number of preserved skins, it is evident
that the Civets of this genus are rare, possibly approaching
extinction. When additional material comes to hand other
differences than those enumerated will probably be discovered.
40. Mosehothera megaspila (Blyth). The Large-spotted Civet.
Viverra megaspila, Blyth, Journ. As. Soe. Beng. xxxi, p. 331, 1862 ;
id., Proc. Zool. Soc. 1864, p. 484; Gunther, Proc. Zool. Soc.
1876, p. 428, pi. 37 ; Blanford, Mamm. Brit. India, p. 99,
1888; Robinson & Kloss, Bee. Ind. Mus. xix, p. 175, 1920;
Lindsay, Journ. Bomb. Nat. Hist. Soc. xxxiii, p. 146, pis. 1 & 2,
1928.
Mosehothera megaspila, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 443, text-figs. 3 & 4, 1933.
Vermicular. — Kyoung-myeng and Kyaung-myeng-Kwet (Bur-
mese) ; He.n-H.awn and Amnge (Shan) ; MlHsang-jebat (Malay).
Locality of the type, Prome in Lower Burma.
Distribution. — Centbal and Sottthbek Burma, Malay
Peninsula, Siam, and southern Indo-Chma.
Soles of the feet partially naked only in front of the plantar-
pad, the webbing between the 3rd and 4th digits hairy as
MOSCHOTHERA. 357
far back as the pad, but the hairy patches on the other
interdigital webs not reaching so far inwards ; the skin at
the sides of the plantar pad and above it entirely covered with
hair, completely isolating the pads of the pollex and hallux
and the carpal pad ; no remnants of metatarsal pads traceable.
Hairs of the crest from about 50 to 100 mm., averaging
about 60 mm. ; contour hairs of the flanks from 25 to 30 mm.
General colour varying from silvery-grey to golden-buff
or tawny, and the pattern from black to brown, the spots
large or comparatively small, separated or sometimes fusing
into blotches or into vertical stripes behind the shoulders
or into a pair of dorsal stripes bordering the median crest,
the white bands on the tail very seldom forming complete
rings, mostly restricted to the sides and lower surface.
Some individual differences between two specimens from
Tenasserim may be recorded. The first, from Victoria Point,
Nov. 27, in early winter coat, has the crest-hairs 48 mm.,
the flank-hairs 30 mm., the general colour grey, with a decided
ochreous-tawny wash on the shoulders and pale tawny
behind, and a pattern of small, scattered, indistinct spots
on the shoulders, of large black well-separated spots on tas
flanks, the uppermost forming on the loins a longitudinal
stripe on each side of the crest ; the tail has seven white
stripes below, the first three reaching some distance up the
sides, the next three restricted to the ventral surface, and the
last completely encircling the tail near its apex. The second,
from Tenasserim Town, March 9, in late winter coat, has the
crest-hairs 67 mm., the tint slightly tawnier, and the tail
with only five white rings, dorsafly incomplete and restricted
to its proximal portion, the distal portion being entirely black.
The only other available Burmese skin, from Allagappa,
30 miles west of Sagaing in Upper Burma, shows the beginning
of the moult, the crest hairs being about 70 mm., the flank-
hairs 25 mm., the general colour much greyer, with only
a faint buff wash on the fore quarters, the pattern black and
standing boldly out against the silvery-grey ground-tint.
The rest of the skins examined, all from countries outside
the limits of the British Indian fauna, are undated. They
show considerable variations in coat, colour, and pattern.
One from Nan in Siam is pale tawny-grey, with the pattern
golden-brown. One from Penang is like the latter, but has
the pattern deeper brown. One from Nahtrang in Annam,
1,200 ft., resembles the Tenasserim Town skin, but has the
•crest-hairs only 40 mm., the flank-hairs 27 mm. Three
from Cochin-China differ greatly from each other. In one
the crest-hairs reach the extraordinary length of 100 mm.,
"the flank-hairs being 30 mm. ; the general hue is grey washed
with tawny, as in the Tenasserim skins, but the spots are
358 VIVERBIDiE.
smaller, showing a tendency to run into blotches and stripes.
In another the contour hairs are of average length, the general
tint much deeper tawny, and the pattern forms two vertical
stripes on the fore part of the flanks. The third also has the
contour hairs normal, but the general hue is golden-buff,
with many of the large, widely separated spots rusty-brown,
those adjoining the crest forming a complete longitudinal
stripe on each side.
The material is quite insufficient to show whether the species
is differentiated into distinguishable local races or not.
The flesh-measurements (in English inches) and weights,
(in lb.) of the two specimens of this species collected by
Shortridge in S. Tenasserim and the measurements of the
only available specimen of the other species, civettina, are as
follows : —
Name, locality, and sex.
Head and
body.
Tail.
Hind
foot.
Weigh*
megaspila.
Tenasserim Town ; ad. $ . .
. . 31J
30
161
14|
5+
18*
19
civettina.
30
13
—
Ui
Except that the tail is a little shorter, only about half the
length of the head and body, the dimensions agree closely
with those of Viverra zibetha.
Habits. — There appears to be very little special information
about the habits of this species, which Blanford states are similar
to those of V. zibetha. According to a note by Shortridge
on specimens observed at Victoria Point, Tenasserim, it is
very similar to V. zibetha in general appearance, except that
the head is more massive and has a swollen appearance about
the muzzle ; but it does not smell nearly so strongly of
" civet " as V. zibetha. Incidentally he added that he had
never seen any kind of Viverra climb a tree, although no
doubt they are well able to do so if they choose. The Utter,
according to Cantor's observation in the Malay Peninsula,
consists of from one to three cubs.
41. Mosehothera civettina (Blyth). The Malabar Civet.
Viverra civettina, Blyth, Joum. As. Soc. Beng. xxxi, p. 332,1862 ;
other references as under the preceding species.
Mosehothera civettina, Poeoel:, Joum. Bomb. Nat. Hist. Soc. xxxvi,
p. 445, text-figs. 3 <fe 4, 1933.
Locality of the type, Travancore.
Distribution. — The coastal district and Western Ghats of
S. India.
MOSCBCOTHEBA.
359"
.3
•a*
§
&
•ca
«
■51
•I
s
02
ft
6J-S
-3 3-
3 "1 IP
"S.-SS
"3 £
9
£
c8
^ *o
c3
cS
* 1
g
£
«3
j360 VIVERBIDiE.
Distinguished principally from M. megaspila in external
•characters by the greater nakedness of the soles of the
feet, the hairs on the interdigital webs being between the
digital pads, forming submarginal patches, with the skin
in front and at the sides of the plantar pad naked, this naked
area extending above the pad on each side as far up as the
carpal pad on the fore foot and to the level of the hallux on
the hind foot, with a hairy median strip between them, reaching
to the middle of the plantar pad behind. On the hind foot
also remnants of the metatarsal pads persist as two naked
spots, the external a little above the level of the hallux, the
internal considerably higher. These differences, however, are
based on the examination of a single example of M . civettina
that died in the Zoological Gardens in Trivandrum, and may
prove to be subject to individual or seasonal variations.
The coat in this specimen is tolerably long and full, the
contour hairs of the crest and flanks being about 50 and
30 mm. respectively. The ground-colour is clear grey,
nearly matching the specimen of megaspila from Sagaing ;
the pattern is black or nearly so, the spots being smaller
and more closely set than in most, but not all, skins of
megaspila, but more spotted on the shoulders and chest.
The tail has five white rings extending farther up the
sides than in any skins of megaspila, the last being only
about two inches from the tip, whereas in megaspila the
terminal six inches or so of the tail are typically black.
There is also more white on the muzzle and chin than in
megaspila, accompanied by slight differences in the bands
of the throat. But on the evidence of a single skin of civettina
definite differences in colour and pattern between the two
.species cannot be established.
The flesh-measurements and weight of this example are
entered above (p. 358). Although the tail is shorter than
in the two specimens of megaspila from Tenasserim, the length
of that organ in individuals of Viverra zibetha is too variable
for it to be trusted as of systematic importance in this instance.
The skull is distinguishable from the three skulls of megaspila
available for examination by having two strongish V-shaped
temporal ridges running forwards from the sagittal crest to
the postorbital frontal bosses, with the waist a little more
constricted, although the animal was younger. Also the
lower edge of the postdental portion of the mandible is
markedly concave just behind the level of the last lower molar.
Habits.— Apparently all that is known of the habits and occur-
rence of this Civet is contained in the paragraph, quoted by Blan-
ford,from Jerdon's volume -.-—"The Malabar Civet Cat is found
throughout the Malabar coast from the latitude of Honore
MAMMALIA.
PLATE XXVII.
^y
*$j. '
0*#*#*"
&& .
, .fitoC ' (*£, '-'.i&sZ.,
Malabar Civet (Moschothera civettina).
Photo W. S. Berridge.
Little Civet (V/verncu/a indica).
MOSCHOTHEEA.
361
Fig. 88.
A. Palate of skull of Moschothera civettina from Trivandrum.
B. Frontal area of skull of the same.
C The same of skull of M. megaspila from Tenasserim TWn.
D, E, F. Posterior portion, of mandible of Raines's skull of M. civettina,
of skull of same species from Trivandrum, and of skull of
M. megaspila from Tenasserim Town. (All figures f nat. size.)
362 VIVEBRDME.
(Honawar) at all events to Cape Comorin. It inhabits the
forests and the richly wooded lowland chiefly, but is occasionally
found on the elevated forest-tracts of Wynaad, Coorg, etc.
It is very abundant in Travancore, and I have procured it
close to my own house at Tellicherry. I have never obtained
it from the Eastern Ghats nor from Central India. It is
stated by the natives to be very destructive to poultry."
Since, however, the collectors for the Mammal Survey failed
to get a single specimen, it seems that the species must be
very much rarer than in Jerdon's time. In the first decade
of the present century the late Harold Ferguson did his best,
at my request, to secure specimens for the Zoological Society
in London, but without success ; and in 1923 A. P. Kinloch
omitted the species from his list of the dominant mammals
of the Nelliampathi Hills, although his remark that there
is " another Civet up here, judging from its droppings con-
siderably larger than the Toddy-Cat, but I have never seen
it," may have referred to this rare animal. The facts suggest
that the species is on the wane.
Genus VIVBRRICULA Hodgson.
Viverrioula, Hodgson, Ann. Mag. Nat. Hist, i, p. 152, 1838 ; id.,
Journ. As. Soc. Beng. x, p. 909, 1841 ; and of most subsequent
authors, including Blanford, Mamm. Brit. Ind. p. 100, 1888 ;
Poeoek, Proc. Zool. Soc. 1915, pp. 136 & 147 ; id., Journ. Bomb.
Nat. Hist. Soc. xxxvi, p. 629, 1933.
Viverrula, Hodgson, Cale. Journ. Sci. ii, pp. 53—5, pi. i, 1842
(by error for Viverricula and misprinted Viricaula in expl. of
plate).
Type of the genus, V. indica (Geoffr.).
Distribution. — Peninsular India from Sind, the Punjab,
and the foothills of the Himalayas, Ceylon - , Burma, S. China,
and thence southwards through the Malay Peninsula to Sumatra
and Java. Artificially imported and now wild in Madagascar,
Sokotra, and possibly elsewhere.
Distinguished externally from Viverra and Moschothera by
the absence of the dorsal crest and by the insertion of the
ears, the anterior edges of which are set closer together on
the forehead, a feature which, in conjunction with the shorter,
more pointed muzzle, gives a more alert aspect to the face.
The feet are as in Moschothera, but the scent-gland is relatively
not so well developed as in Viverra. The neck-stripes are
similar in arrangement, but narrower, more variable, and not
set off by pale interspaces to the same extent ; the body-
pattern consists of small spots on the fore quarters, larger
spots, tending to run into longitudinal lines, on the flanks,
and of six to eight stripes down the back. Skull distinct
from that of Viverra and Moschothera in many particulars.
The muzzle is short and weak, the cranial portion long,
VIVERRICTTLA. 363
narrow, and compressed above posteriorly, the subocular
portion of the cheek is longer, 35m* being set farther back,
the infraorbital foramen approximately above the point
of contact of pm z and pm z , and the suture between the
zygomatic arch and the maxilla very long, about as long as
the median length of the nasals, and half the length of the
upper cheek-teeth and the width across the occipital condyles ;
the auditory bulla is also relatively much longer, exceeding
the width across the condyles. The teeth too are more
compressed and trenchant, and the first upper molar has
a narrower inner lobe.
All the different forms of this Civet are regarded as repre-
senting a single specie? which has the characters and distribu-
tion of the genus.
42. Viverrieula indiea (Geoffroy). The Little Civet.
Civetta indiea, Geoffroy, Cat. Malum, p. 113, p. 1803.
Viverrieula malaccensis of all recent authors to 1933, but not
Viverra malaccensis Gmelin, Syst. Nat. i, p. 92, 1788.
Viverrieula indiea, Pocock, Journ. Bomb. Nat. Hist. Soc. xxxvi,
pp. 629-31, 1933*; for other bibliographical references see
under the subspecific headings.
42 a. Viverrieula indiea mayori Pocock.
Viverrieula indiea mayori, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 632, 1933 ; Phillips, Man. Mamm. Ceylon, p. 168,
1935.
Vernacular. — Urulaera (Sinhalese) ; Poolu poona, Ve®u
(Tamil) ; Pulugoo Poonai (Jaffna Tamil) .
Locality of the type, Maha Oya in the Eastern Province of
Ceylon.
Distribution. — Ceylon. " Common all over the island, both
in the hills and in the low country" (Phillips).
Distinguished from the typical S. Indian race, indiea,
described below, by its slightly smaller average size and
shorter tail and by the absence of noticeable change in colora-
tion and in the length and thickness of the coat, at least from
May to December. The general colour is on the whole
tolerably uniform, varying from grey to buffish-grey ; the
pattern is distinct and blackish-brown or black, but varies
considerably in boldness not only in the width of the dorsal
stripes but in the extent to which they are broken up or more
or less coalescent. No two examples are exactly alike ;
two from the same locality may be dissimilar, and each may
closely match another from a remote part of the island.
* It is needless to repeat here my reasons, given at some length
in this paper, for discarding for this species the name malaecensis
applied by Gmelin to the figure and description of an alleged Civet
from Malacca published by Sonnerat, who used a Genet from the Cape
as his model. Sonnerat's account does not agree with any known
form of Viverrieula, and there is very little evidence that the latter
was included in his composite description.
364 VIVEBEIDJE.
42 b. Viverrieula indica indica (Geoffroy).
Civetta indica,~Q6o3xoy, Cat. Mamm. p. 113, 1803*.
Viverra indica, Desmarest, Nouv. Diet. d'Hist. Nat. vii, p. 170,
1817 ; id., Mamm. i, p. 210, 1820 ; Sykes, & Horsfield, Proc.
Zool. Soo. 1832, p. 22 ; Geoffroy, Mag.' de Zool. 1836, p. 10
(in part, specimens from Malabar).
Viverrieula malaceensis indica, Robinson & Kloss, Eec. Ind. Mus.
xix, p. 177, 1920.
Viverrieula indica indica, Pocock, Journ. Bomb. Nat. Hist. Soe.
xxxvi, p. 633, 1933.
Vernacular.— Punagina-Bekku (Kanarese) ; Fowadi-Manjur
(Marathi) ; Puluk-Philli (Wadari) ; Punkassibekku (Haran Shi-
karis) ; Mushak-billi (Dekhani) ; Pulunguotay Punugu (Coorg).
Locality of the type, " India " ; Western Ghats (Sykes and
Horsfield) ; Dharwar (Robinson and Kloss).
Distribution. — Sotjthebn India from the Western to the
Eastern Ghats and, according to Robinson and Kloss, as far
north as Lake Chilka on the east coast.
Coat very variable in length and texture before and after
the moult, the contour hairs in the fresh coat being from about
•20 to 25 mm. long and gradually increasing in length to
35 or 40 mm. and becoming harsh, brittle, and liable to be
broken at the tip before being shed. Colour also variable,
Tout, when the coat is soft and full, it is on the average relatively
dull, varying from brownish or olivaceous-grey to lighter grey,
occasionally only with a brighter ochreous tinge ; but in the
old coat the hairs of the interspaces apparently bleach to grey
or even silvery. The pattern is usually distinct, blackish
or brownish, but is sometimes obscured somewhat by an
ochreous tinge.
Specimens often vary considerably from the same locality
and at approximately the same time of year. Of five skins
from the typical locality, Dharwar, 2,400 ft. (Shortridge),
one, Nov. 9, has the coat coarse, shaggy, about 37 mm., and
the general tint almost silvery, with the pattern obscure,
especially on the tail. Another, Dec. 8, has the coat, smoother,
but equally thick with underwool and about the same length,
and the colour dull buffy-grey, not bleached, with the pattern
more conspicuous. The others are intermediate. Examples
from other localities in S.W. India, from Satara (Prater),
N. Coorg, 3,550 ft., S. Coorg, 2,000-2,840 ft. (Shortridge),
collected from December to February agree fairly well with the
Dharwar skins, and two from the Mlgiris, 3,000-4,000 ft.,
collected in July, although like the unbleached Dharwar skin of
Dec. 8 in tint, have the coat harsh and short, 25 to 30 mm.
From the Eastern Ghats (Baptista), two from Kumool, May,
* Since only a few copies of this work were printed and privately
given to friends by Geoffroy, its publication is open to dispute, and
Desmarest may be regarded as the author of the name, although he
gave Geofiroy the credit of it.
VIVERRICTTLA. 365
closely resemble the long harsh-coated, bleached November skin
from Dharwar, except that the underwool is scanty and the
pattern more distinct.
Of three skins from the Palkonda Hills (Baptista), two,
July 24 land Aug. 2, have the coat harsh, shabby, and dead,
and are also like the bleached Dharwar skin, but not so silvery,
with the coat shorter and the pattern even more obscure ;
the third, July 15, is very different, and seems to have com-
pleted the moult earlier, the coat being full, soft, and fresh,
25 mm., and the coloration unusually black and white.
A skin from the Vontimitta Range, 325 ft., Sept. 14, also
has the coat fresh, soft, and full, but the general colour
normal ; and one from the Denkanikota Range, 3,062 ft.,
Oct. 22, is similar, whereas a July skin from Salem, 1,000 ft.,
has the coat coarse, brittle, and bleached.
From these skins it seems that the date of the moult is
inconstant. Typically the coat is harsh and more or less
bleached from May to August with the moult in progress,
and full and soft from September to February, but there are
individual exceptions to this generalization.
The following table of flesh-measurements (in English
inches) and weights (in lb.) of the Ceylonese and. S. Indian
races, mayori and indica, shows clearly the average superiority
in size of the latter over the former. The data of mayori
are taken largely from Phillips's volume. The dimensions
of the two $ specimens with localities are respectively those
of the largest and smallest of this sex in the British Museum
collected by Dr. Mayor, who failed to secure an adult $, thus
bearing out Phillips's statement that the <J is more abundant
than the 0.
Head and Hind
Name, locality, and sex. body. Tail. foot. Weight.
mayori.
Phillips's largest ; ad. <? 23| 15 4 6£
Maha Oya (type) ; ad. $ 2a 14$ 3i 6J
"Weligatta ; ad. <? 21 13§ 3f 4f
Phillips's average of 12; ad. #. 21$ 13| 3| 5$
Phillips's largest ; ad. $ 22A 14A 3f 5#
Phillips's average of 5 ; ad. $ 20 j 13+ 3| 5J—
indica.
Dharwar ; ad. <$ : . . 2S| 16f 4 —
Medha, Satara ; ad. <J 24£ 15| 3f 8
Dharwar, ad. <J 234 15| , 3|- —
Benhope, Nilgiri Hills ; ad. «}. 21f 14$ 3+ —
Dharwar ; ad. S 23f 16$ 3f —
Virajpet, S. Coorg ; ad. ? 23i 14? 3| 7
Dharwar ; ad. $ 22| 15f 3J —
Palkonda Hills ; ad. ? 22$ 15$ — 5
The skull-measurements confirm the flesh-measurements in
attesting the average inferiority in size of mayori to typical
indica.
366
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VIVERRICTTLA. 367
42 c. Viverricula indica bengalensis (Gray).
Viverra bengalensis, Gray & Hardwicke, 111. Ind. Zool. i, pi. 4,
1832.
Viverricula malaccensis bengalensis, Robinson & Kloss, Ree. Ind.
Mus. xix, p. 177, 1920.
Viverricula indica bengalensis, Pooock, Journ. Bomb. Nat. Hist.
Soo. xxxvi, p. 637, 1933.
Vernacular. — Mashk-billa, Katus (Hindi), Qundha Gokal,
Gando gaula (Beng.) ; Jdbadio (Cutch).
Locality of the type, " most part of Bengal," according to
Gray ; Calcutta, as restricted by Robinson and Kloss.
Distribution. — The plains of Northern India, south of the
Ganges, from Calcutta to Gujerat, and possibly Sind.
A race provisionally admitted on the evidence of several
skins, from scattered localities, which are on the average
paler and greyer than those assigned to indica from Southern
India, none exhibiting the dark olivaceous or ochreous-brown
hue of some examples of indica, but the grey examples of the
latter are practically indistinguishable from them.
The individual and seasonal variations are very similar
to those of indica. The coat is from 30 to 35 mm. in April skins
from Hazaribagh and Hoshangabad (Crump), in a March skin
from Gujerat, and a December skin from Kathiawar, from
20 to 22 mm. in a July skin from Sehore and a June skin from
Gujerat. The ground-colour in the fresh coat ranges from
yellowish- and greyish-buff to silvery-grey ; the dead harsh
coat, bleaches grey, and in some skins the hind-body is bleached
and the head and fore quarters well coloured. The pattern
may be black and sharply denned or considerably diluted with
ochreous and comparatively inconspicuous.
In addition to the skins from the localities above referred to,
there is one, undated, from Larkana in Sind (Prater). The
coat is shaggy, harsh, and long, 38 mm., the colour is grey,
and the pattern deep brown. It closely resembles two skins
of indica from Kurnool killed in May, but, pending the collection
of additional material, the skin may be provisionally assigned
to bengalensis.
According to the measurements entered below bengalensis
is a little shorter in the head and body than indica with
the same length of tail. The skull, too, is a little shorter
on the average than in indica ; but hardly differs in other
respects. The claim of Robinson and Kloss that the skull
of bengalensis is larger and has larger bullse is not borne out
by additional specimens.
368 VIVBEBID3!.
42 d. Vlverricula indiea deserti Bonhote.
Viverricula malaccensis deserti, Bonhote, Ann. Mag. Nat. Hist.
(7) i, p. 120, 1898.
Vwerricula indiea deserti, Pocoek, Joum. Bomb. Nat. Hist. Soo.
xxxvi, p. 639, 1933.
Locality of the type, Sambhar, near Jaipur in Rajputana.
Distribution. — Rajputana, so far as at present known.
Distinguished from the proceeding races by its longer,
shaggier coat, which in late summer and early winter is longer
A.
B.
Fig. 89.
Upper view of skull of Viverricula indiea wellsi from Kangra.
Lower view of occipital region, of the same, showing the auditory
bullse.
The same of a skull of V. indiea baptwtse from Bhutan Duars.
than the longest winter coats in indiea and bengalensis.
The " waist " or postorbital area of the skull is also narrower.
This race is provisionally admitted on the evidence of
two specimens only. The type, from Sambhar (Adam),
December, has tolerably luxuriant underwool, and the contour
hairs 45 mm. long ; the general colour is grey, and the stripes
and spots form an obscure, irregularly-clouded pattern,
a mixture of grey, ochreous, and dusky brown, but no black,
VIVBBRICULA. 369*
A second specimen, from Nasirabad, August, has the coat
42 mm., the ground-colour buffy-grey, and the pattern
obscure, mixed black and rusty ochre, and hardly forms
any definite stripes.
The skull of the type, the only one known, is very similar
to that of bengalensis and indica except for the exceptional
narrowness of the " waist," which is not an age character ;
the distance between the bullae is also less than in the skulls-
of the previously described races. Until additional specimens-
are procured from Bajputana the precise status of this race
must remain in doubt.
An adult <J from Dagshai, near Simla, 6,000 ft. (Dunn),
Jan. 26, closely matches bengalensis in size, in its grey colour,
and well-defined black pattern, but has the coat a little longer,.
38 mm. The skull is like the smallest of the bengalensis skulls,
an adult <J from Hoshangabad, but the waist is narrower and
the bulla shorter. By its colour, pattern, smaller skull, and
upper carnassial and shorter bulla it differs from the race
from Kangra described below, which from its distribution it
might be expected to resemble ; and apart from colour it
similarly differs from deserti. Additional specimens from
this district of the Upper Punjab are required before the
status of this skin can be determined.
42 e. Viverricula indica wellsi Pocock.
Viverricula indica wellsi, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 640, 1933.
Vernacular. — MaVpusa (Hindi in Kumaun).
Locality of the type, Kangra, 2,000 ft.
Distribution. — Kahstgba, Kumaun, and the United-
Pbovxstces.
Closely resembling deserti in the length of the coat and
the coalescence and confusion of the dorsal pattern, but
differing in being on the average more richly tinted in general
colour.
In a series of five winter skins from Kangra, 2,000 ft.
(Wells), February and March, the coat ranges from 25 to 46 mm.
long ; the general colour is mostly buff, with the pattern very
obscure owing to its ochreous tinge blending with the ground-
colour. In one skin the pattern is more distinct, with some
black in it, and in another the ground-colour is bleached grey.
Two skins from Eamnagar, Kumaun, 1,500 ft., January, and
ten undated skins from the United Provinces (Burke) closely
match the Kangra series.
The following table of flesh-measurements shows approxi-
mate equality in the length of the head and body to the
S. Indian form (indica) and slight superiority to the Plains:
vol. i. 2b
370 VrVBKEIDJB.
form (bengalensis), but the tail is apparently on the average
shorter even than in bengalensis.
The skull is very like those of these two races, but is longer
than in bengalensis, and has the postorbital area narrower
•than in either, although not so narrow as in deserti.
The flesh-measurements (in English inches) and weights
(in lb.) of specimens from the Plains of Northern India
(bengalensis), from Rajputana (deserti), and from Kangra
and Kumaun (wellsi) are as follows : —
Head and Hind
Name, locality, and sex. body. Tail. foot. Weight.
Hazaribagh, Bengal ; ad. <J . . 23f 16$- 4 6J
Sehore, C.I. ; ad. <J 22f 15£ 3£ —
Hoshangabad ; ad. <J 21f 14f 3£ 5
Kathiawar ; ad. $ 22f ISf 3f 5J
Danta, Gujerat ; ad. ? 21£ 16J 3f 5*
Hazaribagh ; ad. $ 21f 15 3{ 6
deserti.
Sarabhar, Rajputana (type) ;
ad. 3 23- 16- 4- —
Kangra, 2000 ft. ; ad. <J 25| 14* 4 —
Kangra ; ad. <J 24$ 15£ 3J —
Kangra ; ad. <J 24f 14f 4 —
Kangra ; ad. ? 22 14f 3£ —
Kamnagar, Kumaun ; ad. $ . 22f 14f 3| 6J
A number of unsexed old skins collected by Hodgson in
Nepal, without further particulars, resemble the next race
rather than the last in shortness of coat and distinctness of
pattern ; but in cranial characters they are nearly inter-
mediate, as might be expected from their distribution.
42/. Viverrleula indica baptists Pocock.
Vwerrieula indica baptistse, Pooook, Journ. Bomb. Nat. Hist.
Soc. xxxvi, p. 43, 1933.
Vernacular. — Saiyar, Bag-nyul (Nepal Terai).
Locality of the type, Hasimara in Bhutan Duars.
Distribution. — Irom Bhutan and Upper Bengal to Assam.
Distinguished from wellsi and deserti by its shorter coat
and more distinct pattern, but closely resembling wellsi in its
bright colouring, which is brighter than in deserti, bengalensis,
and indica. It is also decidedly smaller in its flesh-measure-
ments.
A series of ten skins, collected between Nov. 7 and April 20,
have the coat from 27 to 35 mm. long, the general colour
buff or ochreous, occasionally with a grey or ohvaceous cast,
VIVEEBICULA.
371
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372 VIVBBBID^E.
and the well-defined pattern varying from blackish to rufous-
brown. Specimens representing this race were collected
at Hasimara and Bhamabari, 600 ft., in Bhutan Duars
(Baptista), Haldibari, near Cooch Behar (Crump), Darbhanga,
150 ft. (Inglis), N. Kamrup, 300 ft., and possibly Sadiya
(Blanford) and Golaghat, 250 ft., in Assam. A specimen from
Mokokchung in the Naga HiUs, 4,500 ft. (MiUs), which in 1933
I assigned to this race, agrees better in size with the next race,
although the end of the tail is not so pale. It is probably
an intermediate form.
As the table of flesh-measurements shows, this race is a little
smaller than the other Indian races, coming very near the
Ceylonese race in that respect. The skull is about the same
length as that of mayori, but that of baptistse is more lightly
built, being narrower in all the width-dimensions given.
42 g. Viverricula indica thai Kloss.
Viverricula malaccensis then, Kloss, Journ. Nat. Hist. Sqe. Siam,
iii, p. 352, 1919 : Robinson & Kloss, Bee. Ind. Mas. xix, p. 178,
1920 ; Chasen, J. Siam Soc. Nat. Hist., Suppl. x, p. 41, 1935.
Viverricula indica thai, Pocoek, Journ. Bomb. Nat. Hist. Soc.
xxxvi, p. 650, 1933.
Vernacular. — Koung-ka-do (Burmese) ; Wa-young-kyoung-
byouk (Arakan) ; Kyaung-myin or Kyaung myeng (Burmese,
Mt. Popa).
Locality of the type, Prapatom, west of Bangkok, Siam.
Distribution. — Siam, Indo-Cbina, and Btjbma.
Distinguished from baptistse by the longer winter coat,
the tail conspicuously whiter at the end, owing to defective
black banding, and the skull larger, with the bullae more
widely spaced.
Two names were given to races of this Civet inhabiting
the mainland of S.E. Asia — V. indica thai, based upon specimens-
from Prapatom, about 40 miles west of Bangkok in Siam,
and V. indica klossi, proposed by myself in 1933 for specimens
from Penang, as a substitute for the inadmissible name
V. malaccensis malaccensis adopted by Kloss for specimens-
from the Malay Peninsula . The* Siamese form was distinguished
from the Malayan by having the buffy ground-colour slightly
paler and duller, with more black speckling, and the tail more
nearly white. In 1933 I assigned to thai several specimens
from Tong-king, Annam (Delacour and Lowe), and Cochin-
Chiaa, and one from 60 miles north of Raheng on the Me-wong
River, Siam, which seems to agree closely with the type from
Prapatom.
Provisionally I assign all the Burmese skins in the British
Museum to thai. Their colour, however, is extremely variable,,
and they seem to fall into two categories.
VIVERRICULA. 373
A skin from the Chin Hills, 4,000 ft., in Upper Chindwin,
April 5, with the coat 38 mm., the ground-colour bright
buff, and the pattern rusty, closely matches a skin from Saigon,
Cochin China. Another from Yin in Lower Chindwin, Sept. 18,
with the coat 33 mm., the fore quarters brownish, the flanks
grey, and the pattern black, resembles in its combined features
skins from Tong-king, Annam, and Cambodia ; but two
skins from Mt. Popa, 4,061 ft., Sept. 3, and the Allagappa
Valley, 30 miles west of Sagaing, have the coat thin, short,
harsh, and bleached to whitish-grey on the flanks, but the
pattern black and sharply defined. The extension of this race
to the southern end of the Irrawaddy Valley is attested by
a skin from Rangoon which in its brownish coloration resembles
the fore quarters of the skin from Yin in Lower Chindwin,
and suggests the unbleached phase of the skins from Mt. Popa
and the Allagappa Valley.
Skins of the second category came from the valley of the
Sittang ( J. M. D. Mackenzie) . Six from the delta of the Sittang,
some 40 miles south of Pegu Town, March, have the coat
decidedly less luxuriant and less shaggy than skins from
Upper Burma collected hi February and March ; the ground-
colour is ochreous or buff, the pattern well defined, blackish
or rusty black, and the tail is noticeably whitish at the end.
In brightness of tint these skins differ strikingly from the
skin from Rangoon and from most of the skins from Upper
Burma, except the one from the Chin Hills. One of three
skins from 30 miles north of Toungoo, Jan. 8, has the fresh
winter coat close and soft, not loose, and in colour and pattern
is very like the skins from the Sittang Delta. The other two,
on the contrary, March 18 and 23, have the coat thinner,
shorter, and harsher, the colour buffy-grey or silvery-grey
on the flanks, darker on the back, with the pattern black.
The coat is clearly dead and bleached, with the moult imminent,
and the skins closely resemble those collected on Mt. Popa
and in the Allagappa Valley in September and October.
In colour and shortness of coat the well-coloured Sittang
Valley skins are very like skins of klossi that I have seen from
the Malay Peninsula, and in 1933 I provisionally identified
them as that race ; but the tails are whitish at the end as in
thai, and on geographical grounds they are probably the
same as the Siamese race.
The flesh- and skull-measurements of the Upper and Lower
Burmese specimens are in tolerably close agreement. In
the table of skull-measurements are entered two from Chitta-
gong (B. B. Osmaston), which have no skins. That of the
• $ is the same length and width approximately as the skull
of the type of thai, also $, from Prapatom, and the skull of
the (J is the largest of Viverricula I have measured. The
374 vrvEKBiD-a:.
skulls from Toungoo and the Sittang Delta are a little larger
than skulls of typical khssi from Penang.
Clearly there may be at least two races, a northern and
a southern, represented in Burma. Additional specimens
are wanted, especially from Northern Burma and Tenasserim.
Until these come to hand it seems better to regard them as one.
The flesh-measurements (in English inches) and weights
(in lb.) of specimens from the Western Himalayan area
{baptistee) and from Burma (thai) are as follows : —
Name, locality, and sex.
Head and
Hind
body.
Tail.
foot.
Weight
21*
13f
3*
6
20f
13*
3*
—
20*
18*
3|
—
20f
Hf
3!
—
m
12!
3!
m
16±
3*
25!
16*
4
7*
22}
13!
3!
21
14!
»!
—
m
16!
n
—
Angarakata, N. Kamrup; ad. <J
Hasimara, Bhutan Duars ; ad. <J
Hasimara, Bhutan Duars ; ad. <J
Hasimara, Bhutan Duars ; ad. $>
Hasimara, Bhutan Duars ; ad. $
thai.
Mokokchung, Naga Hills ; ad. $
Mt. Popa, Upper Burma ; ad. $
Toungoo, Lower Burma ; ad. $
Toungoo, Lower Burma ; ad. <j
Toungoo, Lower Burma ; ad. <J
Habits. — The Ceylonese race of this Civet is common, according
to Phillips, all over the island, both in the hills and low country.
Although apparently purely nocturnal in the more thickly
populated districts, lying up by day in some thick clump of
fern, long grass or scrub, very rarely among rocks or boulders,
in the wilder jungle, it may be seen hunting in the daytime.
When chased by dogs, which easily follow its strong scent,
it usually tries to escape by dodging and twisting in and out
through the thickest undergrowth, sometimes going to ground,
but when driven into the open it is not swift enough to escape
its pursuers. Although reported to be a good climber, the
animal was only once seen by Phillips to take refuge in a tree.
It feeds chiefly on birds, small mammals like rats and
squirrels, on frogs, insects, and their grubs, but also on fruits
and certain roots.
It breeds at all seasons, the young, from three to 'five in
number, being born in a chamber at the end of a short burrow
usually under a rock or tree-stump or in a field-drain.
In essentials this account no doubt applies to all the subspecies
of V. indica, except possibly in the breeding at all seasons.
In the main it agrees witb Blanford's report, although according
to the latter the species is said to be " distinctly arboreal."
Phillips's statement about the. infrequency with which it
takes refuge in a tree is in agreement with the structure
of the feet and with Shortridge's report that he never saw
any kind of Civet climb a tree.
VIVBBEICULA.
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376 , viVERKiD-a;.
In Dharwar, according to Shortridge, the typical race is
plentiful, its chief food consisting probably of Gerbils (Tatera)
-and other field-rats, which exist in enormous numbers. He
found it equally plentiful in Coorg, and noted that numbers
■are kept in captivity at Kolar in Eastern Mysore for the sake
of their perfume, -which is largely used in India for flavouring
tobacco smoked by the natives. Incidentally he remarked
that this Civet, like the Toddy-Cat and Common Mongoose,
is not so destructive to domesticated poultry as the smaller
wild cats. This is not from lack of opportunity, because they
habitually live in large numbers near human dwellings.
They will kill fowls when opportunity occurs, but they prefer,
apparently, to prey on rats, lizards, and other small animals,
which are found everywhere in abundance. They are fond
of carrion, and are invariably attracted by dead animals.
The Nepalese race was reported by Hodgson to be solitary,
■even pairs being seldom seen together. It was found in forests,
■detached woods, or copses, whence it wandered freely by day,
•occasionally at least by night as well, into the open country,
feeding upon birds, eggs, snakes, frogs, and insects as well
■as upon some fruits and roots.
Subfamily PARADOXURIN^E.
Resembling the Viverrinse in the presence of well-developed
scent-glands in both sexes, but the glandular pouch represented
externally by a more widely-spread area of naked skin bordered
by tumid lips capable of being folded over to meet in the
middle line, but not so as to constitute a definite closed pouch
for the storage of the secretion. In the male also there is
■a neck of naked skin round the base of the penis, continuous
behind with the glandular area, and in the $ the glandular
area embraces the vulva, which lies near its centre*. The
feet also are adapted for climbing, being short and broad,
with the five digits evenly spaced, except the third and fourth
of the hind foot, which have their pads joined proximally,
And forming a continuously curved line, the first being approxi-
mately on the same level as the fifth and not raised as a " dew-
claw " above the plantar pad. The plantar pad is wide, com-
posed of four distinct, subequal lobes, and continuous behind
with the equally wide bilobed carpal pad on the fore foot
And with the two large metatarsal pads on the hind foot,
of which the underside is naked nearly or , quite to the heel,
the gait being subplantigrade, not digitigrade.
* For figures and descriptions of the glands and other external
characters of the genera of this group see my paper in Proe. Zool.
Soe. 1915, pp. 387-412, where references to the works of other authors
-are discussed.
rAEADOXURINJE.
377
The general shape is also different, the body being longer
and more sinuous, the legs shorter, and the tail typically
much longer.
The skull is generieally variable in form, being sometimes
more, sometimes less, modified in response to the action of the
Pig. 90.
A. Lower view of right hind foot of Paguma larvata larvata : to, internal
metatarsal pad.
B. Lower view of right fore foot of the same : a, inner half of carpal
pad.
C. Anal and genital area of 9 Paradoxurus hermaphroditus : a, anus ;
gl, shallow glandular pouch with row of secreting pores on each
side of the vulva in the centre.
D. The same of <? Paradoxurus hermaphroditus : a, anus ; gl, shallow
glandular pouch, spread open with the scrotum behind it and
the prepuce (p) in front.
'temporal muscles than in the Viverrinse. The dental formula
is typically the same as in that subfamily, but the teeth
themselves, like the shape of the skull, are generieally very
378 VIVEEBID^!.
variable. They may be similar to those of Viverra, except
that the postcanine teeth are slightly less compressed, ' the
lower carnassial (m x ) is typically a little longer than the external
length of the upper (#m 4 ), and always the last two upper cheek-
teeth (m 1 and m 2 ) are less abruptly intumed, so that the
upper carnassial is set a little more forward, not so near
the " point of maximum efficiency " of the jaw, these features
indicating that the teeth are not so specialized for flesh-
eating as in the Viverrinse.
This subfamily, ranging throughout the forested parts of the
Oriental Region, is represented in British India by three
well-defined genera distinguishable by external and cranial
characters.
Key to their Identification by External Gharacters.
a. Tail not prehensile ; hind foot -with the heel
hairy; coat not exceptionally long and
shaggy ; hairs on the backs of the ears
short.
6. A definite pattern of dorsal stripes and
lateral spots, at least in the new coat,
but not infrequently concealed by the [& Geoffr., p. 379.
long black hairs Paradoxtjbus Cuv.
b'. No pattern of stripes or spots on the body. . Pagttma Gray,
a'. Tail prehensile ; hmd foot with heel naked ; [p. 415.
coat long and shaggy, the backs of the ears
clothed with long hairs projecting beyond [p. 431.
their tips as a tuft Aectictis Tenrm.,
Key based on Cranial and Dental Gharacters.
a. Skull with the palate not produced over the
anterior half of the mesopterygoid fossa ;
its postorbital area strongly constricted,
much narrower than the interorbital and [<fc Geoffr., p. 379.
than the maxilla above the canines Pabadoxubtts Cuv.
a'. Skull with the palate produced backwards
over the fore part of the mesopterygoid
fossa ; postorbital area about the same
width as the interorbital.
b. Teeth normal, fourth premolar and first
molar above and below large, nearly as
in Paradoxurus ; muzzle larger, its width
above the upper canines about the width
of the postorbital area ; interorbital and
frontal area not noticeably inflated by (p. 415.
air-cells Paguma. Gray,
£'. Teeth, except canines, reduced, the fourth
upper premolar and first molar above and
below small ; width of maxilla above
canines much less than interorbital width ;
interorbital and frontal regions typically [p. 431.
a good deal inflated with air-cells Acxictis Temm.,
PARADOXURUS. 379
The only other genus of the Paradoxurinse is Mussehenbrok's
Palm-Civet (Macrogalidia musschenhroechii) from N. Celebes.
This large species is most nearly related to Paradoxurus,
resembling P. zeylonensis in the reversal of the hairs on the
neck and in its brown colour, with at most a faint pattern.
The skull is like that of Paradoxurus in its narrow, constricted,
postorbital area and other features due to muscular develop-
ment, and like that of Paguma in the extension of the palate
over the fore part of the mesopterygoid fossa ; but it differs
from both in having the palate parallel-sided instead of
widened behind by the divergence of the rows of the cheek-
teeth. Also the enamel of the teeth, especially of the canines,
is conspicuously pitted (see Pocock, Proc. Zool. Soc. 1933,
p. 1013).
Genus PARADOXURUS Cuvier & Geoffroy.
Pwadozturus, F. Cuvier & Geoffroy, Hist. Nat., Mamm. -at. 24
p. 5, 1821.
Platychista, Otto, Nov. Act. Acad. Caes. Leop., xvii, p. 1089, 1835.
Bondar and Macrodus, Gray, Proc. Zool. Soc. 1864, pp. 531 & 536,
Type of Paradoxurus, typus Cuv. & Geoffr. (=hermaphroditus
Schreber) ; of Platychista, pallasi Otto (=hermaphroditus,
Schreb.) ; of Bondar, bondar Desm. ; and of Macrodus,
macrodus Gray (=javanicus Horsf.).
Distribution. — Over nearly all the forested parts of the
Oriental Region from Ceylon and India, northwards to the
Himalayas, thence eastwards through Upper Burma to
S. China and Hainan, southwards through Indo-China, Siam,
Malaya, the larger and smaller Austro-Malayan islands to
Celebes, Ceram, and the Kei Islands.
Tail nearly as long as the head and body, sometimes quite
as long, and about six times as long as the hind foot. Coat
on the body consisting of long contour hairs and more or less
underwool, varying with latitude, in winter. Pattern of
dorsal stripes and lateral spots usually manifest, sometimes
very faint. Head broad, muzzle narrow, with the rhmarium
large, deeply sulcate in the middle, with prominent angles
above anteriorly ; ears large, rounded at the tip, the interior
ridges and bursa well developed, the posterior flap of the
latter rising behind the edge of the pinna, the anterior flap
deeply emarginate. Feet as described under the subfamily.
The skull exhibits marked muscular moulding as compared
with that of the other British Indian genera, notably in the
postorbital area, which is deeply constricted a short distance
behind the well-developed postorbital processes, and is con-
siderably narrower than the interorbital area and than the
-380
VrVBEBIDiB.
muzzle above the canines ; the sagittal crest is present in the
adult of both sexes, and the zygomatic arches are often strongly
salient ; the palate is not produced behind to cover the anterior
half of the mesopterygoid fossa, and is flat and expanded
between the posterior cheek-teeth. The dental formula is
Fig. 91. — Right half of palate of Paradoxwus liermaphrodiiug, showing
the upper teeth and the anterior part of the mesopterygoid fossa.
(From Blanford.)
'• normal for the Viverridse and the teeth are tolerably similar
to those of Viverra, except for the differences stated under
the subfamily heading.
The three species admitted as British Indian in this volume
may be briefly distinguished as follows : —
■a. Hairs on the neck reversed in direction,
growing forwards from the shoulders to the
head ; contour hairs of the coat typically
less long and shaggy ; pattern at most
obscure.
6. General colour from dark sepia-brown to
golden-brown ; facial vibrissse reddish ;
prepalatine foramina in skull of normal [p. 381.
length zeylonen&is (Schreb. )
£>'. General colour typically darker ; facial
vibrissas from brown to "blackish ; pre-
palatine foramina exceptionally long,
projecting beyond the level of the canines, [p. 383.
except in old skulls jerdoni Blanf.,
■a'. Hairs on the neck directed backwards ; con-
tour hairs typically long and shaggy ;
pattern better defined ; facial vibrissse
black; prepalatine foramina of normal (Schreb.), p. 387.
length hermaphroditus
PARADOXURUS. 381
43. Paradoxurus zeylonensis (Schreber). The Golden Palm-
Civet.
Viverra zeylonensis (Pallas), Schreber, Saug. iii, p. 451, 1777.
Viverra zeylanica, Gmelin, Syst. Nat. ed. 13, i, p. 89, 1788.
Paradoxurus zeylanicus, with var. fuscus or montanus, Kelaart,
Prodr. Faun. Zeylon. pp. 39-40, 1852.
? Paradoxurus aureus, F. Cuvier, Mem. Mus. Hist. Nat. Paris,
p. 48, pi. 4, 1822 ; Blanford, Fauna of Brit. Ind., Mamm. p. 110,
1888 ; and of most subsequent authors.
Paradoxurus zeylonensis* Poeock, Journ. Bomb. Nat. Hist. Soo.
xxxvi, p. 859, 1933*; Phillips, Man. Mamm. Ceylon, p. 174,
pi. 13, 1935.
Vernacular. — Kalawedda (Sinhalese) ; Marum nai (Tamil).
Locality of the type of zeylonensis (=zeylanicus), Ceylon ;
of fuscus or montanus, Newera Eliya, Ceylon ; of aureus*
unknown.
Distribution. — Ceylon.
Hair in front of the shoulders radiating from two whorls
and growing forwards along the nape and sides of the neek
to the head ; on the fore throat also it grows forwards, radiating
from a single whorl. General colour of upper side brown,
but individually variable from dark sepia to ochreous, rusty
or golden-brown, the tips of the contour hairs frequently
lustrous, sometimes greyish ; legs about the same tint as the
back, but the tail and face sometimes noticeably paler, buffy-
grey, the face without grey pattern, but the vibrissse dirty
white. The dorsal pattern characteristic of the typical species
of Paradoxurus absent or represented by faint bands and spots
slightly darker than the ground-colour. The lower side slightly
paler and sometimes greyer than the upper, especially,
apparently, in younger specimens, in which grey may be
dominant over brown even on the back.
There is not sufficient material to establish the existence
of possible local races ; but the following variations may
be noted. Two examples from Nuwara Eliya, collected by
Kelaart and named by him fuscus (montanus), are decidedly
paler on the head and face than others from different localities,
but the type is darker, more rusty brown than the other and
has no pattern, the topotype being more golden-brown with
* Note on the synonymy. — The original specimen of this species was
from Ceylon, as recorded by Pallas, who sent a description of it to
Schreber for publication. G-melin epitomized this description, but
altered the name to zeylanica,, and this inadmissible emendation was
adopted by Kelaart and Blyth. In 1885 Blanford (Proe. Zool. Soc.
1885, p. 612) quoted the species as Paradoxurus zeylonensis ; but in
his volume in the ' Fauna of British India, Mamm.,' he abandoned that
name in favour of aureus given by Cuvier to a young example of Para-
doxurus from an unknown locality. In this he was followed by sub-
sequent writers. In my paper quoted above I gave my reasons, which,
need not be here repeated, for thinking that Blanford's original opinion
on this point was correct.
382 vivbrridje.
faint pattern. In two examples from Mousakanda, Gamma-
duwa, C.P. (W- W. A. Phillips), one, a subadult <$, is rather
darker than the type of fuscus, the hairs of the back having
very dark brown tips, especially on the eroup, and there is
a dark brown band down the upper side of the tail, of which
the underside is grey ; the other, quite immature, is much
greyer than other known skins, grey with a brownish tinge
above and quite grey below. These two specimens bear
out Blyth's statement that two living examples sent to him
by Kelaart gradually darkened with age. An adult $ from
Maha Oya, E.P., is dull greyish-brown above, with no bright
tint ; the tip of the tail is white, and there is a white patch on
the belly. A £ from Koslanda, S. Ceylon, intermediate in tint
between the darker and paler brown specimens, is remarkable
for the very pale hue of the tail, which becomes gradually
white from the base to the tip.
The coat varies from about 30 to over 40 mm. It is longest,
namely, 43 mm., in the undated type of fuscus (montcmus)
from Nuwara Eliya, and shortest, namely, 29 and 27 mm.
respectively, in a skin from Gammaduwa, 3,000 ft., C.P.,
dated September, and one from Maha Oya, E.P., dated
August. No doubt the length varies both seasonally and with
altitude.
The flesh-measurements (in English inches) and weights
(in lb.) of some specimens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
lb. oz.
Phillips's largest ; ad. <J 22f 19f 3£ 7 3
Phillips's average of 7 ; ad. <J 20£ 18 3 6 4
Gammaduwa (Phillips's, in.
Brit. Mas.) ; ad. <J 20f 17f 2$ 5 8
Phillips's largest ; ad. $ 20 17$ 3— 5
Phillips's average of 5; ad. $. 18 i 16| 2£ 3 3
MTaha Oya (Mayor, in Brit.
Mus.);ad. $ 20+ 18 2£ 4
Habits. — This species, according to Phillips, although not
uncommon, is locally distributed in the hills and low country,
but seems to be commoner in the hills round Kandy and in
the Dambulla and Dikoya districts of the Central Province
than elsewhere. It has been recorded from Colombo, but is
not at all common there. It is very rare at Kalutara in the
Western Province, but has been observed at Ratnapura and
several places in the North Central and Eastern Provinces.
In habits it is nocturnal and essentially arboreal, usually
spending the day in the hollow branches of large jungle-trees ;
not infrequently it occupies the roofs of bungalows adjoining
the jungle. It feeds largely on seeds and fruits such as
plantains, guavas, pineapples, and mangoes, but is apparently
PABABOXUKT7S. 383
by preference a flesh-eater, devouring any small mammals,
birds, snakes, lizards, frogs, moths, and other insects it can
catch. The young, generally two or three in number, are
born mostly, apparently, in the later months of the year,
October and November. They are dull mouse-grey or grey-
brown in colour, occasionally snowing a faint pattern of dorsal
stripes.
44. Paradoxurus jerdoni Blanford. Jerdon's Palm-Civet.
Distribution. — Sotxthebn India, the Palni and Nilgiri
Hills, Travaneore, and Coorg.
Resembling P. zeyhnensis in the reversal in growth of the
hairs of the neck and throat, but larger and, although sometimes
very similar in colour, usually considerably darker, either
deep brown all over or brown or black speckled with silvery
or buffy-grey on the back or flanks, and the pattern only
occasionally just traceable on the back ; the facial vibrissse
are blackish,' not rufescent as in zeyhnensis. The skull, larger
and with larger teeth than in zeyhnensis, is distinguished
from that of all the other species of Paradoxurus by the
exceptional length of the prepalatine foramina, which extend
backwards beyond the line of the posterior edges of the
upper canines, although becoming shorter in old skulls.
This southern Indian species appears to be represented
by the following two local races.
44 a. Paradoxurus jerdoni jerdoni Blanford.
Paradoxurus jerdoni, Blanford, Proc. Zool. Soc. 1885, pp. 613 &
802, pi. 49 ; also 1886, p. 420 ; and of subsequent authors,
at least in part.
Paradoxurus jerdoni jerdoni, Pooook, Journ. Bomb. Nat. Hist.
Soc. xxxvi, p. 863, 1933.
Locality of the type, Kodaikanal in the Palni Hills.
Distribution. — Palni Hills, Nilgiri Hills, and Travaneore.
Face uniformly coloured or with faint traces of grey speckling,
but without definite grey pattern ; the body on the average
less conspicuously speckled with grey-tipped or buff-tipped
hairs.
Skins assigned to this race show great individual variation
in colour. The coat in the type is mostly destroyed by moth ;
but three, to all intents and purposes topotypical examples,
collected in the Palni Hills by Mr. C. McCann, indicate the
coloration of the race. An adult $ from the Pamber River,
Kodaikanal, 7,000 ft., May 16, has the coat thick, 42 mm. long,
and the general tint uniformly dark glossy brown all over,
with some faint buffy speckling, whereas two adult (J specimens
from Tiger Shola, 5,600-5,700 ft., April 22 and 27, have the
384
VIVBEEri>^B.
PAEA.DOXURUS.
385-
coat 41 and 43 mm. and the tint blackish-brown, with the
back behind the shoulders, the flanks, and belly speckled with
clear grey, and some grey in front of the ears. One is more
grizzled than the other, has a white tail-tip, and some yellow
at the base of the tail. The difference between the uniformly
brown $ and the grizzled $ specimens is striking ; but
the difference is probably individual, not sexual or seasonal.
The example figured by Blanford, which probably came from
the Anaimalai Hills, is like the Tiger Shola skins, but has the
$&.
Fig. 92.
A. Anterior part of bony palate of very old specimen of Paradoxwus
jerdoni, showing the shortening of the prepalatine foramina with
age.
B. Thesameofyoungerspecimenfrom Tiger Shola, showing the normal,
exceptionally large prepalatine foramina.
C. & D. Upper and lower views of exceptionally well-developed <J skull
of P. h. hermaphroditus from Bellary. All figures X J.
dorsal pattern faintly indicated. An unsexed specimen
from Wellington in the Nilgiri Hills, December 11, has the very
full coat about 40 mm. long and the tint uniformly deep brown,
like the skin from the Pamber River, but somewhat richer.
vol. i. '2c
386 VOTEKJ&rDiE.
On the other hand, an old # from Kateri in the Nilgiri Hills
has the back and flanks distinctly speckled like the <J skins
from the Palni Hills, although the speckling is more buffy-
grey. Another <J, from the northern end of the Travancore
range, January, has the coat only 35 mm., and resembles
the skin from Kateri in colour except that the speckling is
noticeably less conspicuous, hut a $, undated, skin from
Trivandrum differs from the foregoing in being dominantly
buffy-grey owing to the more extensive and profuse paleness
of the hair-tips.
A series of five skins of various ages from Lovedale,
Ootacamund, in the Nilgiri Hills, 7,300 ft. (Phythian Adams),
agrees with those previously described, some being uniformly
brown on the back, others speckled with buffy-grey to a
varying extent.
Conceivably more than one subspecies is represented by
the skins above described. This can only be decided by
additional material.
The flesh-measurements (in English inches) and weights
(in lb.) of the specimens collected by McCann in the Palni Hills
.are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Tiger Shola ; ad. <? 23J 21 3| 8
Tiger Shola ; subad. <J 22£ 19£ 3§ 6
JXodaikanal; ad. $ 21f 17f 3+ —
44 b. Paradoxurus jerdoni caniscus Pocock.
Paradoxurus jerdoni caniscus, Pocock, Journ. Bomb. Nat. Hist.
Soc. xxxvi, pp 803-4, 1933
Locality of the type, Virajpet in S. Coorg, 3,000 ft.
Distribution. — Nqbth and South Cooeg.
Distinguished from the typical race by havings the grey
■and black facial pattern, or mask, well defined, as well as in
some Indian examples of the next species, hermaphroditus,
and consisting of a large grey patch in front of the ear, and
•one below and one above the eye set off by black intervening
areas, the back, flanks, belly, and tail being also more profusely
speckled with grey-tipped hairs.
There is a certain amount of individual variation in the
"three known skins. Two, adult o* and $, from the typical
locality (Shortridge), collected January 13, are alike except
■than the $ has the hairs on the basal portion of the tail paler,
dirty white. The coat is about 35 mm. in both. A young
<J from Haleri, N. Coorg, February 28, has the coat a trifle
longer, 37 mm., and considerably more white on the body,
the tail being mainly white, with a black longitudinal dorsal
stripe and a black tip.
PABADOXUKUS. 387
The flesh-measurements (in English inches) and the weights
(in lb.) of the type and topotype are as follows : —
Head and Hind
Locality and sex body. Tail. foot. Weight.
Virajpet ; ad. <J 24 21 3f 9£
Virajpet ; ad. ? 22* 21 3f 5J
Habits.— According to Shortridge this race is fairly plentiful
in Coorg, although not nearly so abundant as the common
Toddy-Cat. The two species are found side by side in the same
localities, especially round coffee estates, and are not distin-
guished by the natives. Their habits are nearly identical, both
being largely arboreal and to be seen in trees on moonlight
nights ; but this race of Jerdon'sPalm-Civetwas not observed
to frequent the roofs of bungalows like the Toddy-Cat.
45. Paradoxurus hermaphroditus Schreber. The Common
Indian and Burmese Palm-Civets, " Toddy-Cat " of
Anglo-Indians ; Polecat and Black Palm-Civet in
Ceylon.
(Essential bibliographical references are entered tinder the sub-
species. )
Vernacular*. — TJgguduwa, Kalawedda (Sinhalese), Marum
nai (Tamil) ; Mara naie (Jaffna Tamil) in Ceylon ; Ud
(Mahratta) ; Kullibekku (Coorg), Manupilli, Punugupilli
(Telegu) ; Punaginabekku, Kerabekku or Kerabek (Kanarese) ;
Nulla philli (Waddani) ; Mahngutchi (Haran Shikaris in
Dharwar) ; Togot (Singbhum), Khotas, Menuri, Lakdti,
Ghangar, Jhar-ka-Kutta (Hindi) ; Bhondar, Baghdankh, Bham
(Bengali) ; Kyowng-na-ga, Ky owing -won-baik or Kyaung-wwm-
paik (Burmese) ; Tok-Toh, Sapo-mi-aing (Karen) ; Khubbo-
palaing (Talain) ; Musang or Musang Pandan (Malay).
Distribution. — The same as for the genus.
Distinguished from P. zeylonensis and P. jerdoni by the
backward slope of the hairs of the neck and from jerdoni
also by the shorter prepalatine foramina of the skull.
The pattern, too, is, as a rule, better defined than in those
species, especially in the short, new coat, where it consists of
longitudinal stripes on the back, spots on the sides, shoulders,
and thighs, and sometimes on the base of the tail. The head
also typically shows a definite pattern, the " mask," composed
of white patches on a black ground, the fundamental plan
being a patch on each side of the muzzle, one, the suboeular,
* Under this species I have made no attempt to allocate the recorded
vernacular names to the different subspecies. In some cases there
is no doubt to which they apply ; but in others the same name may be
given to two or more races.
2 02
388 VrVERBIDJE.
below the eye, and one, the superciliary, above it ; a larger,
preaural patch, in front of the ear, and a frontal band sweeping
across the forehead ; this " mask," however, is very variable
individually and racially, the white patches being sometimes
nearly obliterated by the extension over them of the black
intervening spaces, sometimes enlarged, and to a great extent
confluent, so that the head may be dominantly black or
largely white. The pattern on the body may also be obscured
by the black pigmentation of the long contour hairs concealing
the underlying under-hair, which is more or less woolly.
Legs black. Tail also black, at least at the end, generally
paler at the base, and sometimes faintly striped.
This species, in accordance with its extensive distribution,
is represented by a very large number of local races dis-
tinguished by size, coloration, and cranial and dental characters.
Individual variations, seasonal or otherwise, are also generally
well marked.
Many local races are now distinguished in the British Indian
fauna. These were assigned by Blanford to two species,
P. niger, occurring in Ceylon and India, from the Himalayas
southwards, and P. hermaphroditus, found to the east of the
Bay of Bengal. He admitted, however, the existence of
intergrading forms, and, in accordance with his principle of
ignoring local races or subspecies, should have followed
Blyth and Jerdon in uniting the two as representing a single
species. There also seems to be no reason to doubt that the
name hermaphroditus was originally given to a South Indian
specimen, and not to a specimen from Malaya or Sumatra.
. The marked individual variation in colour and pattern
in specimens from Peninsular India resulted in the ascription
of a large number of spurious names to these Palm-Civets,
especially by Gray and Hodgson. The individual synonymy
was fully discussed in my papers (Joum. Bomb. Nat. Hist.
Soc. 1933-4), and my reasons for the nomenclature here
adopted need, not be repeated. All that need be said is
that the races here admitted as occurring in Ceylon and
Hindustan from the Himalayas southwards are indistinguishable
by colour and pattern when the contour hairs are moulted
and the new coat, showing a definite pattern of stripes and
spots, is growing.
45 a. Paradoxurus hermaphroditus hermaphroditus (Schreber).
Viverra hermaphrodite/, (Pallas), Schreber, Saug. iii, p. 426, 1778.
Paradoxurus hermaphroditus, Gray, Proc. Zool. Soe. 1864, p. 532
(not of Blanford, 1885 and 1888, and the authors who followed
him).
Viverra nigra, Desmarest, Mamm. p. 208, 1820 (not V. nigra Peale-
& Beauvois, 1796.)
PARADOXURUS. 389
Paradoxurus niger, Blanford, Proc. Zool. Soe. 1885, p. 792 ; id.,
Mamm. Brit. Ind. p. 106, 1888 (in part) ; Wroughton, Journ.
Bomb. Nat. Hist. Soc. xxv, pp. 48-51, 1917.
Paradoxurus typus, F. Cuvier & Geoffiroy St. Hilaire, Hist. Nat.
Mamm., pt. 24, p. 5, pi. 186, 1821.
Paradoxurus hermaphroditus typus, Robinson & Kloss, Reo. Ind.
Mus. xix, p. 178, 1920.
Paradoxurus typus var.fuliginosus, Gray, Proc. Zool. Soc. 1832,
p. 65.
Paradoxurus hermaphroditus hermaphroditus, Pocock, Journ.
Bomb. Nat. Hist. Soe. xxxvi, p. 867, test-fig. 1, A, B (heads)
and 2, C, D (skull), 1933 ; Phillips, Man. Mamm. Ceylon,
p. 171, pi. xii, 1935.
Locality of the type of hermaphroditus unknown, recorded
from " Bombay " ; of nigra (niger) and typus, Pondicherry :
of fuliginosus, S. India.
Distribution. — Ceylon and Southern India as far north
as the Narbada River.
Distinguished when in full coat by the combination of com-
paratively short and thin under wool with long, extensively
blackened contour hairs almost entirely concealing the
underlying pale tint, which may vary in skins from the same
locality from whitish to buff or rich ochreous, but when the
contour hairs are shed or thinned by the moult the ground-
colour is revealed wholly or in part, and with it the distinct
pattern of black stripes and spots. The " mask " on the
short-haired head does not change appreciably with the
season ; it varies very considerably individually in the relative
extent of the black and white.
The long-coated dark phase, agreeing closely with the
description of hermaphroditus, in which the pattern is often
obscured and the pale hue of the under-hair shows as streaks
or mottling when the contour hairs are parted, the phase
described by Gray as a distinct variety, fuliginosus, is repre-
sented by an undated skin from Ceylon, and by many skins
from Coorg, Kanara, Dharwar, Ratnagiri, and Satara in
S.W. India, collected from November to February, when
the coat usually shows no sign of moulting. But in one
from S. Coorg, dated January, most of the contour hairs are
shed from the rump and hind back, revealing the grey under-
hair, which shows pronounced pattern. Specimens collected
at the end of April and early in May in the North Central
and Southern Provinces of Ceylon show moult of the contour
hairs in various stages. One of these from TJdugama, S.P.,
differs greatly from the Ceylonese representative of the
fuliginosus-type above referred to not only in the less abundance
of the contour hairs, but in the ground-colour being a rich,
nearly ochreous-buff instead of grey, and the conspicuous
whiteness of the mask, which in the other is largely obliterated
by black. These two skins are illustrations of the extensive
390 VrVEBBID^l.
individual differences which may occur in the race. Another
series of skins collected from August to October in the Eastern
Ghats, i. e., the Dharmapuri and Denkanikota Ranges,
N. Salem, and the Palkonda Hills in S. Cuddapah, is very similar
in coat and general coloration to those collected in Ceylon
in April and May, and similarly differs from the majority of
skins from the Western Ghats collected from November to
February.
The specimens from the Eastern Ghats, occurring up to
3,000 ft. or more, as well as one from Madras, which closely
resembles the skin from Udugama in Ceylon, are of interest
from the proximity of their localities to Pondicherry, whence
the specimens described respectively by Desmarest and
Cuvier as niger and typus were shipped.
A few skins from the district of Hoshangabad, collected on
March 5 and 21, at 2,500 and 3,300 ft., are in full coat, and in
their black, marbled with grey, coloration are very like the
skins from Satara, whereas one dated April 1, 1,000 ft., is
thinner-coated, showing the beginning of the moult. Like
the skins from Satara, they are somewhat fuller in the coat
than those found farther south, thus intergrading in that
respect with the race from Gwalior (p. 393).
The following table shows the flesh-measurements (in
English inches) and the weights (in lb.) of some of the largest
specimens : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Satara ; <J 22 \ 24-} 3 10
Hoshangabad ; <? 2H 21§ 3J 7§
Palkonda Hills ; <J 22§ 2lf 3* 6
Ceylon, N.C.P. ; <J 2 If 18* 3j 8
Satara; $ . .• 2lJ 19£ 2§ 8
Dharwar ; ? 22$ 23 3& —
Palkonda Hills ; $ 20$ 18 3J 6
Ceylon, S.P. ; $ 23$ 17$ 3* —
Phillips's measurements and weights of Ceylonese examples
are approximately as follows : —
Head and Hind
body. Tail. foot. Weight.
Largest ad. <$ 22$ 18* 3£ 7£
Average of 10 ad. tJ(J 20J 17° 3+ 6£
Largest ad. 9 20J 18$ 3* 6
Average of 4 ad. $°. 18J 17 3" 5
These data agree tolerably closely with those recorded
in my table.
Sktdl.— In adult and old £ skulls the sagittal crest is
well developed, reaching 6 mm. in height in an oldish well-
developed skull from Bellary, and the postorbital processes
MAMMALIA.
PLATE XXVIII.
Photo W. S. Berndge.
Indian Palm-Civet (Paradoxurus hermaphroditus).
A & B. Heads of two examples of Paradoxurus hermaphroditus hermaphroditus,
showing variation in pattern. C. Head of Burmese Palm-Civet (Paradoxurus
hermaphroditus laotum) from Mingun. D. Head of Lesser Palm-Civet (Para-
doxurus hermaphroditus minor) from Tenasserim.
PAKADOXtTBTJS.
391
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392 VTVEBBIDJ3.
are triangular, pointed, and have their posterior edges
approximately in the same transverse line. In adult and
old $ skulls, which are on the average a little shorter than
£ skulls and narrower across the zygomata, the sagittal
crest is complete but low, about 1 mm. high, and is late
in forming, the two temporal ridges in a young adult $ skull
from Dharwar, with complete dentition, being 5 mm. apart
■on the frontals and 10 mm. on the parietals. As shown
in the subjoined table, the skulls vary individually in their
principal dimensions as well as in the size of the bullae and
teeth.
45 6. Paradoxurus hermaphroditus nictitatans Taylor.
i. Paradozurus leueopua, Ogilby, Zool. Joum. iv, p. 300, 1828.
Paradoxurus nietitatans, Taylor, Joum. Bomb. Nat. Hist. Soo.
vi, p. 429, 1891.
Paradoxurus hermaphroditvs nictitans, Pocock, Journ. Bomb. Nat.
Hist. Soo., xxxvii, p. 172, text-fig. 3, A, B (skull), 1934.
Locality of the type, Kondmals, Orissa Division of S. Bengal.
Distribution. — Lowbe Bengal.
Distinguished, on the evidence of a few specimens, from
the preceeding race by its smaller size and possibly average
difference in colour, being greyer, less black, and with the
pattern superficial.
The examples of this Palm-Civet recorded and collected
by Taylor, as well as one or two additional specimens, are
only provisionally admitted as representing a distinct race
pending the collection of more material than is at present
available. Taylor considered the partial albinism of his
type, of which the mother was also similarly coloured, as
a specific feature. The type has only the head and shoulders
normally coloured, the rest being white. Subsequently he
procured two other specimens with the white less extensive,
appearing as a broad belt encircling the hind body and extending
over the ventral surface, the feet and the terminal third of
the tail being also white. These specimens came from
Kondmals in Orissa. Where the coat is pigmented the
coloration is very similar in the three specimens, including
the type, in the British Museum, the ground-tint being clear
grey or slightly tinged with buff and covered to a varying
extent by long black-tipped contour hairs, but black is very
little in evidence on the grey tail. The pattern is indefinite
and superficial, being formed by the confluence of the black
tips of the contour hairs. The coat has rather more under wool
on the average than in typical hermaphroditus.
A just adult $ example collected at Hazaribagh (Crump) in
June, with its long coat still unmoulted, probably represents
PAEADOXTIBTTS.
393
the normally coloured phase of this race. The coat is long
and shaggy, the under-hair is grey, and the black-tipped
contour hairs, 57 mm. long, are tolerably abundant, but there
is no definite pattern. Its measurements are as follows : —
Head and body 1S§ in., tail 19J in., hind foot 3 in., and its
weight was 3 lb. It is thus considerably smaller than adult
2 examples of typical hermaphrodite. Its skull also is smaller
than that of the latter race, as indicated in the table (p. 391) ;
Fig. 93.
A. Skull of adult t? of Paradoxurus hermaphroditus nictitatans from
Khondmals, Orissa.
B. Skull of adult $ identified as Paradoxurus hermaphroditus nictitatans
from. Hazaribagh. Both figures X J.
and the skull of a $, just adult, with complete but low sagittal
crest, from Khondmals, is also smaller than average $ skulls of
typical hermaphroditus, resembling 2 skulls of that race.
45 c. Paradoxurus hermaphroditus scindise Pocock.
Paradoxurus hermaphroditus scindise, Poeock, Journ. Bomb. Nat.
Hist. Soc. xxxvii, p. 176, 1934.
Locality of the type, Guna in Gwalior, about 40 miles
north of lat. 24°.
Distribution. — Gwauob, Bhopal, Centbal India, and
probably Rajptttana, i. e., the country drained by the
394 VIVEEBIDJE.
Chambal and its tributaries flowing north-east into the
Ganges.
Distinguished from typical herrnaphroditm of Southern
India and Ceylon by its slightly larger size and, in winter,
by its longer, more luxuriant woolly coat, which is not con-
cealed to the same extent by the long black-tipped contour'
hairs. The pale ground-colour varies from darker or lighter
grey to buffy-grey, which, combined with the blackness of
the contour hairs, yields a marbled or blotchy pattern.
Skins referred to this race were collected between October
and December at Guna, Binganj, Cachora Fort, and Agar at
altitudes varying from 1,375 to 1,680 ft. in Gwalior and at
Neemuch (Nimach), also in Gwalior, in March and April,
at 1,400 and 1,500 ft. One of the specimens from Neemuch
is remarkable for the tawny hue of the under-hair and the
brown hue of the tips of the contour hairs, which are probably
faded previous to the moult. There is also a skin from
Sehore in Bhopal, April 29, which, like the tawny Neemuch
skin, has the coat thinnish and shaggy, but the ground-colour
is grey. Skins from Samblar, Bajputana, September 15, in
poor coat, also seem to belong to this race.
The difference in coat between this race and hermaphroditus
is as follows : — In scindim the approximate average length
of the wool between December and April is 45 mm., of the
contour hairs 60 mm, the corresponding measurements
for the same months in hermaphroditus being 30 and 53 mm.
The flesh-measurements (in English inches) of some examples,
of scindise are : —
Head and Hind
Locality and sex. body. Tail. foot.
Guna ; yg ad. cj 26£ 22J 3J-
Binganj ; ad. ? 26 24 3J
Cachora Fort ; ad. o 24$ 22 3J
Agar-Malwa ; yg. ad. $ 24$ 20| 3
Bhopal; $ 23£ 21£ 3£
Sambhar ; yg. ad. ^ 24£ 23 3|
The two (J specimens, namely, the type from Guna and the-
one from Sambhar, although not fully adult, are bigger than
the largest <J specimens of hermaphroditus, and the average
dimensions of $ specimens of scindise. are : head and body
24| in., tail 22 in.; of hermaphroditus, 21 J in. and 20 in.
respectively.
45 d. Paradoxurus hermaphroditus laneus Pocock.
Paradoxurus hermaphroditus laneus, Pocock, Journ. Bo
Hist. Soc. xxxrii, p. 178, fig. 4, B, 1934.
Locality of the type, Gopalpur, 5,200 ft. in Kangra.
PABADOXUBUS. 395
Distribution. — Kangra in the Upper Punjab from 2,000 to
7,000 ft.
Closely resembling scindias from Gwalior in size and general
appearance, but -with, the -winter coat fuller, more woolly,
perceptibly more resistant to the touch and keeping the
contour hairs, which are less extensively black terminally,
even more erect than in that race ; the sides of the neck
are also somewhat blacker.
Several examples collected in March at Gopalpur, 5,200 and
7,000 ft., and in the Kangra Valley, 2,000 to 2,500 ft. (Wells).
The series shows stages of the coat-change, the type from
Gopalpur, March 6, being in full winter coat, whereas one from
the Kangra Valley, March 28, is in full moult. The ground-
colour varies from grey to buffy-grey and, combined with the
black of the more or less upstanding contour hairs, produces
in full coat a marbled or mottled pattern of lines and
spots. In the type the white of the mask is a good deal
reduced.
In the $ in full moult a large number of the contour hairs
are shed and the neck, shoulders, and fore-back are covered
with a short, close coat of soft greyish-white new hair showing
a very distinct pattern of black stripes and spots. In its
coloration this skin is not distinguishable from skins of
scindias or hermophroditus in similar coat-change.
A young dark grey specimen showing the pattern was
collected by H. Whistler in September at Bharmsala, 4,000 ft.
In four examples in good coat the wool is about 40 mm.
and the contour hairs about 60 mm. long, the wool being
a little shorter, although thicker, than in scindias.
Flesh-measurements (in English inches) of some quite or
nearly full-sized specimens are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Gopalpur ; ad. (J 27§- 24i 3f
Kangra Valley ; yg. ad. <J 26* 25° 3f
Kangra Valley ; ad. <J 23£ 22$ 3£
Kangra Valley ; ad. $ 25J 24 31
Kangra Valley ; ad. $ 26| 25£ 3|
There is very little difference between the sexes in size.
The average of the two $ specimens, head and body 26 in.
and tail about 25 in., is a little greater than in scindias.
The skull is a little longer than in scindias and hermaphroditus
and has slightly larger teeth. That of the type is exceptionally
well developed muscularly, the sagittal crest, in the middle
of its length being 7£ mm. high, and the width across the
postorbital processes from tip to tip is 40 mm.
396
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397
45 e. Paradoxurus hermaphroditus vellerosus Pocock.
Paradoxurus hermaphroditus vellerosus, Pocock, Journ. Bomb.
Nat. Hist. Soe. xxxvii, p. 181, text-fig. 4, A (skull), 1934.
Locality of the type, Kashmir.
Distribution. — Kashmir, on the evidence of the only known
specimen.
Resembling the type of laneus in colour and pattern ,
but distinguished from that race and from scindiee by its
longer woolly coat and longer skull.
The undated, unmeasured type, the only described specimen.
Fig. 94.
A. Skull of subadult <J of Paradoxurus hermaphroditus vellerosus from
Kashmir.
B. Skull of fully adult <J of P. h. laneus from Kangra. Both figures X |.
of this race, is evidently in full winter coat, of which the
wool is about 50 mm. long and the contour hairs 60 mm.
Although the skull has the basioccipital suture still open, and
probably had not attained its full length, it is decidedly
longer than the skull of the type of laneus, as the table of
measurements shows. On account of its comparative youth
it is less well developed. The zygomatic width is less, the
398 -viVEBErD-ffi.
sagittal crest is lower, only 4J mm. high, the postorbital
processes are smaller, only 34 mm. from tip to tip, and the
cranium is narrower above the posterior root of the zygomatic
arches. No doubt with increased age the skull would have
surpassed that of lemeus in these respects as well as in length.
Probably the examples from Kashmir recorded by Col. Ward
as P. niger (Journ. Bomb. Nat. Hist. Soc. xxix, p. 10, 1927)
were representatives of this race. It is found, he says, in the
outer ranges, and is arboreal and very nocturnal, sleeping
throughout the day. But the measurements he cited — head
and body 23 in., tail 20 in., and skull 100 mm. — apply to
a much smaller animal than the type of velkrosus, which,
judging from its skull, must have exceeded the largest example
of the Kangra race, laneus. There is, however, no evidence
that the example Ward measured was mature. He also referred
to a specimen from Simla with the head and body 24 in. long,
which is racially unidentifiable, but may have been an im-
mature example of the Kangra race.
Under the heading of Viverra zibetha I suggested above that
Col. Ward's record of that species from Kashmir may possibly
have been due to confusion with the large Kashmir Toddy-Cat
(vellerosus). He cited Nil Biral as its native name and said
that it often lives under thatched roofs. This is a very
unlikely habitat for V. zibetha to choose, but agrees precisely
with records of the Toddy-Cat.
45/. Paradoxurus hermaphroditus bondar (Desmarest).
Viverra bondar, Desmarest, Mamm. p. 210, 1820.
Paradoxurus bondar, Gray, Illustr. Lad. Zool. pi. 12, 1833 ; id.,
Proc. Zool. Soc. 1864,p.531,andCat.Carn.B. M. p. 18 {bondar)
Paradooourus pennantii, Gray, Proc. Zool. Soc. 1832, p. 66 ;
id., Illustr. Ind. Zool. pi. 13, 1833.
Paradoxurus arossi, Gray, Proc. Zool. Soc. 1932, p. 67 ; id., Illustr.
Ind. Zool. pi. 7, 1833.
Paradoxurus hirsutus, Hodgson, Asiat. Bes. xix, p. 72, 1836.
Paradoxurus strictus (Hodgson MS.), Horsfxeld, Ann. Mag. Nat.
Hist. (2) xvi, p. 105, 1855 ; id., Proc. Zool. Soc. 1856, p. 396,
pi. 47.
Paradoxurus hermaphroditus bondar, Pocock, Journ. Bomb. Nat.
Hist. Soc. xxxvii, p. 182, 1934*.
Locality oitype of the bondar, Bengal ; of pennantii, Higher
Province of Bengal ; of crossi, India ; of hirsutus and strictus,
Nepal Tarai.
* No two authors out of the many who have attempted to straighten
the involved synonymy of this and the next race of P. hermaphroditus,
both occurring to the north of the Ganges, have reached the same result.
The confusion is due partly to several of the names having been given
to drawings of specimens or to specimens while alive in menageries
JPABADOXUBTIS. 399
Distribution. — Rohilkand and, according to Hodgson,
" the open parts of the Nipalese Tarai and generally in British
districts on the left (north) side of the Ganges to Northern
Bihar."
Closely resembling scindiee and laneus in the length and
luxuriance of the winter coat, the wool being long and thick,
with the black-tipped contour hairs upstanding, but the
pattern is less pronounced, sometimes evanescent, and, when
indicated, formed by the confluence of the black tips of the
contour hairs and therefore readily disarranged ; the ground-
tint of the under-coat varies from nearly white to " yellow "
or tawny. On the evidence of a few skulls and measured skins
bondar is also decidedly smaller than scindise and laneus.
The most western known examples assigned to this race
are two — one an old $, the other unsexed — collected on March 8
in full winter coat at Pilibhit, 800 ft., in Rohilkand (Crump).
The luxuriant woolly hair, from 35 to 37 mm. long, is mostly
white or silvery grey ; this is sparsely oversprinkled with the
black tips of the upstanding contour hairs, from 50 to 55 mm.
long, which at most form irregular elongated patches readily
broken up by the parting of the hair-tips. Except that the
ground-colour and brow-band are white instead of yellow,
these skins closely resemble Hodgson's description of his
hirsutus and Gray's figure of crossi taken from a living mena-
gerie specimen. Both the wool and the contour hairs of the
type of hirsutus were approximately the same as in these
Rohilkand specimens.
from vague localities, partly to discrepancies' between alleged type-
specimens and the original descriptions. My reasons for the opinions
here adopted were given in full in my paper in 1934. All that need be
repeated in connection with this race is that the original description of
bondar was taken from a copy of a sketch of a specimen from ' ' Bengal ' '
in which the coat was apparently very thick, the colour tawny, with the
long hairs black at the tip and the pattern consisting of three dorsal
stripes. As regards pennantii, based upon a coloured sketch of a speci-
men from the higher Province of Bengal, Gray himself (Proc. Zool. Soc.
1864, p. 531) dropped the name as a synonym of bondar. According to
Gray's description and figure of crossi, taken from a specimen
from " India " living in the Surrey Zoological Gardens, the coat was
long and luxuriantly woolly and yellowish or buff in colour, but with
no pattern. This agrees very closely with Hodgson's description of
hirsutus, from the Nepal Tarai, which had the coat very long, full, and
erect, the colour clear yellow, but no pattern. Hodgson himself thought
this a n imal to be the same as bondar. The type of strictus, on the
other hand, was distinguished from hirsutus on account of the distinctness
of the pattern of stripes and spots. But the type-skin is in full moult
and the pattern is shown on the newly erupted coat, and Hodgson was
■apparently unaware of the profound effect of coat-change on the colour
and pattern. I consider strictus to be the same as hirsutus because
both came from the Nepal Tarai, but it may represent palktsii.
400 VIVEBBIDiE.
Some flesh-measurements (in English inches) and weights
(in lb.) are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Nepal Tarai ;? ad. <J 23 22 — 6
Darbhanga ; yg. ad. <J 20 18 3f 8
Rohilkand ; old $ 20 18 3- 4
Darbhanga ; yg. ad. $ 18 19 3+ —
The number of available skulls is too small for profitable
comparison with the skulls of other races. None of Hodgson's
skins has a skull that can be definitely assigned to it. Measure-
ments of these skulls are entered on the table (p. 404).
The race from Lower Bengal recorded above as nictitatans
(p. 392) unmistakably resembles bondar in many particulars,
and the two may prove indistinguishable. But the skulls
of the young adult $ of bondar from Darbhanga and of the °
from Rohilkand are respectively larger than the just adult <j
skull of nictitatans from Khondmals and the adult $ skull from
Hazaribagh.
In. the table of skull-measurements I have provisionally
entered as belonging to this race all the available skulls from
Nepal belonging to Hodgson's collection, although only one
of them, the first on the list, has a skin on the whole more like
bondar in colour and pattern than pallasii. But the identifica-
tion of these skulls is doubtful. Except the one in question,
they are less well developed in the short postorbital processes
than the skulls of pallasii, but they are barely adult.
45 g. Paradoxurus he"rmaphroditus pallasii Gray.
Viverra prehensilis, Desmarest, Mamm. p. 208, 1820 (not of Kerr,
1792).
Paradoxurus pallasii, Gray, Proe. Zool. Soo. 1832, p. 67 ; IUustr.
Ind. Zool. pi. 8, 1834, and subsequent papers.
Paradoxurus quadriscriptus (Hodgson MS.), Horsfield, Ann. Mag.
Nat. Hist. (2) xvi, p. 106, 1856 ; Proc. Zool. Soo. 1856, p. 396,
pi. 48.
Paradoxurus nigrifrons, Gray, Proo. Zool. Soo. 1864, p. 535.
Paradoxurus vicirms, Sehwarz, Ann. Mag. Nat. Hist. (8) vi, p. 230,
1910.
Paradoxurus hermaphroditus pallasii, Pocock, Journ. Bomb. Nat.
Hist. Soc. xxxvii, p. 187, 1934; id., Proo. Zool. Soo. 1934,
p. 615*.
* The typical specimen of prehensilis needs no discussion, since the
name is unavailable. The figure and description of pallasii were taken
from a living specimen in the Zoological Gardens brought by Buchanan
from India, probably from Upper Bengal. The figure represents
it as in good coat, grey in hue, with a decided pattern of stripes and
spots, and a conspicuously white forehead. The type in the British
Museum has the coat thick, but much shorter than in typical skins
of bondar, but the pattern is not so definite as in Gray's figure, being
apparently obscured by the recent make-up of the skin on modern
PARADOXtTRTJS. 401
Locality of the type of prehensilis, Bengal (Buchanan) ; of
pallasii, India (Buchanan) ; of quadriscriptus, Nepal (in the
hills) ; of nigrifrons, India ; of vicinus, probably Assam.
Distribution. — Nepal, Sikkim, Upper Bengal, Assam, and
Upper Burma.
Wintsr coat full, with abundant wool, but shorter than in
bondar, laneus, and scindise, the wool between December and
March from 26 to 34 mm., the contour hairs from 34 to 43 mm.
The comparatively short black tips of the contour hairs
not concealing the ground-colour, which varies from clear grey
to rich ochreous, nor the pattern, consisting of well-defined
stripes and spots ; the mask consisting typically of a white
patch on the muzzle, another below the eye and of the brow-
oand which is usually more or less interrupted in the middle
line by black speckling ; basal half of tail generally striped.
Variations in the tint of the ground-colour, sometimes well
marked in skins from the same locality, are shown by the
following specimens : — One from Narbong, Darjeeling, 2,000 ft.,
one from Sevoke, south of Darjeeling in Upper Bengal (Crump),
one from the Daranga River, N. Kamrup, 400 ft. (Wells),
and one from Cachar are pale buffy-grey, like the type of
quadriscriptus ; a second specimen from Sevoke and one from
Hasimara, Bhutan Duars, 600 ft. (Baptista), are decidedly
more buffy than the first lot ; others from Hasimara are
richer, more ochreous, almost orange-oehreous in hue and
grey or buff below. These and two skins from Golaghat,
Upper Assam, resemble the type of vicinus ; one from the
Garo Hills (Wells) is intermediate between the richer and
duller skins, and one from Haldibari, Cooch Behar, 150 ft.,
is dull-tinted in front, rich-tinted behind. A young specimen
from the Uyu Forest, 60 miles east of Homalin in Upper
Chindwin, resembling in coat and colour the richer-tinted
Assamese skins, attests the extension of this race into Upper
Burma. A pair of adult specimens from Haraincha, Morang,
in the eastern Nepal Tarai (F. M. Bailey), February, resemble
those described above from districts farther to the east. The
<J has the full winter coat thick, but only about 40 mm. long j
the underhair of the back has a decided buffish tinge, the belly
is buffy-grey, and the pattern consists of well-defined dorsal
lines ; the hairs of the brow have been to a great extent rubbed off.
but the area seems to have been extensively white. The type of
quadriservptus is buffy-grey, has a longer, shaggier eoat than the type
of pallasii, and hardly differs from the alleged type of Mrsuius, except
that the pattern of stripes is distinct. It connects the two races
pallasii and bondar, as is not surprising from its locality, the hills of
Nepal. The type of nigrifrons is like that of pallasii, but has a tinge-
of buff and the crown blacker ; and the type of moinvs is similar, but
richly tinted, and has the pattern definitely arranged.
VOL. I. 2d
402 VTVERBIDJS.
stripes and fainter lateral spots. The $ differs in having the
underhair of the back and the belly much richer, almost
ochreous-buff, and the white brow-hand more conspicuous.
The only available flesh-measured skins of adult specimens
are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Easimara, Bhutan Duars ; ad. <J 24$ 23 3f
Morang, E. Nepal Tarai; ad. <J 24£ 20| 3£
Hasimara, Bhutan Duars ; ad. $ 23£ 22| 3|
Sevoke, Upper Bengal ; ad. <J 22 2lJ 3$
Duranga River, N. Kamrup ; ad. <J . . 22 — Z\
Sevoke, Upper Bengal ; ad. ? 22| 22£ 3|
Morang, E. Nepal Tarai; ad. ? 23 20f 3
The weight of a young adult <J from Haldibari, Cooch
Behar, was 5| lb., of the <J from the Duranga River 5 lb.
45 h. Paradoxurus hermaphroditus laotum Gyldenstolpe.
Paradoxurus hermaphroditus laotiwm, Gyldenstolpe, K. Sv. Vet.-
Akad. Handl. lvii, no. 2, p. 26, 1917 ; id., Journ. Nat. Hist. Soo.
Siam, iii, p. 147, 1919 ; Robinson & Kloss, Rec. Ind. Mus. xix,
p. 179, 1920 ; a. M. Allen, Amer. Mus. Novit. 359, p. 4, 1929;
Osgood, Field Mus. Nat. Hist., Zool. xviii, p. 267, 1932;
Pocock, Proc. Zool. Soc. 1934, p. 620, fig. 1, A (skull), pi. 1, A,
B (heads).
Paradoxurus birmanicus, Wroughton, Journ. Bomb. Nat. Hist.
Soe. xxv, p. 51, 1917*.
Paradoxurus hermaphroditus ravus, Pocook, Journ. Bomb. Nat.
Hist. Soo. xxxvii, p. 314, fig. 5, A (skull) , 1934 (not P. ramus
Miller, 1913) .
Locality of the type, of laotum, Chieng Hai, N.W. Siam ;
of birmanicus, Mingun, north of Sagaing, Upper Burma.
* It is a little doubtful if laotum is the correct name for this race.
It was given by Gyldenstolpe to a specimen larger both in its body
and skull-measurements than any of the known Burmese specimens,
as shown in the tables of measurements. But Gyldenstolpe's statement
that the types of laotum and of birmanicus, the former name antedating
the latter by one month, are representatives of the same race is usually
accepted. The type of laotum, however, may be the smallest known
example of a possible race, distinguished from the Burmese by its
larger size, of which two other specimens are known. One of these,
an old o\ collected by Col. C. S. Stoekley„at Mewong in N.W. Siam,
near the type-locality of laotum, has the largest known skull in the genus
Paradoxurus, its total and condylobasal lengths being respectively
130 and 125 mm., a few mm. longer in both respects than the skull
of the type of the Kashmirrace, P.h. veUerosus (p. 397). This Mewong
skull was figured and described with full measurements in my paper
(Proc. Zool. Soc. 1934, pp. 618, 623, 625). The other large Siamese
skull, that of an old ? from Muek Lek, west of Korat in S. Central Siam,
has the total and condylobasal lengths 125 and 121 mm. respectively.
These two Siamese skulls areprovisionaUy regarded asaberrant "giants "
of laotum,. The skull of the type of the latter is nearly intermediate
in size between the smaller of the two and the largest of the Burmese
PAKADOXtr-RTTS.
403
Distribution. — Btjkma from Mandalay and Chindwin to
Tenasserim ; also West and North Siam, Indo-Ohina, and
Hainan.
Resembling pallasii in the distinctness of the pattern
throughout the year, but with the wool of the winter coat
shorter and less luxuriant, and the contour hairs a little
longer, up to 46 mm., so that the coat is shaggier and not
so close. The colour on the average is greyer, rich buff
or ochreous, skins being comparatively rare, and the brow-
band is typically a more conspicuous feature of the mask,
being more extensive, less invaded by black speckling, and
frequently continued backwards over the middle line of the
crown, which, however, varies from black to white in specimens
from the same locality. The skull, on the average, has the
Fig. 95. — The larger Burmese race, Paradoxurus hermaphroditus laotum,
from a drawing by Tickell of a specimen from Moulmein. (From
Blanford.)
muzzle broader, the postorbital processes shorter, the external
pterygoid crest larger, the sagittal crest earlier developed,
and the dentition more robust than in the Indian races.
Specimens have been recorded from the following localities
in British Burma : — Mingun, north of Sagaing ; Lower Chindwin ;
skulls. If there should prove to be a larger Siamese race taking the
name laotum, Wroughton's name birmanicus will come in. for the Burmese
form. But the available evidence is at present in favour of the view
here adopted. In the first of my papers of 1934 I cited both laotum
and birmanicus as synonyms of ravus, a name given by Miller in 1913
(Smiths. Misc. Coll. xlix. no. 21, p. 2) to two specimens from Trang in
Peninsular Siam. I had not then seen examples definitely assignable
to ravus; but later a series, kindly lent to me by the Smithsonian
Institution, showed my identification to be incorrect, ramus being
distinguished from the Burmese race by its smaller size. It does not
occur in British Burma.
2d2
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PARADOXTJEUS. 405
Mt. Popa, 4,960 ft. ; Toungoo, 500 ft. ; Lower Pegu ; Thaton,
north-west of Moulmein ; Tenasserim Town.
The series from Mingun, the typical locality of birmanicus,
exhibits the seasonal and individual variations in this race.
A skin dated January 11 has comparatively abundant
underwool, the ground-colour is whitish-grey, with a faint
buff tinge, but the black tips of the copious contour hairs
somewhat obscure the pattern, giving a marbled coloration.
Other skins, dated July, have little or no underwool, the ground-
colour varying from ashy to buffy-grey, the pattern well
defined and not obscured by the contour hairs of the old
winter coat, which are mostly still retained, although moulted
on the fore quarters in one case ; the brow- band may be
uniformly grey, or mesially speckled with black or completely
divided by a black line, and the crown may be black or mesially
speckled with white as far as the nape.
Skins from the other localities similarly vary in the coloration
both of the head and body. Three skins, namely, one from
Nan in Lower Chindwin, one from Toungoo, and one from
Thaton, are unusually bright-tinted, ochreous-buff like the
majority of skins of pallasii.
The following are some flesh-measurements (in English
inches) and weights (in lb.) : —
Locality and sex.
Tenasserim Town ; just ad. <J .
Tenasserim Town ; just ad. (J .
Mingun (birmanicus type) ; ad.
Kin, Lower Chindwin ; ad. $ .
Thayagon, Chindwin ; ad. $ . .
Tenasserim Town ; ad. $. . . .
Head and
Hind
body.
Tail.
foot.
Weight.
24
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The type of laotum, an adult $, is a little larger than any
of the Burmese skins, the head and body being 25£ in., the
tail 21^ in.
According to Shortridge's observations at Mingun and
on Mt. Popa, this Palm-Civet is apparently plentiful
throughout the dry zone in Upper Burma, where its habits
are the same as those of the typical race of hermaphroditus
that he collected in S.W. India.
45 *. Paradoxurus hermaphroditus minor Bonhote.
Paradosourus minor, Bonhote, Fasc. Malay., Zool. i, p. 8, 1903 ;
Miller, Smiths. Misc. Coll. lxi, p. 2, 1913.
Paradoxurus hermaphroditus minor, Pocock, Journ. Bomb. Nat.
Hist. Soc. xxxvi, p. 870, text-fig. 1, D (head), 1933, and xxxvii,
p. 322, fig. 5. B, 1934 ; id., Proe. Zool. Soc. 1934, p. 616.
Locality of the type, Kampung Jalor in Peninsular Siam.
406
V1VERBIDJE.
Distribution. — Loweb Bttema and S.W. Siam through
Tenasserim and Peninsular Siam to the Malay Peninsula and
possibly Annam.
Closely resembles pallasii in colour and pattern and in the
characters of the skull and teeth, but smaller and "with the
white areas of the mask on the average less distinct, the
muzzle-patch being absent and the brow-band typically
more obliterated by black. Differing from laotum, which it
overlaps in range in Lower Burma, by its smaller size, narrower
muzzle, and less robust, more trenchant teeth. Also the
A B
Fig. 96.
A. Skull of adult <J of the larger Burmese race, Pwadoxurus herma-
phroditm laotum, from 30 miles west of Sagaing, Upper Burma.
B. Skull of adult £ of the smaller Burmese race, P. h. minor, from
Bankaehon, Tenasserim.
ground-colour is on the average brighter and more ochreous,
and the mask dominantly black instead of white.
This race varies considerably in colour, from rich ochreous-
buff to grey, and also in size. Northern specimens are larger
than southern, suggesting the possibility of more than one
race being involved, but the available material hardly warrants
the introduction of new names.
PABADOXUBTJS. 407
Only three skins have been recorded from British. India,
two from Tenasserim, one from Bankaehon, Victoria Point,
the other from Thaget on the Little Tenasserim River (Short-
ridge), and the third from the Zamayi Reserve, 80 miles north
of Pegu Town (Mackenzie). The Tenasserim skins, collected
in December and March, are in full colour and coat, the
contour hairs measuring 35 and 32 mm. respectively, only
a trifle shorter than iapaUasii, and the general colour and pattern
are similar to those of the brightest-tinted specimens of that
race, although the mask is blacker. The specimen from the
Zamayi Reserve, dated April 30, being in full moult, is very
different in colour, apparently from the fading of the dead
hair. The contour hairs are shed and the short-haired coat
is greyish-buff, with the pattern brown and ill denned, and
the neck and head are dull brownish-grey.
Of other known specimens of this race, the type from Jalor
in Peninsular Siam is paler than the Tenasserim skins ; but
one from Perak (June) is brighter coloured, nearly orange-
buff on the body, and has the head largely brownish-grey
as in the Zamayi Reserve skin ; others from Selangor, 4,800 ft. ,
vary from buffish-grey to cream-buff.
The following flesh-measurements (in English inches) and
weight (in lb.) show the variations in this race : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Zamayi Res. ; imm. $ 18f 20 3 —
Bankaehon ; ad. <J 19§ 19 3+ 4*
Selangor ; ad. $ 17| 18- 3- —
Selangor; ad. $ 17£ 17- 2*- —
Jalor (type) ; yg. ad. $ 18 18} 2% —
The specimen from Thaget in Tenasserim is almost exactly
the same size as the one from Bankaehon, but its weight
was 5\ lb. The example from the Zamayi Reserve had not
shed its milk-teeth and would probably, when full grown,
have surpassed the Bankaehon example in size. Both
are considerably larger than the <J from Selangor, which seems
to be the smallest adult $ Paradoxurus yet recorded.
The skull of the adult (J from Bankaehon, included in the
table, is well developed, with a high sagittal crest, narrow
" waist," and strong postorbital processes. The immature g
skull, from the Zamayi Reserve, with the milk-teeth all in use
and m 1 just erupted, has the temporal ridges 10 mm. apart on.
the parietals, although the skull is not far short of its potential
full length, thus showing that the sagittal crest is late in
forming as in pallasii, and not precocious, as indicated by the
skulls of laotum, of about the same age, from Sagaing, Toungoo,
and Tenasserim referred to above. Similar lateness in the
formation of the sagittal crest is shown by $ skulls of minor,,
408
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PARADOXUBTJS. 409
in none of which is the crest formed, the temporal ridges being
9 mm. apart on the parietals in a just adult skull from Selangor
and about 10 mm. in the almost adult $, the type, from Jalor.
Probably only aged females acquire the crest.
The largest (J skull known is that of an adult from 100 miles
north of Bangkok, figured in my paper (Proe. Zool. Soc. 1934,
p. 618, fig. 1, B). With a eondylobasal length of 107 mm.
and a zygomatic width of 60 mm. it is 5 mm. longer than the
<J skull from Bankachon, some 12 mm. longer than adult $
skulls from Selangor, and the same length as the adult $ skull
of pallasii from Sevoke, south of Darjeeling.
The status here accorded to P. h. minor is open to dispute
on the grounds that it inhabits the same countries as two
larger, intergrading races, namely, laotum in Lower Burma
and ravus in the Malay Peninsula, without, so far as is known,
intergrading with them. For these reasons it was considered
by Bonhote and Miller as a distinct species. On the other
hand, it is obviously very closely akin to pallasii, differing
from that race merely in its average smaller size, in my opinion
a subspecific character.
In Tenasserim Shortridge found this Palm- Civet plentiful
about the villages, where it hides by day in the roofs of houses ;
but Mackenzie's specimen from the Southern Zamayi Reserve,
Pegu, was shot in a tree at dusk.
The described Races of Paradoxurus from the
Mergui Archipelago.
Six subspecies of P. hermaphroditus were named by G. S.
Miller from some of these islands (Smiths. Misc. Coll. Ixi,
no. 21, pp. 3-5, 1913). They are known to me only from the
descriptions. Their distributions, names, and alleged distinc-
tive features, epitomized in my paper (Journ. Bomb. Nat.
Hist. Soc. xxxvii, pp. 324-6, 1934), are as follows : —
P. h. senex. Domel Island. Based on a single young
adult $ distinguished from the typical examples of ravus
Miller (cf. supra, p. 403, note), which has page priority, from
Trang in Peninsular Siam, by having less traces of yellow on
the back and sides and greatly reduced auditory bullse, which
.are slightly inflated and only \0\ mm. long. Miller himself
was doubtful about the constancy of the difference in colour
and, considering the recently recorded variations in tint from
clear ashy-grey to brownish-grey and buff in skins of ravus
from Bagnara, Patani (see Proc. Zool. Soc. 1934, p. 636),
no reliance can be placed on the characters derived from
a single specimen. The length of the bulla is also too variable
a feature to be trusted. In one specimen, for example, of
laotum, from Tenasserim Town, the bulla, measured along its
inner surface, is 10J mm., in other specimens it is 13 mm.
410 VrVEBKID.®.
P. h. fuscus. James Island. Based on a single adult $
distinguished from senex by its more extensive black markings
and sooty-brown underside. Both these features are
individually variable in mainland forms.
P. h. pollens. Kissaraing Island. Based on a single
adult <J distinguished from typical ravus by having the
tail distinctly brownish instead of essentially black, its basal
third being like the head and body and contrasted with its
distal portion. In the mainland race laotum this difference
has no systematic value, being attributable to seasonal
fading of black to brown.
P. h. pugnax. Sullivan Island. Based on two specimens
resembling pollens from Kissaraing, but slightly smaller,
and with the dark markings rather more extensive, especially
on the thighs, the upper parts of the fore legs, nape, and cheeks.
The distinctness of the pattern on the areas named is variable
individually in the mainland race laotum.
P. h. sacer. St. Matthew and St. Luke Islands. Based on
five specimens ]ike pattens from Kissaraing, but with the stripes
reduced and broken up into lines of spots, the skull smaller
and scarcely exceeding that of P. h. minor. The disintegration
of stripes into spots is not unknown as an individual character
in some races of P. hermaphroditus. It has been recorded,
for example, in skins of ravus from Patani ; but its occurrence
in the five known examples from these islands suggests that
subspecific importance may be attached to it.
P. h. pulcher. Clare Island. Based on two specimens
like pollens of Kissaraing, but the ground-colour above and
below between " cartridge-buff and cream-buff," more yellow-
ish than in any of the other races from the Archipelago.
Such differences in tint are not uncommon in individuals
of several races of P. hermaphroditus.
The flesh-measurements (in English inches) of the types of
these races are as follows : —
Head and
Locality, name, and sex. body. Tail.
Domel Island (senex) ; yg. ad. (J 21 18|
James Island (fuscus) ; ad. $ 20 —
Kissaraing Island ■pollens) ; ad. $ 21 J 17
Sullivan Island (pugnax) ; ad. <j> 20f 17 —
St. Matthew Island (sacer) ; ad. $ 20f 17 —
Clare Island (pulcher) ; ad. $ 19£ 18J
These measurements do not indicate any reliable differences
in size between the races, but they show that collectively
these insular forms are a trifle smaller on the average than in
the mainland forms laotum and ravus, two adult $ examples
of the latter from Trang and Jalor in Peninsular Siam giving
PARADOXURTTS.
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412 VIVEBBIDJE.
a head and body length of over 23 in. But the dimensions
agree closely with a small race named cochinensis by Sehwarz
in 1911, and the range of this race has been recorded from
Cochin China to Peninsular Siam (see Proc. Zool. Soc. 1934,
p. 626). It is quite possible that cochinensis, which has two
years priority over the names given to the Mergui series, may
come in for one or more of the races from that Archipelago,
assuming that more than one race is admissible. The skull-
measurements also agree, as stated below. But until more
examples have been collected from the islands to show if the
characters relied upon by Miller have the value assigned to
them, and if they are different from those of cochinensis,
the names proposed by Miller must provisionally stand.
The skull-measurements, approximately as recorded by
Miller, appear on p. 411.
The variation in the condylobasal length of the skulls
from 99 to 103 mm. is not greater than that of mainland skulls
assigned to laotum and raws, in which the length is respectively
110 to 117| (omitting " giant "-skulls) and 104 to 108 mm.
The. Mergui series is decidedly smaller than either, but it is
the same as in cochinensis, as stated above.
Habits. — The habits of all the British Indian races of this
Palm-Civet appear to be similar. Although an expert climber
and mainly arboreal in forest and jungle, it sometimes lives
amongst rocks, and is everywhere liable to become partially
parasitic on man, quite commonly taking up its abode under the
roofs of bungalows near the jungle, in old drains, or even in
houses in villages and towns. As lately indeed as the end of the
last century it occurred in Calcutta, as stated in his edition of
Sterndale by Finn, who observed one climbing up the stack-
pipe of his house, and subsequently trapped a pair. The
inducement to attach itself to man is no doubt the food it
thereby secures. The roofs of bungalows harbour rats
and other vermin, and as a destroyer of these the Palm-Civet
might be welcomed were it not for its unclean habits and,
being nocturnal, for the disturbing noise it makes overhead
catching its prey at night. Prater, too, reported that at
Satara it was a nuisance to the shopkeepers by raiding their
stores of provisions, for, being omnivorous, it will eat fruits of
many kinds as well as animal food, and is a notorious poultry
thief. It preys, in fact, on any small mammals and birds it
can overcome, as well as on snakes, lizards, frogs, and insects.
In Ceylon, according to Phillips, it is very fond of the fruit
of the passion flower vine, and may cause considerable loss
to the growers of pineapples by raiding the plantations just
before the fruit is ready for plucking. At Dharwar and Coorg
Shortridge found it frequently in the neighbourhood of coffee-
PAKADOXURTJS. '±13
estates, attracted by the berries, which it eats in quantities
when ripe. But the damage done in this case is not so serious
as might be expected, because the animal can digest only the
soft pulp of the berries, and excretes the hard beans unaltered.
It has too the habit of repeatedly depositing its droppings
in heaps on the same spot — it may be on a path, or rock, or
fallen tree-trunk in the jungle ; and the coolies, after gathering
the harvest from the estate, hunt for these accumulations
and add the recovered beans to their store for the market.
Seeds of the peepal, according to Finn, are passed and deposited
in the same way. It is probably the attempt to masticate
these and other hard vegetable substances that accounts
for the manner in which the grinding teeth of this Palm-Civet
are not infrequently worn down to the gum even in skulls not
noticeably very old. As has been repeatedly told, in districts
like Ceylon and S. India for example, where the " Kitual "-
palm flourishes and is tapped for its juices, or " toddy," this
Civet's habitual robbing of the pots in which the natives
collect the sweet stuff has earned for it the title "Toddy-Cat."
As stated above, the animal is active at night. It spends
the day curled up asleep in a hollow tree or any suitable
sheltered place.
In Ceylon, according to Phillips, the young, three or four to
the litter, are born in holes in trees, the " Kitual "-palm, where
available, or in recesses under overhanging rocks. They
may be found at all times of the year, but more commonly
in the latter part before the onset of the north-east monsoon.
In India, also, judging from the evidence supplied by 2 skins
with the teats enlarged and the area round them naked, the
young may be produced at all months of the year. Skins
in this condition were collected in Coorg and Kanara on
January 11 and February 9, and in Dharwar on November 3
and December 8 ; at Hazaribagh on March 8, at Dharbanga
on July 28, Rajputana on September 15, Kangra on March 28,
and Lower Chindwin on June 4. The statement that the
litter consists of from four or five to six in India has been
repeatedly copied ; but it is unlikely that the number exceeds
that of Ceylon.
Some additional observations on this species are of great
interest. McMaster ('Notes on Jerdon's Mamm. of India,'
p. 44, 1870) recorded that on returning one moonlight night
to his garden at Russellkonda in Orissa he found that his
dogs, three or four eager, hardy terriers and spaniels, had
treed a Paradoxurus*. Unwilling to shoot the animal,
* He identified the animal as Paradoxurus grayi (=Paguma larvata
yrayi, see p. 420), a Himalayan species, and this was accepted
by Blanford. But from the locality there is little doubt the Palm-Civet
was an example of P. hermaphroditus nictitatans.
414 vivebbim;.
he sent his attendant up the tree to dislodge it and drive it
to the ground. " It did not appear to notice his approach
till he was almost within arms' length," whereupon " the
man uttered an acute shriek of mingled horror and rage as
he was suddenly sprinkled all over with some detestable
fluid it discharged at him, so keenly fetid that it disabled him
from using hand or eye, and at the same time covered the
retreat of the beast as it bolted down the tree past me and
through the dogs, driving me back in discomfort, and preventing
them from closing with it . . . The stench was vile and
abominable beyond any other I have had the ill-fortune to
experience, and I am certain that by it the beast saved its life."
For many years this was the only recorded evidence that the
Toddy-Cat can discharge from its anal glands a nauseating
defensive fluid. This was confirmed by Crump who, at Sevoke,
found a pair of P. h. bondar in a tree. When shot, " both
gave out an intensely fetid odour, and when they were being
carried to camp through the forest all the jungle fowl became
very excited, and with much cackling flew up into the bushes
and trees." *
From these two cases it may be inferred with certainty
that all examples of this species of Palm-Civet are similarly
protectively endowed. The apparent infrequency with which
they make use of the secretion, at all events where man is
concerned, is in keeping with the general unwillingness of
mammals protected in the same way to employ the secretion
except as a last resort.
A note by Col. D. D. Chmningham (' Some Indian Friends
and Acquaintances,' 1903) has a bearing upon this subject.
He wrote of the Indian Toddy-Cats as " wonderfully fearless
animals," adding that " their eyes are strangely luminous in
dim light, much more so than those of almost any other
animal save death's-head moths." The interest of these
remarks lies in fearlessness and conspicuousness being well
known attributes of distasteful, poisonous, or nauseously
■odorous animals ; and the knowledge that this Palm-Civet
comes into the last category throws light upon its coloration,
which, apart from the not always manifest stripes and spots,
ahows, especially in some races from Ceylon and Hindustan,
marked resemblance to that of the typical Polecats (Putorius),
■a resemblance which no doubt suggested that name for the
species in Ceylon. In both the legs are blackish, the face
has a distinct " mask " of dark and light patches, and the
back is covered with long, black-tipped contour hairs which,
when erected, display the whitish or yellowish underhair.
* It is not certain ■whether the perturbation of the fowls was due to
the stink or sight of the Palm-Civets.
PLATE XXIX.
MAMMALIA.
Photo F. W. Bond.
Burmese Palm-Civet (Paradoxurus hermaphroditus /ootum).
Phoio F. W. Bond.
Masked Palm-Civet (Paguma larvata larvata) from Szechwan.
PAGUMA.
4=15
In the Polecat this coloration is known to be associated with
the protective nauseous secretion of the anal glands, and is
believed to have a warning significance, making the animal
conspicuous, readily recognized, and avoided by enemies
after one or two experiences, precisely as in the Skunks,
which are similarly equipped and coloured for advertisement.
No doubt the same explanation applies to the colour of this
Palm-Civet.
Genus PAGUMA Gray.
Paguma, Gray, Proc. Zool. Soc, 1831, p. 94 ; and of most recent
authors, including Pocoek, Journ. Bomb. Nat. Hist. Soc. xxxvii,
p. 326, 1934, and Proe. Zool. Soc. 1934, p. 665.
Ambliodon, Jourdan, C. R. Acad. Sci. Paris, v, p. 445, 1837.
Paradoxurus, Blanford, Mamm. Brit. Ind., p. 18 (in part).
? Osmetectis, Gray, Ann. Mag. Nat. Hist, x, p. 260, 1842.
Type of Paguma, larvata Hamilton-Smith ; of Ambliodon,
jourdanii Gray ; of Osmetectis, fuscus Gray *.
Fig. 97. — Right half of the bony palate of Paguma larvata grayi, show-
ing the dentition and the fore part of the mesopterygoid fossa, to
compare with fig. 91, p. 380. (From Blanford.)
* By Anderson and Thomas Osmetectis fuscus was cited, but obviously
wrongly, as a synonym of Herpestes urva, the Crab-eating Mongoose.
The generic name was given by Gray to a species he had previously
figured as Viverra fusca (Hardw. ULustr. Ind. Zool. pi. i, 1830). This
figure, however, with a long snout, large ears, and cat-like feet, is not
the least like any species of Mongoose. It may be a caricature of
Paguma larvata grayi.
416 viverbijxs:.
Distribution. — From Kashmir and Tibet through the
Himalayas into China, Formosa, and Hainan, Indo-China,
Burma, the Andaman Islands, Malay Peninsula, Sumatra,
and Borneo.
Distinguished from Paradoxurus by the entire absence of
the body-pattern, except sometimes, at least, in the newly-born
young (see p. 430), and by the presence of two pairs of mammae
instead of three. Also by the skull, which has the palate
produced posteriorly so as to overlap to a greater extent the
anterior portion of the mesopterygoid fossa, and is everywhere
less affected by the action of the masticatory muscles, the
postorbital area, or " waist," being broad and comparatively
unconstricted, and the postorbital processes consequently less
salient ; the teeth, too, are reduced in size and less trenchant.
Nearly all the described forms of this Palm-Civet are regarded
as local races of a single species, P. larvata ; but there is an
imperfectly known form which on the available evidence may
provisionally be granted specific status.
The two may be briefly distinguished as follows : —
a. Tail about half the length of the head and body ; [p. 416.
winter coat woolly and matted lanigera (Hodgs.),
6. Tail over two -thirds the length of the head and
body, typically only two or three inches Smith), p. 417.
shorter ; winter coat much thinner larvata (Ham.-
46. Paguma lanigera (Hodgson).
Paradoxurus lanigerus, Hodgson, Asiatic Res. xix, p. 79, 1836.
Paradoxurus laniger, Hodgson, Journ. As. Soc. Beng. x, p. 909,
1841, and xi, p. 279, 1842 ; Blanford, Mamm. Brit. Ind. p. 114,
1888.
Paguma grayi, Wroughton, Journ. Bomb. Nat. Hist. Soc. xx, p. 50,
1918 (not grayi Bennett).
Paguma lanigera, Pocock, Journ. Bomb. Nat. Hist. Soc. xxvii p.
327, 1934.
Locality of the type, " the northern region of Nepal," subse-
quently said to be Tingree in Tibet.
Distinguished from the other known forms of Paguma
and from all the species of Paradoxurus by the shortness of the
tail, which is at most only a little more than half the length
of the head and body.
In the only known specimen, the type, an imperfect, no
doubt immature skin, without skull (B.M. no. 43.1.12.103),
the dorsal coat from the crown backwards is very thick',
woolly, and matted, the moult apparently being imminent •
the wool is copiously mixed with fine whitish hairs with curled^
frizzled ends hardly projecting beyond the summit of the wool^
which is about 40 mm. long ; the hairs of the face, paws'
throat, and belly are sleek and comparatively short. General
colour pale brown, darker and more rufous on the back than
PAGUMA. 417
on. the flanks ; the legs and upper side of the tail like the back ;
the -ventral surface from the throat backwards mostly creamy-
white. The tail, which is woolly like the back, is thick at
the base and gradually tapers to the point.
A peculiarity of the coloration is the entire absence of black
pigment in the pelage, and there is no trace of the " mask "
on the imperfectly preserved skin of the head. But the con-
dition of the coat suggests that many of the contour hairs
were shed at the time of death, and that those still retained
were dead, with the terminal portion broken, the whole
coat being no doubt seasonally faded. Probably the fresh
summer coat has black and buff or grey contour hairs. Another
peculiarity is the thickness of the base of the tail, suggesting
an accumulation of fat in the winter months, during which
the animal possibly hibernated.
The approximate measurements of the skin, are : head and
body 20 in., tail, apparently complete, 9 J in., but Hodgson
gave its length as 12 in. Possibly Hodgson had more than
one example of this species. His unpublished drawings
contain three illustrations, and one of them is more richly
coloured than the others.
It is doubtful if this species comes within the limits of the
fauna of British India, although originally recorded by
Hodgson from the " northern region of Nepal." In his second
paper he cited Tingree in Tibet as its locality. Of its habits
nothing has been recorded.
47. Paguma larvata (Hamilton-Smith).
Gulo larvatus, Hamilton-Smith, Griffith's Anim. Kingd. ii, p. 281,
pi., 1827.
Paguma larvata, Gray, Proc. Zool. Soc. 1831, p. 94.
Distribution as under the generic name, with the omission
of Tibet.
Distinguished from P. kmigera, as stated above, by its
considerably longer tail and less luxuriantly woolly winter
coat.
The typical southern Chinese race of this species, P. larvata
larvata, which is not known to occur within the limits of the
British Indian fauna, is a rather small, thick-coated form,
with very conspicuous black-and-white mask, the general
colour varying from foxy-red to clear grey, with the tips of
the long contour hairs inconspicuously darkened.
Of the British Indian races here admitted, Blanford (Mamm.
Brit. Ind. pp. 112-14, 1888) referred specimens representing
wrougMoni, grayi, neglecta, and tytlerii to Paradoxurus grayi.
There is no evidence that he was acquainted with intrudens,
robusta, a,nd janetia. If he had known them he would, no doubt,
vol. i. 2 E
418 -VIVERBID.2E.
have assigned the first to Paradoxurus larvatus and the
others to P. leucomystax. He regarded the Palm-Civets of
this geims, which he did not distinguish from Paradoxurus,
as belonging to three distinct species — P. grayi, occurring
in the Himalayas, Burma, and the Andamans; P. larvatus,
in S. China ; and P. leucomystax, in Malacca. There is now
evidence that these forms intergrade. The reasons for the
synonymies given below were fully discussed in my two
papers quoted under the subspecific headings, and need
not be repeated.
Key to the British Indian Subspecies of Paguma larvata.
a. The preaural crest of hair comparatively close
to the ear and some distance behind the eye.
b. Head mainly Hack, the pale areas of the [nov., p. 424.
mask greatly reduced in extent nigriceps, subsp.
6'. Head not mainly black, pale areas of mask
extensive and typically well defined,
c. Pattern of mask moderately emphasized,
the median band confluent on the fore-
head with the pale area on the cheek,
and not continued all along the nape.
d. Larger, winter coat longer and thicker,
pale areas of mask lighter than tint
of back.
e. Nape nearly black, sharply con-
trasted with grey of crown and [p. 418.
darker than back wroughtoni Schwarz,
e'. Nape typically copiously speckled
with grey and similar in tint to
crown and back
/. Winter coat longer and thicker. . . grayi (Bern.), jj. 420.
/'. Winter coat shorter and thinner. . neglecta Poc, p. 422.
d'. Smaller, winter coat thin and short,
pale areas of mask similar to tint of
back tytlerii (Tytl.),p. 424.
o'. Pattern of mask strongly emphasized,
black and white, the median band
bordered throughout with black, not
confluent with the pale area of the cheek [p. 425.
but continued along the nape intrudens Wrought.,
a'. The preaural crest of hair reaching nearly to
the eye and far in front of the ear.
g. General colour tawny, nape at most pale [p. 426.
brown robusta (Miller),
g'. General colour much darker, nape blackish-
brown , janetta Thos., p. 427.
47 a. Paguma larvata wroughtoni Schwarz.
Paguma grayi wroughtoni, Schwarz, Ann. Mag. Nat. Hist. (8) xii,
p. 289, 1913 ; Wroughton, Journ. Bomb. Nat. Hist. Soc. xx,
p. 51, 1913.
Paguma larvata wroughtoni, Pocock, Journ. Bomb. Nat. Hist.
Soc. xxxvii, p. 329, fig. 6, 1934.
Locality of the type, Gharial near Murree, Upper Punjab.
PAGTTMA. 419
Distribution. — Kashmir and the Upper Punjab, eastward
to Garhwal and Kumatjst, where the race blends with the-
next.
Mask tolerably well defined, the head dominantly grey above
and in front of the ears, with the sides of the muzzle, the area
round the eye, the chin, fore-throat, and lower cheek blackish-
brown, but relieved by a conspicuous grey patch below the
eye ; the grey of the crown invading the fore part of the nape
more or less as a median streak set off by the generally blackish
hue of the rest of the nape and shoulders, which are at most
inconspicuously speckled with grey ; the back darkish grey,
black speckled with grey in front, with bumsh-grey behind ;
tail like the back mostly, but gradually darkening towards,
the tip ; the underside whitish ; the legs like the flanks,
but with paws dark brown ; the dorsal contour hairs of the
coat vary seasonally from aboiit 30 to 55 mm., but are usually
about 50 mm. (2 inches).
No flesh-measurements of adult specimens are available.
Skins assigned to this race have been collected at Gharial
near Murree (H. N. Dunn) ; at Sopor, near Lake Walar in
Kashmir ; at Chamba, 6,000 to 9,000 ft. (H. W. Wells) ;
at Dharmsala, 5,000 ft. (H. Whistler) ; and at Simla (A. O-
Hume). But the Simla skin approaches the next race, grayi,
in having the grey of the mask less conspicuous than in typical
wroughtoni. Of two skins from Salim, near Lansdowne in
Garhwal, 2,750 ft. (D. E. Lowndes), one, September, is very
like the type of wroughtoni, but has the coat shorter, only
30 instead of 45 mm., and the back less speckled with buff,
whereas the other, May, has the coat about 43 mm,, the back
much more buff, and the nape and shoulders speckled with
buff as in grayi. Two skins from Dhakuri in Kumaun, 9,000 ft.
(Crump), similarly connect the two races, the nape of one
being blackish as in wroughtoni, in the other speckled as in
grayi.
The only known adult skull of this race, that of the type,
<J, has a condylobasal length of 125 mm., a little longer than
any of the skulls assigned to the next race, grayi, and since
a young adult $ from the same locality near Murree, in the
Upper Punjab, is slightly longer than the longest known
skull of grayi, an old $ from E. Kumaun (Stockley), it may
be inferred, although the data are scanty, that typical
wroughtoni is a little larger than grayi ; but, as in the case of
the skins, no doubt the two races intergrade completely in
cranial characters.
2e2
420 viveebid^:.
47 6. Paguma larvata grayi (Bennett).
Paradoxurus grayi, Bennett, Proc. Zool. Soc. 1835, p. 118.
Paradoxurus nipalensis, Hodgson, Asiafc. Bes. xix, p. 76, 1836.
Paguma grayi, Gray, Proc. Zool. Soo. 1864, p. 541 ; and of most
subsequent writers, at least in part.
Paguma larvata grayi, Pooock, Journ. Bomb. Nat. Hist. Soo.
xxxvii, p. 330, 1934.
Locality of the type of grayi, " India " ; of nipalensis,
Nepal, mainly in the hilly region (Hodgson).
Distribution. — Nepal, and thence westwards to KtrMATn*
and Gakhwal, where the race Mends with wroughtoni.
Nearly resembling wroughtoni, but distinguished from
typical forms of that race by the duller, less silvery-grey
hue of the pale areas of the mask, by the profuse pale speckling
of the nape and shoulders, which, instead of being contrasted
by their blackness, are nearly the same general tint as the
back and loins ; also the winter coat on the average is fuller
owing to the longer thicker underwool, and not so loose,
although the contour hairs are approximately the same length.
The flesh-measurements (in English inches) and weights
(in lb.) of two examples from Dhakuri in Kumaun are as
follows : —
Head and Hind
Sex. body. Tail. foot. Weight.
Ad. <? 25£ 24£ 4+ Hi
Yg. ad. cj 23£ 22| 4- 8£
The type of this race, ticketed " India, Zool. Soc," was
a menagerie-reared specimen ; but, since it closely resembles
one of Hodgson's skins from Nepal, I accept the traditional
identification of the specimens that Hodgson named nipalensis
as grayi. Hodgson's skins, seven in number, show considerable
individual variation in colour and very marked, apparently
seasonal, differences in coat. The specimen I regard as the
type of Hodgson's nipalensis closely resembles the type of
grayi, but has the back less buffy, the hind back more speckled
with black, and the tail dark in its posterior half, not buffy
throughout. Two others are less buffy than the type of
nipalensis, the pale speckling being grey. The coat of these
varies from 40 to 46 mm. long and is thickened with under-
wool. They are probably early winter skins. A fuller-coated
Nepalese skin (Dhuleep Singh) has the coat 50 mm. and the
underwool 35 mm. long. Four additional skins collected
by Hodgson are moulting and short-coated, the underwool
being scanty. They are on the average darker than the full-
coated skins. One is speckled above with rusty ochreous,
the belly rich ochreous-buff, and even the pale areas of the
mask are buffy. Possibly it is artificially stained by smoke
or scorching. Blanford thought these skins represented
a lowland race, but no doubt the scantiness of the coat is
purely seasonal.
PAGUMA.
421
In addition to the skins from Kumaun and Garhwal referred
to above as intermediates between grayi and wroughtoni,
there is one from Swankot, 3,569 ft. (C. A. Crump), which
■*v*** ^-^
TO*
Fig. 98.
A. Head of Paguma larvata wroughtoni, drawn from skin from Gharial,
near Murree, in the Upper Punjab.
B. Head of P. I. intrudens, drawn from skin from the North Shan States.
C. Head of P. I. janetta, drawn from skin of type from Bankaehon,
Tenasserim.
also has the nape blackish, as in wroughtoni ; but two from
Nairn Tal (E. A. Smithies) and one from eastern Kumaun
(C. H. Stockley) resemble typical grayi.
422 VIVEBBIDiE.
An adult $ skull from Dhakuri, Kumaun, is only 114 mm.
in condylobasal length, considerably less than the skull of
the type of wroughtoni. Anunsexed skull from Nepal (Hodgson)
is 117 mm., whereas an old <j> skull from eastern Kumaun
(Stockley) is 118 mm., the longest observed in the race, and,
as stated above, is nearly the same length as a young adult °
.skull, a topotype of ivroughtoni.
47 c. Paguma larvata negleeta Pocock.
Paguma larvata negleeta, Pocock, Journ. Bomb. Nat. Hist. Soc.
xxxvii, p. 334, 1934, fig. 6, B ; id., Proc. Zool. Soc. 1934, p.
672.
Locality of the type, Mokokchung, 4,500 ft., in the Naga
Hills, Assam.
Distribution. — Assam, and thence apparently westwards to
.Sekkim and the low-lying districts of Nepal and southwards
■to the Chin Hills and Arakan, in Burma.
Closely resembling typical grayi in having the dorsal
surface tolerably uniformly speckled from the crown to the
tail and in the pattern of the mask, but differing in having the
proximal portion of the tail more ochreous, at least in Assamese
skins. Also the winter coat is much shorter and less luxuriant,
the contour hairs, when fully grown, being between 30 and
40 mm. long.
3?lesh-measurements (in English inches) and weights (in lb.)
-of some specimens assigned to this race are as follows : —
Head and
Hind
Locality and sex.
body.
Tail.
foot.
Weight.
Narbong, Darjeeling ; ad. <J
26-
S3*
4+
9£
Hathiban, Nepal ; ad. ? ...
25-
25-
4—
11
•Garo Hills, Assam ; ? ad. $ .
. 25-
24
4—
o*
The type, from the Naga Hills (H. W. Wells), has the dorsal
.surface speckled with black and buff, the buff being richer
•on the hind back and rump than anteriorly, the tail is rich
ochreous with the terminal two inches black, the legs are
grizzled, and the pale areas of the mask are clear silvery-
grey and the tint of the crown blends with that of the nape.
A skin from the Mishmi Hills, 2,250 ft. (H. W. Wells), is like
the type, but the mask is not so white ; another, from Duragiri,
in the Garo Hills (H. W. Wells), also has the mask duller,
the pale speckling on the back greyer and finer, and the tail
not so bright. Evidence for the extension of this race west-
wards to Nepal is supplied by two skins from Hathiban and
Thankot (Baptista), which are on the whole more like negleeta
than grayi, and to Darjeeling by a skin from Narbong, 2,000 ft.
(Crump) ; but another from the Gopaldhana Valley, Darjeeling
<Stevens}. is not definitely assignable to either race. Skins
PAGUMA.
423
g
1
o o ra o ct
a
1 &£
§3 IP
I I
oc oo 30 qo oo t- c-
CI <M *•! (M C<I <M <M
■^h >* (M ^h o I •-<
<>) (M <M CI ^ | O)
01
.60
"33
e
e
J
o
O
CO
3
.o
•8
a
a
a
©
toSS
O -3 a
f H i-H (M CI
(M <N (M (Ml CM
-H
° H ■■*
l> t~ US
T3 "3 $
g § SP
8 to
13
£
lOM'* I (^
5a
2*
~ *
«> -e
Pi S
SB
CH-
5> •Sch-Cw-
§>*°
OH- -S
°« d"3
M
° a "S"S
PHI?: fc
SHI
3 S'S-ci
I S 3 g
424 VIVEBBID^!.
from the Chin Hills, 4,000 ft., 150 miles west of Eondat
(J. M. D. Mackenzie), attest the extension of neglecta into
Upper Burma, and no doubt Blanford's record of grayi from
Arakan referred to this race, which thus has an extended
range to the west of the Chindwin and Irrawaddy.
The skull of the specimen above referred to from Narbong,
Darjeeling, an adult <J, has a condylobasal length of 120 mm.
The skull of the type, a young adult $ from the Naga Hills,
and of an old, probably <?, specimen from the Chin Hills have
the basioccipital cut away, but the total length, probably
1 or 2 mm. in excess of the condylobasal length, is 123 mm.
These data suggest that the skull of neglecta may be a trifle
larger than in typical grayi, but the evidence is far from
satisfactory.
47 d. Paguma larvata nigrieeps, subsp. no v.
Locality of the type, Nam Tamai in Upper Burma.
Distinguished from all the other known races of Paguma
by the nearly complete obliteration of the pale areas of the
mask, the head being mainly black apart from a narrow buffy-
grey stripe between the eyes, but dying out on the forehead,
the merest trace of the patch below the eyes, and a yellowish
preaural area ; the nape is blackish, finely speckled with buff ;
the back has an abundance of black and buff speckling,
the general effect being dark brown ; the tail in its basal
half is black above, yellowish-brown at the sides, and its
terminal half is black.
Although only a single adult skin, collected byLord Cranbrook
in the above-mentioned locality, is available, it differs so
noticeably from the other Burmese forms, neglecta and
intrvdens, in the general blackness of the head that I do not
hesitate to regard it as representing a distinct race of Paguma.
47 e. Paguma larvata tytlerii (Tytler).
Paradoxurus tytlerii, Tytler, Journ. As. Soe. Beng. xxxiii, p. 188,
1864 ; Miller, Proe. U.S. Nat. Mus. xxiv, p. 772, 1902.
Paguma larvata tytlerii, Pocock, Journ. Bomb. Nat. Hist. Soe.
xxxvii, p. 338, 1934 ; id., Proe. Zool. Soo. 1934, p. 672, fig. 11, A
(skull).
Locality of the type, Port Blair, S. Andaman Island.
Distribution. — Andaman Islands.
A small race with the coat thin and short, up to about
25 mm. long, and the general colour pale, the dorsal surface
from the crown nearly to the end of the tail being tolerably
uniformly speckled grey and black and the mask without
strongly contrasted hues, the pale areas being grey, like the
nape and back, and tbe dark areas pale brown.
PAGUMA. 425
The following are the flesh-measurements (in English
inches) of two specimens : —
Head and
Hind
body.
Tail.
foot.
Ad. <J (Miller) .
22
20
n
Type (Tytler) . .
21
20
—
This race was not distinguished from grayi by Blanford.
Only a few specimens of it are known. The type and the
example recorded by Miller were from South Andaman
Island- Another was collected in Rutland Island (C. G.
Rogers). Probably it occurs throughout the Archipelago.
The only adult (J skull known to me, that of an old specimen,
differs from the skulls of other races of Paguma by having
the postorbital area relatively deeply constricted and the
postorbital processes salient. These features, associated
with a high sagittal crest, combine to give it a resemblance
to the skull of Paradoxurus. But the specimen came from
the Zoological Society, and it is almost certain that the peculi-
arities are due to rearing in captivity. The old $ skull
from Rutland Island has the postorbital area parallel-sided
and the postorbital processes small, as in typical Paguma ;
the sagittal area is narrowly lyrate and the muzzle is excep-
tionally wide outside the first upper molars.
4:7 f. Paguma larvata intrudens Wroughton.
Paguma larvata intrudens, Wroughton, Journ. Bomb. Nat. Hist.
Soe. xix, p. 739, 1910, and xx, p. SI, 1918 ; G. M. Allen,
Amer. Mus. Novit. no. 359, p. 7, 1929 ; Osgood, Field Mus.
Nat. Hist., Zool. xviii, p. 259, 1932 ; Pocock, Joxirn. Bomb.
Nat. Hist. Soc. xxxvii, p. 340, fig. 6, C, 1934 ; id., Proo. Zool.
Soe. 1934, p. 670.
Paguma larvata vagans, Kloss, Journ. Nat. Hist. Soc. Siam,
iii, p. 73, 1919.
Paguma larvata yunalw, Thomas, Ann. Mag. Nat. Hist. (9) viii,
p. 617, 1921.
Locality of the type of intrudens, Sima in Myitkyina,
Upper Burma ; of vagans, Sikawtur, north-west of Raheng,
W. Siam ; ofyunalis, Yenyuensien in S. Szechwan.
Distribution. — From S. Szechwan, Yunnan, andN.B. Btjema
to the Shan States, Siam, Laos, and Tong-king.
Distinguished from the races previously described by the
very sharply contrasted black and white pattern of the mask,
the median band being bordered throughout by black and
passing over the nape as a white stripe which may reach
the shoulders or surpass them. Also by the contour hairs of
the upper side behind the shoulders being at most a little
blackened at the tip, so that the general hue is paler, varying
from buffish-grey to foxy-red.
426 vrvBBBiD^:.
The flesh-measurements (in English inches) and weights
(in lb.), where known, of three specimens from British
Indian territory are as follows : —
Head and Hind
Locality and sex. body. Tail. foot. Weight.
Myitkyina (type) ; yg. ad. <j> . . 24 23 3& —
N. Shan States ; yg. ad. <? . . 27 £ 24 1 4| 11
N. Shan States ; imm. $ 22£ 22- 4+ 6 §
This race is very closely related to typical larvata from
S. China, from which it only differs in being a trifle larger,
the condylobasal length of the skull ranging from 116 to
121 mm., whereas in the typical form it is up to about 112 mm.
The characters upon which yunalis from S. Szechwan and
vagans from W. Siam were based have been shown to be
too variable for the admission of the races.
Up to the present time the only localities within our limits
where this race has been collected are Sima in Myitkyina,
KE. Upper Burma (A. W. Kemmis), and Pyaunggaung in
the North Shan States, 2,794 ft. (G. C. Shortridge) ; but it
has also been recorded from Tengyueh, 9,000 ft., in Yunnan,
from the Lichiang Eange, Yunnan, 11,000-12,000 ft., from
Yenyuensien in S- Szechwan, and from Laos and Tong-king in
Indo-China (Delacour and Lowe).
47 g. Paguma larvata rolmsta (Miller).
Paradoxurus robustus, Miller, Pioe. Biol. Soc. Wash, xix, p. 26,
1906.
Paguma larvata robusta, Pocock, Journ. Bomb. Nat. Hist. Soe.
xxxvii, p. 343, 1934 ; id., Proc. Zool. Soc. 1934, p. 674, text-
fig. 10 (skull), pi. 2, D (head).
Locality of the type, Trang in Peninsular Siam.
Distribution. — Tenassebim, Peninsular Siam, northern Malay
Peninsula.
Distinguished from all the preceding races by having the
curved preaural crest of hairs reaching beyond the middle
of the cheek and nearer to the eye than to the ear. It is
also, on the average, slightly larger, and has the coat uniformly
short and thin throughout the year. The general colour,
hardly differing from that of some examples of grayi or
tytlerii, varies from paler or darker tawny to buffy-grey,
the back sometimes showing rusty-oehreous speckling and the
nape may be brownish ; the mask has its dark areas brown,
the pale patch below the eye is an indistinct grey smear,
and the conspicuous median band may stop short on the crown
or reach the nape, and is typically confluent with the white
of the cheek.
PAGUMA. 427
The flesh-measurements (in English inches) of two examples
are as follows : —
Head and Hind
Locality and sex. body. Tail. foot.
Tenasserim (Miller) ; ad. <J . . 25$ 22$ 4
Trang (Brit. Mus.) ; ad. $ . . . 30+ 21f 4+
The evidence that this race comes into the fauna of British
India rests upon the record by Miller of the adult (J, of which
the measurements are give above, which was collected at
Telok Besar in Tenasserim. This specimen I have not seen,
but the above-given description is based upon five examples
collected by Boden Kloss in Peninsular Siam and the northern
part of the Malay Peninsula, the measured example from
Trang being a topotype.
The skull is well developed, with a tolerably high sagittal
crest in the adult and a noticeable depression on the crown
close to the crest where the postorbital area passes into the
expanded portion of the cranium. It is larger than in the
subspecies described above, the condylobasal length in the
adult <J varying from 126 to 135 mm., and in the $ from
120 to about 126 mm.
47 h. Paguma larvata janetta Thomas.
Paguma robusta, Wroughton, Journ. Bomb. Nat. Hist. Soo. xxiii,
p. 710, 1915 (not P. robusta Miller).
Paguma leucomystaz janetta, Thomas, Ann. Mag. Nat. Hist. (10)
ii, p. 101, 1928.
Paguma larvata janetta, Poeock, Journ. Bomb. Nat. Hist. Soo.
xxxvii, p. 344, 1934, fig. 6, D (head), fig. 7 (skull) ; id , Proo.
Zool. Soc. 1934, p. 673.
Locality of the type, Bankachon, Victoria Point, S . Tenasserim.
Distribution. — S. Tbnassbeim.
Resembling robusta in the situation of the crest on the cheek,
in its short, thinnish, winter coat, and in cranial characters,
but distinguished by its much darker colour, the back being
closely speckled with black and ochreous or buff, the pale
speckling passing into grey on the flanks but dying out on
tne shoulders and nape, which are brownish-black with at
most some white continued back from the crown in the middle
of the nape. The facial pattern varies. In the type, adult $,
there is no pale patch below the eye, the grey fronto-nasal
band is narrowly continuous with the large pale preaural
area, and there is no white on the crown or nape. In an
immature <J the mask is more emphasized, black and white,
the median band is more separated from the preaural area,
there is some white on the crown and a good deal on the nape.
An adult $ has a small patch below the eye, and further differs
in having the crest on the cheek nearer the ear, thus approaching
428
vrvERnnxa:.
the condition seen in intrudens and other more northern races ;
but possibly this is due to the make-up of the skin.
The flesh-measurements (in English inches) and the weights
(in lb.) of an adult £ and $ from Bankachon (G. C. Shortridge)
are as follows : —
Head and Hind
body. Tail. foot. Weight.
Ad. S (type) 26+ — 4-} 12i
Ad. $ 25J 23 4 9£
B A
Fig. 99.
A. Upper side of skull of the type of Parjuma larvata ianetta, an old A,
from Bankachon, Tenasssrim.
B. Lower side of the same. Both figures x J.
These dimensions agree tolerably closely with those of
Miller's example of robusta from Tenasserim ; but the type of
the latter, an adult $ from Trang, is as large as the adult $
(type) of janetta and two adult <J examples of robusta, namely
™f ow from Trang, above measured, and another from
Klong Wang Hip in the Malay Peninsula, which is 29i in
m head and body, are a good deal larger. These facts suggest
that typical robusta is, on the average, a larger race, and that
the Tenasserim form of it links robusta with janetta, at all events
PAGTJMA.
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430 VIVBRBID^.
in dimensions. The differences in colour between robusta and
janetta probably indicate a wetter environment for the latter.
The skull has the same depression on the crown at the
junction of the frontal and parietal bones as in rohusta.
It is more muscularly developed than in the northern races
of Paguma, the sagittal crest in the $ being high, adjoining
the depression. In the $ there is no crest, the temporal
ridges being about 5 mm. apart.
Habits. — The habits of Paguma are very similar to those of
Paradoxwrus. According to Hodgson the race grayi, found in
Nepal, where it inhabits the mountain forests, is more frugi-
vorous ; but it is omnivorous, feeding partly on vegetable, partly
on animal food, including birds and small mammals. The
dwarf Andamanese race is said to be very destructive to
pineapple plantations, and Mackenzie was told that Tieglecta,
the race found in the Chin Hills, where it is eaten by the
natives, who call it " Sa Gyaw," feeds on fruits and roots.
Shortridge found janetta very plentiful inland from Victoria
Point, Tenasserim, where it is a " great ratter," but is seldom
destructive to poultry.
Apparently the only information about the breeding habits
in the wild is supplied by Hodgson's statement that in
Nepal this Pahn-Civet breeds in holes in trees and that a
litter of four young was found on one occasion. But a pair
of the typical race (P. larvata larvata) from Szechwan, which
bred in the Zoological Gardens, as I recorded (Proc. Zool. Soc.
1911, p. 621), produced a Utter of three in the early summer.
In colour they differed from their parents in having the
" mask " less emphasized and the general hue of the body
greyer, with the underside white. They also showed on the
back and sides faint traces of the pattern characteristic of
Paradoxurus. Their eyes, at first closed, opened on the ninth
day after birth. The growth of the young was much more
rapid than in kittens and puppies. When only three months'
old they almost equalled their parents in size. We also owe
to Hodgson the evidence that this animal, like Paradoxurus,
is protected by the nauseous secretion of its anal glands.
A tame specimen, he said, was " very cleanly and its body
emitted no unpleasant smell, though when it was irritated it
exhaled a most fetid stench, caused by the discharge of a thin
yellow fluid from pores ... on each side of the anal orifice."
Nearly all the races have conspicuously marked heads, and this
" mask " reaches the maximum of contrast in black and white
in typical larvata from China and its near ally intrudens of
Burma. It is significant that it is least conspicuous in the
race from the Andamans, where there are practically no
carnivorous enemies to be " warned off."
ABCTITIS. 431
Genus ARCTICTIS Temminek.
Arctictis, Temininck, Mon. Mamm. i, Tabl. Method, p. 21, 1824 ;
and of all recent authors, including Blanford, Mamm. Brit. Ind.
p. 117, 1888 ; Pocock, Proc. Zool. Soc. 1915, pp. 387-412, and
1933, p. 1015.
Ictides, F. Cuvier, Dents des Mamm. p. 252, 1824 ; Valenciennes.
Ann. Sci. Nat. iv, p. 57, 1825 *.
Type of Arctictis and of Ictides, binkurong.
Distribution. — The Eastern Himalayas, Assam, Bttema,
Indo-China, Siam, Malay Peninsula, Sumatra, Java, Borneo,
and Palawan.
Essentially like Paradoxurus and Paguma in general build
but more massive, in the length of the tail and legs, and in
the structure of the scent-glands f and of the feet, except
that the hind foot is naked throughout below, instead of having
the heel hairy. The contour hairs of the coat also are much
longer and coarser, and the long hairs clothing the whole of
the back of the ears project beyond the tip as a definite
tuft %. The rhinarium is very large and more convex above
than in Paradoxurus and Paguma, the median groove being
much narrower above the philtrum. The ear is as in the other
genera, except that the anterior flap of the bursa is more
widely and less deeply emarginate. The tail is more muscular,
especially at the base, and is prehensile at the tip, although
hairy throughout §. There are two pairs of mammae.
The skull in its main features is like that of Paguma, having
a similar extension of the palate over the anterior half of the
mesopterygoid fossa and a long, wide, little-constricted
postorbital area, but this area and the area between the
postorbital processes is more inflated by air-cells ; the extent
of the inflation is, however, very variable individually. It
differs from the skulls both of Paguma and Paradoxurus in
having the palate less flat ; it is a little elevated on each side
alongside the larger crushing cheek-teeth and grooved between
these ridges. In this respect the skull approaches that of
Arctogalidia (see p. 441). There is also a resemblance
between the teeth of these two genera, but those of Arctictis
are more decadent and differ a good deal from those of Paguma y
* These two names date from the same year ; and, not knowing
which was published first, I adopt Arctictis, which has been in common,
use for about a century.
f In 1915, writing from memory, I wrongly stated that the ? genera-
tive orifice is in front of the glandular area. It is encircled by it, as
stated by Carlsson.
{ This tuft does not arise from the tip of the ear like that of a Lynx,
as has been stated.
§ The only other Carnivore which has a truly prehensile tail is the-
tropical American Kinkajou (Potos flavus), a member of the Proeyo-
nidffi, which closely resembles the Binturong in habits.
432
VIVERBIDiE.
being much less trenchant and less massive. The upper
incisors are separated from each other and form a tolerably
strongly curved line ; the upper canine is very large and sharp,
and has a sharp cutting hinder edge ; the upper carnassial
(#m 4 ) and the first upper molar have the inner lobe (protocone)
more -widely rounded than in Paguma, and almost as wide
basally as the outer part of the crown, which in pm* has the
posterior cusp very small and hardly larger than the anterior
cusp (parastyle) ; the last upper molar is sometimes absent,
and in the lower jaw the first premolar is usually absent, and
minute when present.
The genus is represented by a single species.
Kg. 100.
A. Skull of adult, probably 2, specimen of Arctictis binturong aibijrons,
from Sikkim.
B. Left half of underside of the same, pm 1 , pm i , and m\ first and fourth
premolars and first molar.
48. Aretietis Wnturong Baffles. The Binturong or Bear-Cat.
(For principal bibliographical references see under the subspecific
Main characters and distribution as under the genus. The
tail is a little shorter than the head and body, and from about
five to six times the length of the hind foot. Colour of the
body very variable, generally mostly black, but ■ more or
ABCTICTIS. 433
fewer of the hairs frequently partly whitish or buff, giving
a speckled appearance to the pelage, sometimes so extensively
pale that the whole body is mostly straw-coloured or grey,
the young being often at all events paler than the adults,
but the head is always closely speckled with grey or buff;
the long mystacial vibrissse are conspicuously white, and there
is a white rim on the summit of the otherwise black ear.
The limbs are typically speckled externally ; the tail is
generally like the body, but commonly paler at the base
beneath, and the glandular area, at least in the <£, is whitish.
Tentatively the British Indian Binturongs are assigned
to two subspecies, a northern and a southern ; but, owing to-
the rarity of the animal, the available material is too scanty
to justify more than the provisional conclusion that the two
differ in size, the northern being the smaller of the two.
48 a. Arctictis binturong albifrons (Cuvier).
Paradoxurus albifrons, F. Cuvier, Mem. Mus. d'Hist. Nat. ix,
pp. 44-8, pi. 8, 1822 *.
Ictides albifrons, F. Cuvier & Geoflroy, Hist. Nat. Mamm. iii,
no. 201, 1824.
Arctictis binturong albifrons, Pocoek, Proe. Zool. Soc. 1933,
p. 1030, figs, of skull, pp. 1024 and 1027.
* There has been great confusion in the application of this name,
which was given by Cuvier in 1822 to a sketch sent to him by Duvaucel
of a specimen living in the Barraekpore Menagerie, Calcutta, which
Duvaucel ascertained had been captured in Bhutan. This sketch
■was reproduced in colour by Cuvier and Geoffroy two years later.
The truth of Duvaucel's account of the specimen has been doubted
and even denied, but, in my opinion, without warrant. Temminek
thought the specimen sketched by Duvaucel was one captured in Malacca
and described by Raffles in 1 822 as Viverra binturong. It is admittedly
possible that the owner took it to Calcutta and deposited it in the
menagerie after Raffles had seen it, and also that the animal changed
from black to grey in the interval. But this opinion leaves unexplained
Duvaucel's statement, no doubt derived from the Superintendent at
Barraekpore, that the animal came from Bhutan. Cuvier himself
was partly responsible for further doubts about the matter, because
in 1824, in conjunction with Valenciennes, he identified as albifrons
an example sent to him from Java ; and this was apparently the-
source of Oustalet's statement (Bull. Mus. d'Hist. Nat. vii, p. 318, 1901)
that between 1820 and 1826 Diard and Duvaucel sent the type of
albifrons to Cuvier from Java. Accepting this view, Kloss (Treub.
Buitenz. x, p. 497, 1929) adopted albifrons for the Binturong of that
island. But the original description of albifrons was taken from a drawing,
not from a specimen ; and since Duvaucel is known to have sent to Cuvier
from Assam descriptions and sketches of purely Indian animals, e. g.,
Melursus ursinus, there is no reason to doubt he was in Northern India,
and may very well have seen the Bhutan Binturong in Barraekpore,
as he stated. The matter is of some importance because Blanford
doubted the existence of the Binturong in the Himalayas, although
admitting its existence in Assam, and did not consider the original
history of the alleged Bhutan specimen described as albifrons.
VOL. T. 2l
434 VTVEBEIDiB.
Vernacular. — Young (Assamese) ; Myouh-Kyd (Burmese).
Locality of the type, Bhutan.
Distribution. — Nepal, Stkktm, Bhutan, Assam *, Uppeb
BtnEMA, and Tong-king.
Coat very long and shaggy and thickened in winter with
abundance of underwool ; colour varying from dominantly
tawny or grey to jet black, with comparatively few pale
annulations on the hairs. Skull small and lightly built.
The figure of the type of this race shows the coat to have
been unusually long and wavy, and, with the exception of
most of the tail, the ears, and the toes, which are blackish,
the colour was mainly pale grey, with some dark streaks
where the contour hairs were parted, the forehead being
•a little lighter and the muzzle a little darker than the body.
An unmeasured skin from Assam has the general colour
.streaked tawny and black in about equal proportions, the hairs
.having long, greyish, or foxy-Ted annulations, with evidence,
especially on the rump, flanks, and the base of the tail, of new
.grey-tipped hairs erupting at the roots of the long old hairs,
but there is no appreciable amount of underwool ; the short
.hairs of the face and limbs are similarly tipped, these parts
being practically wholly tawny owing to the concealment
of the black bases of the hairs by the pale tips ; on the upper
.side of the tail the pale annulations gradually disappear
distally, the end being black, but the lower side of the tail
and of the body are tawny. The coat is loose, rather harsh
and very long, the hairs on the rump being up to about
110 mm. (over 4 in.), and on the shoulders 65 mm. (over 2J in.).
This skin, apparently still carrying its long winter coat,
although the wool is moulted, agrees tolerably closely with the
figure and description of the type of albifrons from Bhutan,
except that its general hue is tawnier.
A skin from Indawgyi Lake in Myitkyina, Upper Burma
(Capt. J. H. Whitehead), dated February 2, is in full winter
coat, and differs strikingly from the Assamese skin by being
mainly black and by the presence of abundant long under-
wool ; but the contour hairs of the coat are about the same
length ; the shoulders, back, and upper side of the tail are
wholly black, the hairs being unspeckled throughout their
length, but there is some buff speckling on the rump and the
top and sides of the head, and the outside of the fore and
hind legs, except for the black toes, are finely speckled with
grey ; the underside from the throat to the belly is black,
and the underside of the tail is the same, except that the hairs
at its base are grey close to the skin.
* No specimens from these Indian States have been recorded for
many years. Gray, apparently on Blyth's authority, stated that the
jspeoies occurs in Nepal ; but Hodgson did not secure it.
AKCTIOTIS. 435
The only other specimen, I have seen assignable to this
xace is an immature $ from Pouine, Tong-kmg (Delacour
and Lowe), September 19. This is almost a duplicate of the
skin from Myitkyina in colour, but the pale speckling on the
head and nape is a trifle more buffy and on the nape more
profuse, and there is no speckling on the rump, only on the
thighs and feet. The coat is very soft, with a tolerable
amount of underwool, the long hairs being 60 mm. on the
rump and about 50 on the shoulders. The difference between
this skin and the one from Myitkyina in the length and
luxuriance of the coat is no doubt largely seasonal, but partly
due to the age of the specimens.
The only skull of a British Indian specimen I have seen is
that of an adult, probably $, without skin, ticketed " Sikkim "
(Ei. Lydekker). It is comparatively small, with the muzzle
narrow ; the temporal ridges are strong, 11 mm. apart at the
postorbital constriction and nearly parallel to the occiput
behind it ; the postorbital area is very little inflated
dorsally and laterally, the sides being approximately parallel,
with a shallow posterior constriction, and the dorsal profile
is less steeply sloped anteriorly and posteriorly from its highest
point than in any skull of the genus I have seen. It disagrees
entirely with the statement of J. A. Allen (Bull. Amer. Mus.
Nat. Hist, xxviii, p. 15, 1910) that " Indian " skulls are larger
than Sumatran skulls, but resemble them in the elevation
and inflation of the frontal region. In its comparatively
small size and low crown it resembles skulls from Palawan,
Philippine Islands, described by Allen as Arctictis whitei
(=Arctictis binturong whitei) ; but at least differs from them
in having the inner lobes of pm 4, and m x narrowed and not
widely rounded.
48 b. Arctictis binturong binturong (Raffles).
Viverra ? binturong, Raffles, Tr. Linn. Soo. xiii, p. 253, 1822.
Ictides ater, F. Cuvier & Geoffroy, Hist. Nat. Mamn, iii, no. 202,
1824.
Arctictis gcdrd.ne.ri, Thomas, Ann. Mag. Nat. Hist. (8) xvii, p. 270,
1916.
Arctictis binturong binturong, Kloss, Journ. Fed. Mai. St. Mus. vii,
p. 293, 1917 ; Pooock, Proc. Zool. Soo. 1933, p. 1030.
Vernacular. — Untarong (Malay) ; Beruong (Malay at Perak).
Locality of the type of binturong and ater, Malacca ; of
■gairdneri, Sai Yoke, S.W. Siam.
Distribution. — S. Siam, Tenassbeim, Malay Peninsula, and
Sumatra.
Distinguished from the northern race, albifrons, by its
shorter, much less luxuriant winter coat, and apparently
by its more consistently blackish colour, the long hairs never
2i2
436
VIVEKEID^!.
being so extensively and profusely annulated as in the examples
of albifrons from Bhutan and Assam. The skull also, judging
from the available material, is considerably longer, more
robust, and more inflated.
Baffles's type of binturong, a living specimen seen in Malacca,
was black, except on the face and legs, which were brown.
Cuvier's type of ater, a coloured sketch, by Duvaucel of a
specimen also seen in Malacca, was very similar, being altogether
black, with the upper side of the muzzle yellowish, and some
white hairs on the forehead and legs. Kloss described
specimens from near Raheng in Siam and from Patani in
Peninsular Siam which were also mainly black, but speckled
slightly and to a varying extent with grey or buff on the
back. Of Malayan specimens in the British Museum the
4
i«
Fig. 101.-
-The Binturong, Arciiotis binturong binturong.
Blanford.)
(From
most interesting is an adult $ from Ulu Ifok, Perak (Vernay's
coll.), February 29, killed in the same month as the example
of albifrons from Myitkyina, but differing from it mainly
in having the wool of the coat very short and scanty and the
hairs on the rump much shorter, only 85 mm., although those
on the shoulder are about 75 mm. The colour is nearly the
same, but the crown between the ears is richer ochreous-buff,
the fore leg is more speckled, but the hind leg and thigh less.
No importance is attached to the slight difference in tint,
but the difference in the coat is very marked, and at once
perceptible to eye and touch.
The only skin from British Indian territory I have seen is
one procured by Shortridge from a native near Tenasserim
Village. Itcloselyresemblesincolourtheskins from Myitkyina
Head and
Hind
body.
Tail.
foot.
30f
26
4
29
25
**
ABCTICTIS. -437
and Perak, but is rather more speckled on the fore back and
flanks.
An adult $ from Sanderam Agong, 2,450 ft., in Sumatra,
collected by Kloss and Eobinson, attests, as they claimed,
the extension of this race to that island.
The following are the flesh-measurements (in English
inches) of two of the specimens above referred to : —
Locality and sex.
Ulu Ifok, Perak ; ad. <J
Sanderam Agong, Sumatra ; ad. $
The weights of an adult <J from Aru Bay, an old <J from
Pulo Payong, and a young adult $ from Sungei Mandan, all
in Sumatra, were recorded by Lyon as 25J, 20, and 23 lb.
respectively.
Of the skulls recorded in the table on p. 438, that of the $
from Sanderam Agong may be compared with the skull of
albifrons, also no doubt a °, from Sikkim. The two differ
greatly. The Sumatran skull is considerably longer, although
the facial portion in front of the postorbital processes, including
the nasal bones, is about the same ; but the nasals and the
muzzle are much wider. There is little difference in the
width of the postorbital area at its narrowest point, which
is remote from the postorbital processes, but in front of the
constriction this area both laterally and above in the Sumatran
skull is strongly inflated with air-cells, so that it is markedly
swollen at the sides and above and is much more steeply
sloped anteriorly and posteriorly from its highest point
a little distance behind the level of the postorbital processes.
Some of these differences may prove to be of subspecific
value when the skull of albifrons is better known. But
skulls of the typical southern race, binturong, differ profoundly
in shape in the same country owing to the great variation
in the extent to which the postorbital area is inflated with
air-cells*. These are most highly developed in the old <J
skull from Sai Yoke in S.W. Siam, the largest recorded skull
of Arctictis, which Thomas regarded as representing a distinct
species, gairdneri. But I agree with Kloss that this is nothing
but an unusually large skull of A. binturong binturong. All its
peculiarities are foreshadowed in the $ skull from Sanderam
Agong in Sumatra.
Habits. — The Binturong lives in dense forests, is mainly
arboreal and nocturnal, and probably nowhere abundant. Its
* This, I believe, was first pointed out by Lyon in connection with
Sumatran skulls (Proc. U.S. Nat. Mus. xxxiv, p. 651, 1908).
438
VIVEKBTIXE.
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ABOTOGAUDim-^i. 439
habitat accounts for the infrequency with which it is seen, let
alone shot or trapped, by Europeans. The only specimens
secured by the collectors for the Mammal Survey of British
India were the two skins from near Tenasserim Village, bought
from natives by Shortridge, who reported that the species
was not at all common, and was quite unknown to many of
the inhabitants. The animal, however, has been frequently
exhibited in captivity, and I observed the behaviour of several
in the London Zoological Gardens. When resting they lie,
as a rule, curled up, with the head tucked under the tail.
They never leap, but climb skilfully, albeit slowly, progressing
with equal ease and confidence along the upper side of branches
or, upside down, beneath them, the prehensile tail being
always in readiness as a help, and they descend the vertical
bars of the cage head first, gripping them between their
paws and using the prehensile tafl. as a check. They feed
upon bread and milk, rice, bananas, and other fruits, as well
as upon eggs and fowls' heads. When irritated they growl
fiercely, and when on the prowl may periodically utter a series
of low grunts or a hissing sound made by expelling the air
through partially opened lips. In the forest the animal is
said to howl loudly. This sound is probably a sexual call, if the
story be true. The natural diet, Blanford says, on whose
authority I know not, consists of small mammals, birds,
fishes, earthworms, insects, and fruits ; but it is probably
mainly of a vegetable nature, since the Binturong has not the
attributes of a predatory mammal, and fishes and earthworms,
it seems, can form but an unimportant item in the food of
an essentially arboreal species which is neither aquatic nor
fossorial.
Subfamily ARCTOGALIDIDWE.
Arctogalidiinee, Pooock, Proo. Zool. Soo. 1933, p. 977.
Distinguished from the preceding subfamilies of Viverridse
by the absence in the male of the perfume gland, the long
perineal area between the scrotum and the prepuce being
continuously hairy, and by the position of the gland in the
female in front of the vulva and clitoris, where it is represented
externally by two low ridges of naked skin diverging anteriorly,
uniting in front, capable of being folded over to meet in the
middle line to form a closed pouch for the secretion, and con-
tinuous behind with a rim of naked skin surrounding the
vulva.
Feet essentially as in the Paradoxurinse, scansorial and
semiplantigrade, but with the pads of the third and fourth
440
VrVBBEID^!.
■digits of the hind foot unfused, the carpal pads of the fore
foot relatively longer and narrower, and the surface of all
the pads smooth, not granular or papillate ; the heel of the
hind foot hairy.
"When I described and figured in 1915 the external features
•of a $ Arctogalidia, no male example was available for com-
parison, and I kept the genus associated with Paradoxurus,
Fig. 102.
A. Lower side of right hind foot of Arctogalidia trivirgata from Borneo
m, inner metatarsal pad.
B. Lower side of right fore foot of the same, c, external carpal pad.
C. Anal and genital area of the same (?). a, anus ; v, vulva • gl
expanded glandular pouch in front of vulva. ' '
D. Glandular poueh {gl), and vulva (v), with clitoris in front of it.
E. The glandular poueh, closed.
as had always been the custom ; but I have since seen many
well-made skins of the male, and can confirm Blanford's
statement that " there is no bald space in front of the scrotum
or round the genital orifice."
ABCTOUALIDIA. 441
Genus ARCTOGALIDIA Merriam.
Arctogale, Gray, Proe. Zool. Soo. 1864, p. 542 ; and of subsequent
authors, including Blanford, until 1897 (not Arctogale Kaup,
1829).
Aretogalidia, Merriam, Science, v, p. 302, 1897 ; and recent authors,
including Poeoek, Proe. Zool. Soc. 1915, pp. 390-412, and 1933,
p. 977.
Type of the genus, Paradoxtirus trivirgatus Gray.
Distribution. — Assam, Burma, Laos, Tong-king, the Malay-
Peninsula, and thence eastwards to Borneo.
Closely resembling Paradoxurus in external form and in the
length of the legs and of the tail. The rhinarium also is very
similar, being deeply grooved both in front and above, but
it has a wider angular emargination above and the infranarial
portions are not so deep. The ear differs in its less well-
developed bursa, M-hieh has the edge of its posterior flap
continuous above and below with the margin of the pinna,
the anterior flap less emarginate and not produced into
a prominence above and below. In addition to the characters
connected with the feet mentioned above under the subfamily,
the fore foot is narrower and longer than in Paradoxurus,
the plantar and carpal pads combined being much longer than
wide, and the distal margins of the four elements of the
plantar pad form a much less widely curved line ; the area
between the plantar and digital pads is naked. The pattern,
when present, consists of three dark dorsal stripes and of
a white streak on the nose. There are two pairs of abdominal
teats.
The skull differs in many respects from that of Paradoxurus.
It is wider in its mastoid region, has less robust but more
salient zygomatic arches, much longer postorbital processes,
and the muzzle longer and wider in front above the canines ;
the palate is not flat, but has an undulating surface owing to
its convexity between the carnassials and first molars of each
side ; behind the last molar it is produced into a broad plate
defined by a deep notch on the inner side ; the mesoperygoid
fossa is long and narrow, and roofed in its anterior half by
a backward extension of the palate ; the bullae are flatter
and the divisional line between their two component bones is
early obliterated. The dentition too is different, the back
teeth being smaller, more crushing, and less trenchant in type-
The main cusp of the first three upper premolars is high
and pointed, but the crowns as a whole are not compressed,
that of pm 3 having a distinct inner lobe. This tooth is only
a little smaller than the upper earnassial (pm 4 ), which has the
inner lobe (protocone) nearly as wide basally as the outer
portion of the crown, which is three-cusped, with a well-
developed parastyle in front, a larger paracone in the middle,
and a sharp cuspidate, not blade-like, metacone behind.
442
VIVEBBID.2E.
The first molar (m 1 ) is about the same size as pm i and is
tolerably like it, but has two outer cusps (paracone and
metaoone) and an inner lobe (protocone) as long at the base
as the crown. In the lower jaw the first three premolars
have a high, piercing main cusp, the fourth has three cusps
in front and a well-developed " heel " behind, the lower
carnassial (m x ) is larger than pm, v and has the normal three
anterior cusps and a large cuspidate heel, and the last molar
Fig. 103.
A. Left half of lower side of skull of Arctogalidia trivirgata. as, anterior
orifice of alisphenoid canal ; o, foramen ovale ; c, carotid foramen.
B. Bight half of lower jaw of the same, showing the upper view of the-
teeth. (From Blanford).
which is four-cusped and irregularly oval, is inserted externally,
so that its inner edge is out of line with the inner edge of the
first molar. The inner margin of the mandible adjoining
this tooth is usually thickened into a crest-like ridge which
ends in a low angle behind the level of the last molar.
AEOTOGAUDIA. 443
The shape of the palate in this genus is an exaggeration
of the condition seen in Arctictis. The reduction in the size
of the back teeth is also a point of resemblance between
them. But these likenesses are no doubt adaptive and not
due to affinity, considering the differences between the two
genera in external and most cranial characters.
Probably no Oriental genus of Carnivora abounds in so
many nominal forms, mainly insular, to which full specific
rank has been given on the evidence of single specimens,
as Arctogalidia. In my paper above quoted I tentatively
reduced the species to two, namely, A. trivirgata Gray (Proc.
Zool. Soc 1832, p. 67, and 1864, p. 543), which ranges from
the Malay Peninsula to Borneo, and is represented by several
local races, many of them found in the smaller islands, and
A. leucotis, which ranges from Assam and Indo-China to
Tenasserim and the Mergui Archipelago, and is the only one
represented in British India. Since the differences between
them, however, are comparatively slight, and there is evidence
of intergradation between them on the mainland of Asia,
where their distribution is continuous, I now adopt the view
of Blanford that the mainland forms, at all events, may be
regarded as representing a single species. Further research
may show that some of the insular forms, falling outside the
limits of the British Indian fauna, are worth specific status.
In the earliest described form, A. trivirgata, found in the
Malay Peninsula *, the back of the ear is clothed with black
hairs and has its integument pigmented black throughout.
In the type of A. leucotis from Tenasserim, on the other hand,
the upper half of the back of the ear has the integument pink
and the hairs clothing it white. This character obtains
in all British Indian skins, which differ in this respect from
skins collected in numerous localities from the Malay Peninsula
to Borneo. But in an example from Baheng in Siam only
the upper fourth of the back of the ear is white. It is thus
intermediate between typical leucotis and typical trivirgata
in this respect. The most northern locality in the Malay
Peninsula whence the black- eared form has been collected
is Trang in Peninsular Siam. A specimen from this spot
was named A. major by Miller (Proc. Biol. Soc. Wash, xix,
p. 25, 1906), and the locality is very near the southern
limit of the known range of the white-eared form.
* The type, in the Leyden Museum, was recorded by Gray from the
" Moluccas," where the species does not occur. This mistake was
probably due to a misprint for "Malacca," or to Gray's misreading
of the locality entered on the label. At all events his description exactly
fits skins from the Malay Peninsula, and Gray himself in 1843 and later
assigned the name trivirgata to specimens in the British Museum
from Malacca and Singapore.
444 VIVERRIM3.
49, Arctogalidia trivirgata (Gray). The Three-banded Palm-
Civet.
Paradoxurus trivirgatus, Gray, Proo. Zool. Soc. 1832, p. 67.
Paguma trivirgata Gray, List Mamm. Brit. Mus. p. 55, 1843.
Arctogale trivirgata, Gray, Proo. Zool. Soc. 1864, p. 543 ; id., Cat.
Cam. Brit. Mus. 1869, p. 75.
(For bibliography and synonymy of the British Indian races see
under the subspecies).
Since only one species is here admitted, its characters and
distribution are as entered under the generic heading.
The following three local races, all with white tips to the
back of the ears, are represented in the British India fauna : —
<z. Size larger, with the three dorsal stripes wide, [p. 447.
black, and equally conspicuous millsi Wrought.,
a'. Size smaller, the three dorsal stripes narrower,
the laterals typically less conspicuous than
the median and not infrequently broken up
into spots. [p. 444.
b. Size larger leucotis (Horsf.),
b'. Size smaller macro, Miller, p. 446.
49 a. Arctogalidia trivirgata leueotis (Horsfield).
Paradoxurus leucotis, Horsfield, Cat. Mamm. E. I. Co. p. 66,
1852 ; Blyth, Journ. As. Soc. Beng. xxvii, p. 274, 1858.
Paradoxurus prehensilis, Sclater, Proc. Zool. Soc. 1877, p. 681,
pi. 71 (not Viverra prehensilis Desm.).
Arctogale leucotis, Blanford, Proc. Zool. Soc. 1885, p. 789 ; id.,
Mamm. Brit. Ind. p. 115, 1888 (in part).
Arctogahdia leucotis, Miller, Proc. Biol. Soc. Wash, xix, p. 25, 1906 ;
id,, Smiths. Misc. Coll. lxi, p. 6, 1913.
Arctogalidia leucotis leucotis, Pocock, Proc. Zool. Soc. 1933, p. 979.
Vernacular. — Kyoung-na-rwek-phyn (Arakan) ; Thit-Te-Kyee
(Burmese) ; Taw Hsee (Karen) ; Kyoung-na-ga (Tenasserim) ;
Musang Ahar (Malay).
Locality of the type of leucotis, Tenasserim ; of prehensilis,
Assoun Hounderaw River, 90 miles from MouLmein.
Distribution. — Abakan, S. Burma, W. Siam, throughout
Tenasserim, Mergui Archipelago, and some islands off
the north-west coast of the Malay Peninsula ; ? Malay Penin-
sula.
General colour of the nape, back, proximal portion of the
tail, and the outside of the limbs tawny, but varying individually
from dusky greyish-tawny to bright ochreous-buffy tawny ;
the head usually darker and greyer, but with no definite
brown area on the crown at the base of the ears ; the paws,
the distal portion of the tail to a varying extent, and the
muzzle brownish ; a median white streak on the muzzle
and front of the forehead ; the three dark blackish or brownish
stripes running backwards from behind the shoulders and
ARCTOGAUDIA.
445
sometimes traceable on the neck, the median complete and
distinct, the laterals usually more or less broken up into
spots, less clearly denned and occasionally almost obsolete ;
the proximal portion of the tail obscurely striped transversely ;
the underside mostly greyish- white or cream-buff, with a whiter,
creamy patch variable in size on the chest, and the hind
throat richer in tint than the fore throat.
The greatest contrast in colour in this race is shown by two
skins from Western Siam. One from Hat Sanuk, near
Fig. 104. — The White-eared Three-banded Palm-Civet, ArctogalitMa
trivirgata leucoUs, from Burma. (From Blanford.)
Koh Lak, Rajburi, April 15 (Robinson and Kloss),is brightish
ochreous-buff all over the upper side up to the head, and the
long hairs have an almost golden sheen ; the stripes are
brownish. The other, from the Mee Taw Forest, Raheng,
1,200 ft., April 22 (C. S. Barton), is ashy olivaceous-grey on
the back, only faintly tinged with buff on the flanks, the
pale area of the hairs being clear greyish- white, not golden ;
the pattern is blacker, and only about one-fourth, of the back
of the ear is white instead of half of it as in the Rajburi skin.
446 VIVERBIDiE.
The skins from British Indian territory are intermediate
between these two extremes, with the exception of an old <J
from King's Island, Mergui Archipelago (Primrose), which
very closely matches the skin from Raheng in its general
dusky hue. A young adult <J from Tenasserim Town, April 30
(G. C. Shortridge), which is tawny-buff, especially on the flanks
and the sides of the neck, with the head and cheeks dusky grey
and the lower side whitish-grey, comes nearest to the bright-
hued skin from Rajburi. Two others from Tenasserim Town,
March 7 and 14, are greyer, the three forming an interlinking
series between the two Siamese skins. Skins from Lower Burma
(J. M. D. Mackenzie), namely, from the Southern Zamayi
Reserve, 60 miles north of Pegu Town, March 12, and a pair
from 20 miles east of Toungoo, 3,000 ft., May 7, fit in with the
Tenasserim skins, except that those from Toungoo show
changes in the coat and colour from the moult. One is
bleached and paler than the Tenasserim series, and the other
has the rump and hind back denuded of contour hairs and
covered with pale wool, showing hardly a trace of pattern,
but there is a patch of new greyish-buff hair showing pronounced
black pattern on the fore back, with a good deal of buff on the
shoulders and sides of the neck, and the pattern is visible on
the nape.
49 h. Arctogalidia trivirgata maera Miller.
Arctogalidia macro, Miller, Smiths. Misc. Coll. bri, p. 6, 1913.
Arctogalidia leuootis rnacra, Pocock, Proc. Zool. Soc. 1933, p. 982.
Locality of the type, Domel Island in the Mergui Archi-
pelago.
Distribution. — Domel Island and possibly Langkawi and
Terutau off the north-west coast of the Malay Peninsula.
Distinguished from leucotis by its slightly smaller size and
shorter tail, the colour and pattern being apparently the same.
I have seen no example of this race from Domel Island.
The type and only certainly known example of it is a $
distinguished specifically by Miller from a single $ example
of leuootis from Tenasserim. I provisionally assign to macro,
two adult (J examples from Langkawi and Terutau (Robinson
and Kloss) which resemble the duller-tinted skins from
Tenasserim Town, but are smaller, smaller even in their
flesh-measurements than the type of macro,, although a little
larger in the skull. They are of interest as being the most
southerly representatives of the white-eared type of A. tri-
virgata, which has not been recorded actually from the Malay
Peninsula. Judging from the flesh- and skull-measurements
recorded on pp. 447 & 448, these three specimens suggest the
existence of a small southern insular race of the White-eared
Three-banded Palm-Civet.
ABCTOGAXIDIA. 447
Flesh-measurements (in, English inches) of specimens
assigned to these two races are as follows : —
Head and Hind
Name, locality, and sex. body Tail. foot.
leueotis.
Toungoo, Lower Burma ; ad. $ 21 — 22f 3f
Raheng, Siam ; ad. <J 20f 26f 3£+
Tenasserim Town ; yg. ad. <J . 19| 24* 3£—
Tenasserim Town ; ad. 9 20& 24f 3i+
Toungoo ; ad. $ 20£ 25f 4-
macra.
Domel Ial. (type) ; ad. $ 19 21f —
Langkawi Isl. ; ad. <$ 18J 20* 3|
Terutau Isl. ; ad. <J 17* 22- 3i
Two young adult $ specimens of leueotis from Tenasserim
Town weighed 4J and 4 lb respectively, and a gravid $ 4 J lb.
49 c. Aretogalidia trivirgata mills! Wroughton.
Arctogalidia millsi, Wroughton, Journ. Bomb. Nat. Hist. Soe.
xxvii, p. 599, 1921.
Arctogalidia leueotis rnillsi, Poeoek, Proc. Zool. Soo. 1933, p. 978.
Locality of the type, Mokokehung in the Naga Hills, Assam.
Distribution. — Assam, Laos, andTong-king.
Distinguished from leueotis by its larger size, judging from
the skull, and by the much more strongly emphasized dorsal
pattern, the three stripes being more intensely black and the
laterals to all intents and purposes as conspicuous as the
median and not broken up into spots on the back, although
on the nape they are more or less interrupted and narrower.
In the type the general hue of the back is dull tawny, like the
duller-tinted skins of leueotis ; but in a specimen from Xien
Quang Koo, Laos (Delacour and Lowe), which resembles
the type in pattern, the ground-colour is more richly tinted
with buff, especially on the flanks and sides of the neck.
These two unmeasured skins are the only representatives
of this race I have seen ; but there is no reason to doubt
that the specimens recorded as leueotis from Sylhet by Blyth,
from Assam by Sterndale, and from Tong-king by Osgood
belong to it.
Habits. — Owing to its being nocturnal and arboreal, and
living in dense forest, comparatively few observations have
been made on the mode of life of this Palm-Civet. The
specimens collected by Shortridge near Tenasserim Village,
by Mackenzie at Toungoo and in the Zamayi Reserve, and by
Barton at Raheng were killed in trees, the last after it had
bolted for safety into a hole in the trunk of a teak. It is an
exceedingly active climber. A tame specimen belonging to
44S
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AROTOGiULIDIA. 4A9
Sterndale would leap from branch, to branch of trees with
great activity, and was wonderfully agile amongst the ropes
on board ship. This specimen attested the omnivorous taste
of the species by its liking for most fruits, especially plantains,
and for milk ; but on one occasion it pounced on a pigeon and
would have killed it if allowed to do so. The Toungoo
specimens were eating fruit in a tree when shot, and the intes-
tines of the one from the Zamayi Reserve contained fruit and
some Ficus. On the other hand, the stomachs of the specimens
Shortridge examined contained remains of Squirrels. Inci-
dentally, he remarked in this connection that although to some
extent frugivorous, all the Palm-Civets live largely on Squirrels
and are a considerable factor in keeping down the numbers
of these rodents, which are so destructive to coconuts.
Shortridge 's discovery of a couple of foetuses in the $ from
Tenasserim Village on May 7 shows that there may be only
two young to the litter and that they are born in the early
summer. It is not unlikely that a second litter is produced
later in the year. From the behaviour of Barton's specimen,
above recorded, it seems probable that the young are reared
in hollow trees. Although these observations apply to
typical leucotis, there is no reason to doubt that the habits
of all the races are similar.
A representative of the dark-eared races of this Palm-Civet
which may turn up in Tenasserim is Arctogalidia trivirgata major,
described by Miller from Trang in Peninsular Siam (Proc.
Biol. Soc. Wash, xix, p. 25, 1906). According to the description,
of the one known specimen, this race differs from typical
trivirgata from the southern districts of the Malay Peninsula
by its larger size, much shorter tail, and by some details of
colour and pattern, the upper side being " light broccoli-
brown, with a silvery gloss," the crown and cheeks " clear
grizzled grey," the stripes strong and extending nearly to the
ears, and the tail marked with ten rings. In typical trivirgata
the general hue varies from deep brownish smoky-grey to
clear ashy- or tawny-grey, and the head is darker than the
back ; the stripes vary in distinctness and occasionally
extend over the nape, and the basal portion of the tail, when
pale, may show a few faint bands. The flesh-measurements
(in English inches) of Miller's type, an adult $, compared with,
those of the largest (J of typical trivirgata known to me, from
Singapore, are as follows : — Head and body 27§, 23§ ; tail 22f ,
25| ; hind foot 4— , 3|. Probably, however, there is a misprint
in the figures indicating the length of the tail in major, because
no other example of A. trivirgata known to me has the
tail shorter than the head and body. In the length of the
head and body major is larger than any of the flesh-measured
vol. I. 2q
450 VIVBEKID^!.
examples of the white-eared races found in British India.
Probably it was about the same size as millsi, although the
skull is a little shorter.
A species of the Viverridse briefly referred to by Blanford
•on account of the possibility of its future discovery in British
Indian territory was Gynogale bennettii, the Otter-Civet,
which is found in the southern part of the Malay Peninsula,
Sumatra, Java, and Borneo. It has not yet been found in
'Tenasserim ; but the possibility of the genus coming into the
British Indian fauna has been increased since Blanford's time
by the discovery by Delacour and Lowe of a second species,
C. lowei Pocock (Proc. Zool. Soc. 1933, p. 1034), in Tong-king.
This may turn up in Upper Burma. The genus, representing
a special subfamily, Cynogalinse, does not differ from other
Viverridas by having webbed feet, as has been frequently
stated, but very noticeably by its greatly expanded, prominent
muzzle, undivided upper lip, flattish rhinarium with dorsal
valvular nostrils, but no philtrum, its abundant bristle-like
facial vibrissse, and small ears, all connected with its semi-
a,quatic fish-eating habits ; also by its short tail, which is
about one-fourth the length of the head and body and less than
twice the length of the hind foot *.
Subfamily HEMIGALIN^}.
HemigaHna, Gray, Proe. Zool. Soo. 1864, p. 524.
HemigaUnse, Pooook, Aim.. Mag. Nat. Hist. (8) xvi, p. 349, 1915 ;
id., Proc. Zool. Soo. 1933, p. 999.
Resembling the Viverrinse in having the scent-glands
present in both sexes and wholly perineal, but differing by
their simpler structure, consisting in the (J of a shallower,
smaller pouch, with less tumid lips, situated mid- way between
the scrotum and the penis, but not extending to either, and
in the 5 of a pair of swellings, each with a slit-like orifice,
situated one on each side of the vagina and a little behind
it and on a common eminence, the perineal area behind
this eminence being naked. This description, based on a fresh
specimen, does not agree with Mivart's statement that in the
$ the gland extends from " near the anus to the vicinity of
the vagina." The prepuce is long and pendulous. Feet
nearly intermediate in structure between those of the digiti-
grade Viverrinse and the semiplantigrade Paradoxurinse,
but more like the latter, both the carpal and metatarsal pads
being well developed, double, and joining the plantar-pad
* See my papers in Ann. Mag. Nat. Hist. (8) xv, p. 351, 1915, and
Proe. Zool. Soo. 1933, p. 1031.
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HEMIGAMN^E.
451
below, and as wide as it is at the point of contact ; but the feet,
with the pads, are considerably narrower, the carpals and
metatarsals converging and meeting above so that a much
larger area of the under surface is hairy; the area between
Fig. 105.
A. Lower side of right hind foot of the Banded Palm-Civet, Hemigalus
derbyanus. m, inner metatarsal pad.
B. Lower side of right fore paw of the same, c, external carpal pad.
C & D. Front and side views of the rhinarium of the same.
E. Anal and genital areas of young (J of the same, a, anus ; gl, orifice
of glandular poueh ; p, tip of penis.
F. Anal and genital areas of adult § of the same, a, anus ; «, vulva ;
gl, orifice of gland on left side of vulva ; ag, anal gland of right
side dissected to show on its inner wall some of the scattered
eminences which secrete the creamy fluid, the arrow indicating
a bristle passed from the gland through the orifice in the anus.
the four main digits and the plantar pad is covered with short
hair, and the pads of the third and fourth digits of the hind
foot are separated as in the Viverrinae, not confluent as in the
2g2
452 VrVERKIDiE.
Paradoxurinse ; the retractile claws are not protected by skin-
lobes.
The skull and teeth are described under the genus.
To this subfamily belong the three genera Hemigalus,
Diplogale, and Ghrotogale. The last two have never been
examined in the flesh, and the structure of their glands and
feet has merely been inferred from the examination of dried
skins, not a satisfactory method. Hemigalus is the only
genus which comes into the British Indian fauna. It was
not referred to by Blanford, who never even suspected the
likelihood of its occurrence in Tenasserim. Its discovery
there was one of the most interesting results of the Mammal
Survey.
Genus HEMIGALUS Jourdan.
Hemigalus, Jourdan, C. B. Acad. Soi. Paris, v, p. 442, 1837.
Hemigalea, Geoffroy St. Hilaire & Blainville, ibid. p. 595.
Hemigale, Gray, Proo. Zool. Soc. 1864, p. 542 ; and of most subse-
quent authors till 1915.
Tolerably similar to the other so-called Palm-Civets in shape
of body, length of limb and of tail, but more lightly built
than Paradoxurus or Paguma, and with a shorter, thinner
coat and more tapering tail. The coloration is very charac-
teristic, the pattern consisting of longitudinal dark bands on
the head, a thin median band running from the black muzzle
to the occiput and one, broader, on each side extending from the
muzzle to the base of the ear on the inner side and interrupted by
a white spot over the eye ; of two broad stripes from the nape
to the shoulders, where they break up or expand laterally into
transverse bands"; and on the back between the shoulders
and the root of the tail of five broad transverse bands ; the
base of the tail has normally two or three transverse bands,
the rest of the organ being black. Everywhere on the dorsal
surface this pattern is emphasized by the pale ground-colour,
and the legs and underside are pale and without pattern.
The rhinarium is very like that of Paradoxurus and Paguma,
but has the infranarial portion wider ; the upper surface from
the front view is noticeably biconvex from the deep median
groove, and the angular emargination is even deeper than in
those genera. The ear is in no respects degenerate, and has
the bursa and the basal ridges and thickenings as well
developed as in Prionodon.
The skull is long and low, with constricted postorbital area,
occasionally a complete but low sagittal crest, frontal postorbital
processes angular, but none on the zygomatic arch, palate
extending over the anterior half of the mesopterygoid fossa,
and tympanic bulla typically low and flat. Dentition not
so trenchant as in Paradoxurus and Paguma, more like that
HEMIGALUS.
453
of Arctictis or Arctogalidia ; the row of upper incisors curved ;
pm 1 conical, one-rooted, main cusp of pm? d and pm s high,
compressed, and sharp, the latter with small inner lobe ;
pm 4, (upper carnassial) with inner lobe (protocone) nearly
as long at its base as the rest of the crown, which has the
anterior cusp (parastyle) larger than the posterior cusp
(metacone) ; m 1 with inner lobe rounded and as long at base
Fig. 106.
A. Left side of bony palate of Hemigalus derbyanus. pm 1 , pm*, m 1 ,
first and fourth premolars and first molar ; ppf, posterior pala-
tine foramina.
B. Right half of mandible of the same, pm^, m lt fourth premolar and
first molar.
C. Outer side of upper carnassial (pm*) of the same.
D. The same of lower carnassial (in{).
as outer part of two-cusped crown ; m 2 about half the area of
■ml. In the lower jaw pm^ to pm 3 have compressed crowns
with high, sharp main cusp ; pm 4 much larger than pm 3
and very like m 1 (lower carnassial), which has the three normal
cusps in front and a large " heel " with one outer cusp and
454 VTVEKBID2E.
a few smaller inner cusps ; ra 2 about as large as " heel " of
m x and ■with five small cusps.
All the described forms of this genus represent in my
opinion a single species.
50. Hemigahis derbyanus (Gray). The Banded Palm-Civet.
Paradoamrus derbyanus and P. zebra, Gray, Charlesw. Mag. Nat.
Hist, i, p. 599, 1837 (Nov.).
Paradoxurus derbianus, Gray, Proe. Zool. Soe. 1837, p. 67 (Jan. 22,
1838).
Viverra boiei, S. Miiller, Tijdschi. Nat. Gesch. v, p. 144, 1838.
Viverra derbyi, Ternminck, Mon. Hamm. pt. 2, p. 343, 1841.
Hemigalea or Hemigale Jiardwickei or hardwickii of most later
authors (not Viverra hardwickii Gray, Spicil. Zool. p. 9, 1830).
HemigaVus derbianus, Thomas, Journ. Bomb. Nat. Hist. Soe. xxiii,
p. 613, 1915.
Hemigalus derbyanus, Pocock, Ann. Mag. Nat. Hist. (8) xvi,
p. 160, 1915 ; id., Proc. Zool. Soe. 1933, p. 1000*.
Locality of the types of derbyanus and derbyi, the Malay
Peninsula ; of zebra " India " ; of boiei, Borneo.
Distribution. — Tenassebim, the Malay Peninsula, Sumatra,
and some of the smaller adjoining islands ; Borneo.
Tail about three-quarters the length of the head and body
or rather less, and about four times the length of the hind
foot. The ground-colour and the pattern are individually
very variable. The coat consists of short underwool, varying
in hue from nearly white, through various shades of buff, to
orange-buff, and of smooth-lying contour hairs which, except
on the dark pattern, are mainly whitish, greyish or buffy,
those on the back having a small dusky tip. The individual
colour variations depend on the intensity and extent of these
hues and upon the degree of exposure of the underwool.
Individual variation in the pattern is well illustrated by two*
specimens, $ and $, collected by Cantor in the Malay Peninsula,
very likely in the same locality. In the $ the nuchal stripes
are comparatively narrow and widely separated and break
* This species has been mostly quoted under the specific name
hardwickii because of the belief that it was the same as the Malayan
animal described as Viverra hardwickii by Gray in 1830. But in the
first place this name -was preoccupied by Viverra hardwichii Lesson,
1827, which is a synonym of Prionodon gracilis (see, p. 342),
preoccupied because the two animals were alike dedicated to General
Hardwieke, whom Lesson called Hardwich. In the second place the-
description of V. hardwickii, taken by Gray from a painting by
Major Farquhar of a specimen from the Malay Peninsula, disagrees with
au the known specimens of the genus in having the basal three-fourths
of the tail marked with six or seven dark rings and the terminal fourth
Wack, the others having at most three rings restricted to the basal
third of the tail, of which the terminal two-thirds are black. In my
paper in 1933 I proposed the name H. de/rhyanus invisus to replace-
hardwKkti. r
HEMIGALUS. 455
up behind into three or four irregularly paired transverse
stripes. Behind these, on the shoulders, is a large isolated
scapular band narrowly divided in the middle line. These
are followed by five dorsal bands, the last, at the root of the
tail, being transversely divided into two narrow stripes. In
the ^ the two nuchal stripes are wide, narrowly separated, both
being continuous with the scapular bands, the right being more
broken up than the left, which is confluent with the first dorsal
band, and all the dorsal bands differ from those of the $.
In general colour these two specimens also differ profoundly.
In the $ the dorsal interspaces and the limbs are clear whitish-
grey with silvery sheen, only faintly dimmed by brown
speckling, and the throat, chest, and belly are decidedly buff.
In the cj the dorsal interspaces and the limbs are not nearly
so grey, being very perceptibly darkened by the deeper, more
extensively brown tips of the hairs, the interspaces on the nape
and, to a less extent, on the head being so heavily pigmented
as partially to obscure the stripes ; the belly is greyer than in
the $. These two specimens show clearly that differences
in the ground-tint and in the shape, width, and degree of
disruption of the bands cannot be relied on for the differentia-
tion of races of this species. These features have been used for
that purpose.
I am unable to distinguish, even subspecifically, Bornean
specimens, described as boiei, from the typical form from the
Malay Peninsula. Hence the synonymy given above applies
to this race, H. derbyanus derbyanus. Miller described as
representing a distinct species, H. minor, some specimens
from S. Pagi Island, off the west coast of Sumatra (Smiths.
Misc. Coll. xlv, p. 43, 1913). This I regard as a valid sub-
species, distinguished by its slightly smaller size and by the
indistinctness of the pattern of the head and neck ; but
the only race of real importance in connection with the Indian
fauna is the following, from Tenasserim.
50 «. Hemigalus derbyanus incursor Thomas.
Hemigalus derbitmus incursor, Thomas, Joum. Bomb. Nat. Hist.
Soc. ssiii, p. 613, 1915.
Hemigalus derbyanus derbyanus, Poeook, Proe. Zool Soc. 1933,
p. 1000 (in part).
Locality of the type, Bankachon, Tenasserim.
Distribution. — Only known from the type-locality.
Distinguished provisionally from the typical race on the
strength of some evidence that the tail is a little longer, the
teeth on the average a little smaller, and the bulla possibly
a little more expanded on the average in its anterior part.
The bulla, however, is individually very variable in the
species.
456 VTVEEEIDJE.
The evidence for the admission of this subspecies is not
very satisfactory. In my revision of the genus in 1933
I was unable to examine the skull of the type of incursor,
an adult <J, and was dependent on the young adult $ skull
of a topotype. In this the unworn teeth were the same size
as in a $ specimen of the typical race from Malacca ; and since
Thomas had based the race upon some supposed dfferences
in the colour and pattern, which do not exist, and upon its
smaller teeth, I regarded incursor as a synonym of typical
derbyanus. But in the skull of the type which has come
to hand the teeth are decidedly smaller, pm s , pm*, and m 1
being about 1 mm. narrower, owing to the reduced inner
lobes, than in Malayan skulls. Provisionally, therefore, the
Tenasserim specimens may be given the status assigned to
them by Thomas.
The external differences relied on by Thomas were the
generally lighter, more silvery hue, and the unbroken, con-
tinuous nuchal stripes. It is true that the type is slightly
lighter and greyer in ground-colour than the greyest Malaecan
specimen he examined, but the topotype is decidedly more
bufiy than in two Malaecan specimens. As for the nape-
stripes, they, as stated above, may be either greatly broken up or
form broad, uninterrupted bands in the typical Malayan race.
The flesh-measurements (in English inches) and weights
(in lb.) of the two examples of this race collected by
Shortridge are entered in the following table, together with
the measurements of two specimens of the typical race, to
show especially the apparently longer tail in the former.
Head and Hind
Locality, name, and sex. body. Tail. foot. Weight.
Bankaehon (incursor type) ;
ad. d 20* 15 3£ 5
Bankaehon ; yg. ad. <J 201 15+ 31 4
Pahang, Malay Peninsula
(derbyanus) ; ad. (J 19f 13£ 3 J —
Tapanuli Bay, Sumatra (derby-
anus) (Miller) ; ad. $ 20* 12f 3- —
For the skull- and tooth-measurements of incursor and of
derbyanus see p. 457.
Habits. — Of the habits of Hemigalus very little is known.
In Tenasserim, according to Shortridge, it is apparently not at
all plentiful, the Malay villagers at Bankaehon having no name
for it. It is probably, he thinks, one of the most active of
all the Viverridse *, and is without doubt largely arboreal.
* That it excels Paradoxurus and Pagwma in this respect might be
inferred from its lighter build and narrower, more delicate paws ;
but it is very doubtful if it similarly surpasses Prionodon and Arato-
galidia.
HEMIG-AirrS.
457
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458 YIVEKEID^l.
His specimens, however, appear to have been trapped on the
ground. When caught alive it is very savage and growls
like a cat ; but it has absolutely no smell. Its tongue,
he adds, is very rough, much more so than in any other small
Carnivore.
Despite Shortridge's observation that the animal is absolutely
•without smell*, I suggested in 1933 that its very aberrant
and striking coloration, -which theoretically must have some
special significance, may be for advertisement, indicating
nauseous protective secretion of the anal glands, such as is
known to exist in Paradoxurus and Paguma (see pp. 414 & 430),
although so infrequently detected by collectors. The truth of
this surmise was established on a fresh $ specimen in which
the anal glands showed superficially as a swelling on each
side of the anal orifice. Each consisted of large oval sac
of which the inner walls were studded with scattered emi-
nences from which a creamy fluid was secreted into the sac,
which was filled with a darker, more liquid secretion. This
secretion had a most unpleasant '' musteline " odour, reminding
me forcibly of that of the Stoat (Mustela erminea).
Of the other genera of Hemigalinse, Ghrotogale Thomas
(Proc. Zool. Soc. 1912, p. 47, and 1927, p. 47, and of subse-
quent authors) may be discovered in Upper Burma, since its
only known species, owstoni, has been recorded from Tong-
king and Laos. It resembles Hemigalus in its pattern of
broad bands on a light ground-colour, but has in addition
conspicuous black spots on the sides of the neck, the upper
part of the limbs, and a few on the flanks. But it differs
essentially from that genus in the peculiar elongation of
the muzzle of the skull, accompanied by broad upper incisors,
forming a strongly curved line, and other dental pecularities.
The other genus, Diplogale Thomas, occurs in Borneo, and is
not at all likely to turn up in British Indian territory. It
differs from the others in being tolerably uniformly otter-
brown in hue, when adult, and although the skull in most
of its features is like that of Hemigalus, it differs in a few
structural details as well as in some dental peculiarities.
Further particulars about these genera may be found in my
paper (Proc. Zool. Soc. 1933, pp. 1009-12).
* A puzzling remark, because from the presence of the perfume -
glands it may be inferred with certainty that Hemigalus has the ordinary
" Civet odour " of the species that possess them.
ALPHABETICAL INDEX.
achates (Semnopitlieeus entellus),
103.
aehilles (Semnopitheceus entellus),
95.
Aeinonychinse, 322.
Aeinonyx, 323.
adusta (Macaea nemestrina), 59.
-<Eluroidea, 190.
seneas (Semnopithecus entellus), 106.
affinis (Palis chaus), 294.
aharonii (Caracal caracal), 307.
ajax (Semnopitheeus entellus), 96.
albibarbatus (Maeaea silenus), 66.
albifrons (Aretictis binturong), 433.
albinus (Kasi senex), 154.
anchises (Semnopitheeus entellus),
101.
andamanensis (Macaea nemestrina),
59, 63.
Anthropomorpha, 15.
Aretictis, 431.
Aretogalidia, 441.
AretogaUdiinae, 439.
Arctoidea, 190.
arctoides (Macaea speciosa), 69, 75.
asiatieus (Panthera Ieo), 212.
assamensis (Macaea), 52.
assamensis (Macaea assamensis), 52.
Assamese Macaque, 52.
atrior (Trachypithecus pyrrhus),
143.
aurea (Macaea irus), 79.
aureus (Paradoxurus), 381.
aurifrons (Macaea sinica), 37.
Banded Linsang, 340.
baptistae (Viverricula indica), 370.
barbei (Trachypithecus phayrei),
136, 143.
bengalensis (Nyeticebus eoucang),
166.
bengalensis (Panthera leo), 212.
bengalensis (Prionailurus), 267.
bengalensis (Prionailurus bengalen-
sis), 268.
bengalensis (Viverricula indica),
367.
Binturong, 432.
binturong (Aretictis), 432.
binturong (Aretictis binturong),
435.
birmanious (Paradoxurus herma-
phroditus), 402.
blythii (Macaea nemestrina), 62.
bondar (Paradoxurus hermaphro-
ditus), 398.
Bonnet Macaque, 38.
brahma (Trachypitheeus pileatus),
128.
eaniseus (Paradoxurus jerdoni), 286.
Caracal, 306.
caracal (Caracal), 306.
caracal (Caracal caracal), 307.
Carnivora, 186.
Catarhini, 15.
Cats, Feral Domesticated, 304.
Cercopithecidse, 32.
charltoni (Pardofelis marmorata),
256.
chaus (Felis), 290.
chaus (Felis chaus), 292.
cinereus (Nyeticebus eoucang), 166.
civettina (Mosehothera), 358.
Clouded Leopard, 247.
460.
AHPHABETICAL INDEX.
cochinensis (Paradoxurus herma-
phrodites), 412.
Colobidae, 83.
Common Palm-Civet, 387.
eonstantina (Felis), 286.
coolidgei (Maeaea assamensis), 53.
corax (Traehypithecus obscurus),
140.
couoang (Nyctieebus), 166.
eoucang (Nyctieebus coueang), 171.
Crab-eating Macaque, 78.
crepusculus (Traehypithecus
phayrei), 134.
crossi (Paradoxurus hermaphro-
dites), 398.
cucullatus (Kasi), 147.
Cynailurus, 323.
Cynamolgus, 34.
eynomolgus (Macaca irus), 79, 82.
Cynomorpha, 31.
derbyanus (Hemigalus), 454.
deserti (Viverricula indica), 368.
diluta (Macaca radiata), 42.
durga (Traehypithecus pileatus),
125.
dussumieri (Semnopitheeus entellus),
107.
elissa (Semnopitheeus entellus), 113.
entelloides (Hylobates lar), 26.
entellus (Semnopitheeus), 90.
entellus (Semnopitheeus entellus),
faseieularis (Macaca irus), 79.
Felidae, 190, 191.
Felinas, 243.
Felis, 285.
femoralis (Presbytia), 159.
ferox (Macaca silenus), 66.
ferrugineus (Otoeolobusmanul), 319.
Fishing-Cat, 281.
flavieauda (Traehypithecus ob-
seurus), 140.
fulvidina (Felis ehaus), 303.
fasca (Panthera pardus), 226.
fuscus (Paradoxurus hermaphro-
ditus), 410.
Gibbons, 17.
Golden Cat, 261.
Golden Palm-Civet, 381.
goojratensis (Panthera leo), 212.
gracilis (Loris tardigradus), 181.
grandis (Loris tardigradus), 184.
grayi (Paguma larvata), 420.
hector (Semnopitheeus entellus),
92.
Hemigalinse, 450.
Hemigalus, 452.
hermaphrodites (Paradoxurus), 287.
hermaphrodites (Paradoxurus her-
maphrodites), 388.
Herpestidss, 190.
Hoolock Gibbon, 19.
hooloek (Hylobates), 19.
horsfleldi (Prionailurus bengalen-
sis), 271.
Hysenidas, 190.
Hylobates, 19.
Hylobatidse, 17.
hypoleucos (Semnopitheeus entel-
lus), 108.
ineanus (Nyctieebus coueang), 167.
ineursor (Hemigalus derbyanus),
455.
indioa (Viverricula), 363.
indica (Viverricula indica), 364.
indicus (Panthera leo), 213.
mdochmensis (Macaca nemestrina),
60.
insulana (Maeaea nemestrina), 59.
intrudens (Paguma larvata), 425.
irbis (Uncia), 240.
irus (Maeaea), 79.
isabellinus (Lynx lynx), 311.
iulus (Semnopitheeus entellus),
104.
janetta (Paguma larvata), 427.
jerdoni (Paradoxurus), 383.
jerdoni (Paradoxurus jerdoni),
383.
Jerdon's Palm-Civet, 383.
johnii (Kasi), 147.
jubatus (Aeinonyx), 324.
Jungls-Cat, 290.
Kasi, 146.
keatii (Presbytis femoralis), 161.
kelaarti (Felis ehaus), 300.
kephalopterus (Kasi senex), 154.
kutas (Felis ehaus), 297.
laneus (Paradoxurus hermaphro-
dites), 394.
Langur, 84.
lanigera (Paguma), 416.
laotum (Paradoxurus hermaphr .
ditus), 402.
ALPHABETICAL INDEX.
461
Large Indian Civet, 346.
Lar-Gibbon, 26.
lar (Hylobates), 26.
larvata (Paguma), 417.
Leaf-Monkey, 84.
Lemuroidea, 163.
leo (Panthera), 210.
leonina (Maeaea nemestrina), 59, 62.
Leopard, 222.
Leopard-Cat, 267.
leuootis (Aretogalidiatrivirgata),
444.
Linsang, 334.
linsang (Prionodon), 339.
Lion, 210.
Lion-tailed Macaque, 66.
Loris, 174.
Lorisidae, 164.
lydekkerianus (Loris tardigradus),
177.
Lynx, 310.
lynx (Lynx), 311.
Lyssodes, 34.
Macaca, 32.
Macaques, 32.
memahoni (Macaca raulatta), 50.
macra(Aretogalidia trivirgata), 446.
macrosceloides (NeofeKs nebulosa),
250.
maculosus (Prionodon linsang), 339.
Malabar Civet, 358.
malabaricus (Loris tardigradus),
181.
malaceensis (Viverra), 352, 363.
manul (Otocolobus), 317.
Marbled Cat, 255.
marmorata (Pardofelis), 255.
mayori (Viverricula indica), 363.
megaspila (Mosehothera), 356.
melamerus (Traehypitheeus phay-
rei), 136.
melanotus (Macaca speeiosa), 73.
millardi (Panthera pardus), 233.
millsi (Aretogalidiatrivirgata), 447.
minor (Paradoxurus hermaphro-
ditus), 405.
Monkeys, 31.
moormensis (Profelis temminckii),
261.
Mosehothera, 354.
mulatta (Macaca), 44.
mulatta (Macaca mulatta), 45.
nebulosa (NeofeKs), 248.
neglecta (Paguma larvata), 422.
nemestrina (Macaca), 58.
Neofelis, 247.
nestor (Kasi senex), 153.
Nicobar Crab -eating Macaque, 82,
nictitatans (Paradoxurus herma-
phroditus), 392.
niger (Paradoxurus), 389.
nigriceps (Paguma larvata), 424.
nigripectus (Otocolobus manul), 319.
nordicus (Loris tardigralus), 182.
Nyctieebus, 165.
obscurus (Traehypitheeus), 138.
ornata (Pelis constantina), 287.
Otocolobus, 315.
Ounce, 240.
Paguma, 415.
pallasii (Paradoxurus hermaphro-
ditus), 400.
pallens (Paradoxurus hermaphro-
ditus), 410.
pallipes (Semnopithecus entellus),
109, 115.
Palm-Civet, 387.
Panther, 222.
Panthera, 196.
Pantherinas, 195.
Paradoxurinse, 376.
Paradoxurus, 379.
pardicolor (Prionodon), 337.
Pardictis, 334.
Pardofelis, 253.
pardus (Panthera), 222.
pelops (Macaca assamensis), 55.
pernigra (Panthera pardus), 231.
persica (Panthera leo), 212.
phillipsi (Prionailurus rubiginosus),
278.
phayrei (Traehypitheeus), 129.
phayrei (Traehypitheeus phayrei),
130. J '
philbricki (Kasi senex), 154.
picta (Viverra zibetha), 350.
Pig-tailed Macaque, 58.
pileatus (Macaca), 34.
pileatus (Traehypitheeus), 121.
pileatus (Traehypitheeus pileatus),
Pithecoidea, 14.
prateri (Felis chaus), 298.
Presbytis, 158.
priam (Semnopithecus entellus),
109.
priamellus (Semnopithecus entellus),
112.
Primates, 13.
Prionailurus, 265.
Prionodon, 334.
Prionodontinas, 332.
462
ALPHABETICAL INDEX.
problematieus (Maeaoa assamensis)
55.
Profelis, 258.
Protective odour of Macaca speciosa,
77; of Paradoxurus, 414; of
Paguma, 430 ; of Hemigalus,
458.
pruinosa (Viverra zibetha), 352.
pugnax (Paradoxurus hermaphro-
ditus, 410.
puleher (Paradoxurus hermaphro-
ditus), 410.
pyrrhus (Traehypithecus), 142.
quadriscriptus (Paradoxurus herma-
phrodite), 400.
radiata (Macaca), 38.
radiata (Macaca radiata), 40.
rheso-similis (Macaca assamensis),
55.
Ehesus, 34.
Bhesus Macaque, 44.
robusta (Paguma larvata), 426.
rubiginosus (Prionailurus), 276.
rubiginosus (Prionailurus rubigino-
sus), 277.
Busty-spotted Cat, 276.
sacer (Paradoxurus hermaphro-
ditus), 410.
sanctorum. (Traehypithecus obsou-
rus), 140.
saturatus (Traehypitbecus pileatus),
125.
saxicolor (Pantbera pardus), 234.
schistaceus (Semnopitbecus entel-
lus), 92.
sehmitzi (Caracal caracal), 307.
seindias (Paradoxurus bermapbro-
ditus), 392.
Semnopitbecus, 88.
senex (Kasi), 150.
senex (Kasi senex), 154.
senex (Paradoxurus hermaphro-
dite), 409.
shanicus (Traehypithecus phayrei),
136.
shawiana (Pelis), 290.
shortridgei (Traehypitbecus pilea-
tus), 128.
sigillata (Viverra zibetha), 352.
Silenus, 33.
silenus (Macaca), 66.
sindica (Pantbera pardus), 233.
sinica (Macaca), 34, 38.
sinica (Macaca sinica), 36.
Slender Loris, 176.
Slow Loris, 165, 169.
Snow-Leopard, 240.
smithi (Traehypithecus obscurus),
140.
speciosa, Macaca, 69.
speciosa (Maoaca speciosa), 71.
Spotted Civet, 356.
Spotted Linsang, 337.
strictus (Paradoxurus hermaphro-
dite), 398.
Stump-tailed Macaque, 69.
surdaster (Viverra zibetha), 350.
tardigradus (Loris), 175.
tardigradus (Loris tardigradus), 181.
temminckii (Profelis), 260.
Temminck's Cat, 261.
tenasserimensis (Nycticebus cou-
eang), 169.
tenebricus (Traehypithecus pilea-
tus), 126.
thai (Viverrieula indioa), 372.
thersites (Semnopitheeus entellus)
115.
Tibetan Lynx, 311.
Tiger, 197.
tigris (Panthera), 197.
tigris (Panthera tigris), 199.
Toddy-Cat, 387.
Toque Macaque, 34.
torquata (Felis constantina), 287,
305.
Traehypitbecus, 120.
trevelyani (Prionailurus bengalen-
sis), 273.
Triebselurus, 315.
tristis (Profelis temminckii), 263.
trivirgata (Arctogalidia), 444.
tulliana (Panthera pardus), 234.
tytlerii (Paguma larvata), 424.
umbrosa (Macaca irus), 82.
Uncia, 239.
uneia (Uncia), 240.
vagans (Paguma larvata), 425.
veflerosus (Paradoxurus herma-
phrodite), 397.
venaticus (Acinonyx jubatus), 325.
vetulus (Kasi senex), 151.
vicinus (Paradoxurus hermaphro-
ditus), 400.
villosa (Macaca mulatta), 49.
ALPHABETICAL INDEX.
463
vitiis (Macaca irus), 79.
Viverra, 344.
Viverricula, 362.
Viverridifi, 190, 330.
Viverrinas, 342.
viverrinus (Prionailurus), 281.
wardi (Lynx lynx), 311.
Warning coloration of Pcvradoxwus,
414, of Paguma, 430 ; of Hemi-
galus, 458.
wellsi (Viverricula indica), 369.
wroughtoni (Paguma larvata), 418.
yunalis (Paguma larvata), 425.
Zati, 34.
zeylonensis (Paradoxurus), 381.
zibetha (Viverra), 346.
zibetha (Viverra zibetha), 347.
PRINTED BY TAYLOR AND FEANCIS, LTD.
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The Fauna of British India,
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Published under the Patronage of the Secretary of State
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LIST OF VOLUMES PUBLISHED AND IN PREPARATION.
MARCH, 1939.
{Those marked * are out of print. Except where publication is known
to have been earleir, dates quoted are those on which the
volumes were first received at the India Office.)
VERTEBRATA.
MAMMALIA.
[FmsT Edition.] By W. T. Blanfoed.
*Part I. [Primates, Carnivora, Insectivora]. Pp. i-xii, 1-250, text-figs.
Aug. 31, 1888.
*Part II. [Chiroptera, Rodentia, Ungulate, Cetacea, Sirenia, Edentata]. Pp. i-xx.
251-617, text-figs. Dee. 18, 1891.
Second Edition.
Vol. I. [Primates and Carnivora]. By K. I. Pooock. Pp. i-xxxiii, 1-464, 31 pis.,
map, text-figs. 30/- March 31, 1939.
This edition will probably occupy three volumes.]
BIRDS.
[First Edition.]
*Vol. I. [Passeres]. By Euqene W. Oaths. Pp. i-xx, 1-556, text-figs.
Dec. 30, 1889.
*Voi. II. [Passeres, concluded]. By Eugene W. Oates. Pp. i-x, 1-407, text-figs.
Dec. 8, 1890.
Vol. III. [Eurylsemi, Pioi, Zygodactyli, Anisodactyli, Macrochires, Trogones,
Coccyges, Psittaci, Striges, Accipitresj. By W. T. Blanfobd. Pp. i-xiv,
1-450, text-figs. 21/- Oct. 2, 1895.
Vol. IV. [ColumbEe. Pterocletes, Gallmse, Hemipodii, Grails, Limicols, Gavise,
Steganopodes, Tubinares, Herodiones, Phcenicopteri. Pygopodes]. By W. T.
Blaotobd. Pp. i-xxi, 1-500, text-figs. 21/- April 25, 1898.
Second Edition. By E. C. Stuart Baker.
Vol. I. [Passeres, Fam. I. Corvidse— VIII. Troglodytidse]. Pp. i-xxiii, 1-479, 8 col.
pis., text-figs. 30/- Aug. 24, 1922.
Vol. II. [Passeres. Fam. IX. Cinclidss — XVTI. Regnlidsel. Pp. i-xxiii, 1-561, 8 col.
pis., text-figs. 30/- April 30, 1924.
Vol. III. [Passeres, Fam. XVIII. Irenidse— XXXIII. Eurylaimidse]. Pp. i-xx.
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[1]
Vol. IV. [Coraciiformes]. Pp. i-xxiv, 1-471, 7 col. pis., text figs. 30/- July28, 1927.
Vol. V. [Accipitres, Columbas, Pterocletes, Galling, Hemipodii]. Pp. i-xviii, 1-469,
6 col. pis., text-figs. 30/- March 21, 1928.
Vol. VI. [Grallse, Charadriiformes, Steganopodes, Tubinares, Herodiones, Phceni-
copteri, An8eres, Pygopodes], Pp. i-xxv, 1-499, 3 pis., text-figs. 30/-
March 26, 1929.
Vol. VII. [Synonymical Catalogue, PaBseres— Grallse]. Pp. i-viii, 1-484. 30/-
March 30, 1930.
Vol. VIII. [Synonymical Catalogue, Grallse — Pygopodes ; Corrigenda and Addenda;
Index]. Pp. i-iv, 485-801. 15/- Sept. 25, 1930.
REPTILIA and BATRACHIA.
[*Frasx Edition, complete in 1 vol.] By George A. Boltlenger.
Pp. i-xviii, 1-541, text-figs. Sept. 4, 1890.
Second Edition. By Malcolm A. Smith.
Vol. I. Lorioata, Testudines. Pp. i-xxviii, 1-185, 2 pis., map, text-figs. 15/-
March 27, 1931.
Vol. H. Sauria. Pp. i-ix, 1-440, 1 pi., 2 maps, text-figs. 30/- Feb. 7, 1935.
[Vol. III., Ophidia, by Dr. Malcolm A. Smith, is in course of preparation.]
FISHES.
[First Edition.] By Francis Day.
Vol. I. [Chondropterygii, Teleostei (Physostomi ; Acanthopterygii : Percids)].
Pp. i-xviii, 1-548, text-figs. 28/- July 11, 1889.
Vol. II. [Teleostei (Acanthopterygii excl. Percidaa ; Anacanthini ; Lophobranchii ;
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[A second edition, by Dr. Sunder Lax Hora, is in course of preparation. It is
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ARTHROPODA.
LEPIDOPTERA.
MOTHS. By G. F. Hampson.
Vol. I. [Fam. 1, Saturniidse — 23, Hypsidse]. Pp. i-viii, 1-527, text-figs. 28/-
Jan. 10, 1893.
Vol. II. [Fam. 24, Arctiidse; 25, Agaristidse; 26, Noctuidse], Pp. i-iv, 1-609,
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Vol. TIL [Fam. 26, Noctuidse (Subfam. Foeillinie, Deltoidinse) ; 27, Epicopiidse ;
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Vol. IV. [Fam. 31, Pyralidse ; additions and corrections to Fam. 1-30]. Pp. i-xxviii,
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Vol. V [Sphingidae]. By. E. D. Bell and F. B. Scott. Pp i-xviii, 1-537, 15 pis.,
text-figs. j une i 5j 1937,
[2]
BUTTERFLIES. [Fibst Edition.] By C. T. Bingham.
■"Vol. I. [Nymphalidss, Nemeobiidse]. Pp. i-xxii, 1-511, 10 col. pis., text-figs.
March 2, 190S.
Vol. II. [Papilionidas, Pieridse, LyeasnidaB (part)]. Pp. i-viii, 1-480, 10 col. pis.,
text-figs. 28/- March 25, 1907.
[Vol.111, of the first edition was never completed.]
[Second Edition.]
Vol. I. [Papilionidse, Pierids.] Pp. i-xxix. 1-600, 3 pis., map, text-figs. 35/-
March 8, 1939.
[This edition, by Mr. G. Talbot, will embrace all the Butterflies and will probably
extend to five volumes.]
COLEOPTERA.
ADEPHAGA.
General Introduction, and Cicindolidse and Paussidaa. By W. W. Fowleb. Pp. i
xx, 1-529, text-figs. 28/- Received in Brit. Mm. {Nat. Hist.) Feb. 18, 1912.
Carabidse: Vol. I. Carabinaa. By H. E. Andeewes. Pp. i-xviii, 1-431, 9 pis.,
text-figs. 22/ 1 May 15, 1929.
Carabidse : Vol. II. Harpalinse — I. By H. E. Andeiswes. Pp. i-xvi, 1-323, 5pls.,
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[A volume on Dytiscidas, Gyrinidas, and Haliplidae, by Mr. J. BAiroua Bbownb,
is in preparation.}
STAPHYLINOIDEA.
Staphylinidse. By Malcolm Camebon.
Vol. I. [Subfam. Micropeplinse, Oxytelinse, Oxyporinae, Megalopinae, Steninse,
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Vol. II. [Subfam. Psederinie.] Pp. i-viii, 1-257, 2 col. pis., text-figs. 15/-
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Vol. III. [Subfam. Staphylininse, Trichophyinas, Termitodiscinse, Pygosteninaa,
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[Vol. IV. is in course of preparation.]
CLAVICORNIA.
Erotylidse, Languriidse, and Endomychidae. By G. <T. Abbow. Pp. i-xvi, 1-416,
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PHYTOPHAGA.
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Chrysomelidae.
Vol. I. [Eupodes, Camptosomes, Cyclica]. By Maetqj Jaooby. Pp. i-xx, 1-534,
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Vol. II. [Hispinas and Cassidinae]. By S. Mauxbs. Pp. i-xi, 1-439, text-figs. 21/-
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[3]
RHYNCHOPHORA.
Curculionidae. [Part I. Brachyderins, Otiorrhynehinse.] By Guy A. K. Marshall.
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[A volwne on Platypodia, by Dr. C. F. C. Beeson, is in preparation, and wUl
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Scarabseidse By G. J. Arrow.
Part I. Cetoniinse, Dynastinae. Pp. i-xiv, 1-322, 2 col. pis., text-figs. 14/-
Sept. 13, 1910.
Partll.Rutelinse, Desmonycinee, Euchirinffi. Pp.i-xiii, 1-387, 5 pis., text-figs. 21/-
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HYMENOPTERA.
Vol. I. Wasps and Bees. [Fossores, Diploptera, Anthophila.] By C. T. Bingham.
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Vol. II. Ants and Cuckoo-Wasps. [Formicidse, Chrysidids.] By C. T. Bdtoham.
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Vol. III. Ichneumonidse : I. Ichneumones Deltoidei [Pimplinse, Tryphoninse, Ophio-
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DIPTERA.
[Vol. I.] Nematocera, excluding [Cecidomyiidas], Chironomidse, and Culicidse. By
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[Vol. IL]Brachycera, Vol. I. [Stratiomyiidaa, Leptidse, Nemestrinidas, Cyrtidse, Bomby-
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APHANIPTERA.
[A Volume on the Fleas, by Dr. M. Sharif, is in course of preparation.]
BHYNCHOTA.
By W. L. Distant.
Vol.1. Heteroptera [Pentatomidas, Coreidse, Berytidse]. Pp. i-xxii, 1-438, text-figs.
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Vol. H. Heteroptera [Fam. 4, Lygaeidse— - 16, Capsidse,] Pp. i-xvii, 1-503, text-figs.
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[4]
Vol. III. Heteroptera — Homoptera [Anthocorida, Polyetenidsa, Cryptoeerata,
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midse, Coreidse, and Berytidse]. Pp. i-xv, 1-501, text-figs. 28;- 1907-8.
[ First published intwo parts: Part I, pp. 1-264, in Nov. 1907 ; Part II, pp. 265-501,
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Vol. V. Heteroptera : Appendix [Lygseidsa to Cryptoeerata]. Pp. i-xii, 1-362,
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Vol. VI. Homoptera : Appendix [Cicadidaa, Fulgoridse, Membracidse, Ceroopidae,
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Vol. VII. Homoptera : Appendix [Jassidte (pt.)] ; Heteroptera : Addenda [Penta-
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ORTHOPTERA.
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DERMAPTERA.
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ODONATA.
Vol. I. [Coenagriidas]. By P. C. Fbasee. Pp. i-xiii, 1-423, map, text-figs. 25;-
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Vol. H. [Agriidae and Gomphidae]. By F. C.Fbaser. Pp. i-xxiii, 1-398, 4 col. pis.,
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ECHINODERMATA.
[A volume on the Echinoidea, by Dr. Th. Mobtensen, is in course of preparation.]
MOLLUSCA.
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[5]
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CCELENTERATA, etc.
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PORIFERA.
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