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DEER OF ALL LANDS

Peer oh Oe erO-r VP ELEM FAMILY CERVIDA
EMNVING AND EXTINCT

BY

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RY LYDEKKER

LONDON
ROWLAND WARD, Lunrep
y 1898

YRAREL
YEOLOOL CU 08. Eli
CEM BOOHEN NE

INO

CEE OUCH ros, OF BEDFORD.
THIS ACCOUNT
OF A GROUP OF ANIMALS
IN WHICH

HER GRACE TAKES A SPECIAL INTEREST

IS DEDICATED
BY

Trt AWMHOR

- |

PRE PACE

Many groups of animals are of interest chiefly or solely to the naturalist,
but those coming under the designation of big game claim a much more
extensive list of admirers. And among these the members of the deer tribe
occupy not the least important position. Not only are the majority objects
of eager pursuit to the sportsman, but three are denizens of our larger
British parks, and two of these are the only British representatives of big
game. Moreover, many of them are capable of more or less complete
acclimatisation in this country, and in that condition add largely to the
attractions of our deer parks.

It is now many years since a complete monograph of the existing
members of the group has appeared, among the latest being one by the
late Sir Victor Brooke in 1878, and a second by the late Dr. L. J. Fitzinger,
which, although commenced in 1873, was not completed till 1879. Since
these dates several new species have been described, and much revision of
others, as well as of the general grouping, has taken place. And the present
volume is an attempt to bring up to date the existing state of our knowledge
of an exceedingly difficult group. Although the author has availed himself
of all the accessible means of information, there are still many points
requiring further elucidation, and much is necessarily left for future
investigation.

Since the author is a palzontologist first and a zoologist second, he could
not treat the subject without some reference to the extinct members of the
family. Many of these, however, have been named on the evidence of such

unsatisfactory fragments that nothing would be gained by their mention

Vill Preface

here ; and the notices of extinct species are consequently limited to such as
can be located among the existing forms with a fair approximation to
certainty.

Being essentially popular in its treatment of the subject, the work makes
no pretence to rank as a strictly scientific treatise, and should not therefore
be judged from such a standard. Anatomical details and technical terms
are in most cases so far as possible avoided. For instance, the characters
of the skull are in general not mentioned ; but in a scientific work it would
have to be stated that the common American deer and its numerous local
races are easily distinguished from the mule-deer by the shallowness of the
pits in the skull for the reception of the face- glands. In spite of its
popular treatment, it is nevertheless probable that the professed naturalist
will not fail to find much matter of interest, and possibly some aid in his
investigations.

An exceedingly difficult and unsatisfactory part of the subject has been
that relating to scientific nomenclature. And both the sportsman and the
amateur naturalist will probably be disagreeably surprised to find a large
proportion of the species figuring under titles with which they are totally
unfamiliar. At the present time zoological nomenclature is undergoing a
complete revolution, the currently accepted idea being that the names first
given to animals must be used, unless they have been previously employed in
another sense, or there is some other bar to their employment. With some
misgivings, the author has felt it advisable in the main to follow this ruling,
although he cannot at present reconcile himself to the use of duplicative
titles, such as Capreolus capreslus. Moreover, when a specific title indicates
a geographical error, it is reyected. And the same course is followed when
a name is founded on a misconception. An instance of the latter kind is
afforded by the name Hppocamelus, given to the Chilian guemal on the
supposition that it was intermediate between a horse and a llama. In this

case the name Mazama, although later, is selected for use.

Preface 1X

In view of these changes, special prominence has been given in this
volume to the popular titles of the various species, which are less subject to
such emendations, and these alone have been employed in the legends to
the plates and text-figures.

A new feature in the treatment of the group is the large number of
forms ranked as sub-species, or races, instead of as full species. ‘The principle
of this is that when certain representatives of a genus, or a group of a genus,
are more nearly allied to one another than they are to the other members
of the same, they are regarded as sub-species. The Asiatic and American
wapitis, for instance, are obviously more closely related to one another than
they are to the red deer, while the Caspian maral and the North African
deer come nearer to the latter than to any other members of the same group.
Consequently all the forms of wapiti are classed as geographical races of a
single variable and widely spread specific type; while the European, the
Caspian, and the North African red deer are ranked as sub-species of a
second. In like manner the numerous modifications of the sambar type
show mutual resemblances not shared either by the chital or the para, and
all the former are accordingly brigaded under one specific title. Although
in some cases such geographical races, or sub-species, may pass imperceptibly
into one another, the absence of intergradation is not considered any bar to
classing any particular form as a sub-species. Where such sub-species are
separated by the ocean, such intergradation must obviously have ceased after
the disruption of the land connection between their respective habitats.

Generic terms are likewise, for the most part, used in a wide sense,
although they are frequently split up into minor groups, or sub-genera,
which many writers would probably prefer to rank as genera. If, however,
genera are used in a more restricted sense we are very apt to lose sight of
many broad generalisations. For instance, if Cervus be split up, we lose
sight of the fact that all the large brow-antlered deer are essentially an Old

World type, with western representatives in the American wapitis. And

x Pretace

similarly, if the American deer be referred to two or more genera, the fact
that all the deer peculiar to the New World are characterised by the
possession of antlers of the bifurcate type, accompanied by the division of
the nasal passage of the skull into two chambers, is not obvious.

The author’s personal acquaintance with the animals of which he treats
is by no means equal throughout. It is least in the smaller deer of Central
and South America, and it is consequently among these that subsequent
revision will probably be most necessary.

To the kindness of numerous friends the author is greatly indebted for
much valuable assistance in various ways during the progress of the work,
and to all of these his best thanks are collectively tendered. Special mention
must, however, be made of his indebtedness to the Duke and Duchess of
Bedford, whose unrivalled collection of living deer at Woburn Abbey has
been unreservedly placed at his disposal for study. It was owing to this
collection that the idea of the work originated, and without it there would
have been no possibility of bringing the history of this group of animals to
such approach to completeness as it may haply possess.

To the Duchess herself the author’s especial thanks are likewise due for
the exquisite photographs of many specimens and groups in the collection
at Woburn with which the work is embellished. Not the least interesting
among them being the series of antlers of individual deer at different ages.

Neither must mention be omitted of the care and attention bestowed by
Mr. Smit on the execution of the coloured lithographs, many of which
have been drawn from living examples in the collection at Woburn Abbey,
while others have been taken from specimens in the Zoological Society’s

Menagerie.

HARPENDEN, Ist March 1808.

———

GON LEN TS

Part J.—INTRODuUCcTORY

DEFINITION OF DEER
ANTLERS

OTHER STRUCTURAL coe TURES
CLASSIFICATION

DisTRIBUTION

Hagsits

Economic Uses

Part JJ.—DescriptTive

A. THe True Deer—Sub-Family Cervine

I. RemnpeER—Genus Rangifer ;
The Reindeer, or Caribou—Rangifer rian
a. Scandinavian Race—Rangifer tarandus typicus

b. Spitzbergen Race—Rangifer tarandus spetzbergensis
c. Woodland Race—Rangifer tarandus caribou

d. Newfoundland Race—Rangifer tarandus terra-nove .

e. Greenland Race—Rangifer tarandus granlandicus
jf. Barren-Ground Race—Rangifer tarandus arcticus

II. Er—x—Genus Aces :
. The Elk, or Moose—ces is
2h ee s Elk— Alces latifrons (extinct) .
3. Scott’s Elk— Alces scott? (extinct)

1H ve TypicaL DEER—Genus Cervus
. The Red Deer, or Elaphine Group—Sub- Cone (on
. The Red Deer—Cervus elaphus
a. European Race—Cervus elaphus typicus
b. Corsican Race—Cervus elaphus corsicanus
c. North African Race—Cervus elaphus barbarus

d. Caspian Race—Cervus elaphus maral .

NJ

real

XI

Contents

III. THe Typrcat DEER—continued

ill.

iv.

N

. The Duke of Bedford’s Deer—Cervus xanthopygus
3. The Hangul—Cervus cashmirianus

a. Kashmir Race—Cervus cashmirianus typicus

6, Yarkand Race—Cervus cashmirianus yarcandensis
4. The Shou—Cervus affinis
5. Thorold’s Deer—Cerwus albirostris
6. The Wapiti—Cervus canadensis .
a. East American Race—Cervus canadensis typicus
b. West American Race—Cervus canadensis occidentalis
ce. Manchurian Race—Cervus canadensis luehdorfi
d. Altai Race—Cervus canadensis astaticus

. The Bokhara Deer—Cervus sp.

a |

. The Sikine Group—Sub-Genus Pseudaxis . ;

1. The Common Sika—Cervus sica
a. Japanese Race—Cervus sica typicus
b. Manchurian Race—Cervus sica manchuricus
2. The Formosan Sika—Cervus taévanus
3- The Pekin Sika—Cervus hortulorum
4. Mandarin Sika—Cervus mandarinus
5. Uhe Pliocene Sika—Cervus perrieri (extinct)
Incerte Sedis—Cervus pseudaxis
The Damine Group—Sub-Genus Dama
1. The Common Fallow Deer—Cervus dama
2. Brown’s Fallow Deer—Cervus browni (extinct)
3. The Mesopotamian Fallow Deer—Cervus mesopotamicus
4. The Giant Fallow Deer—Cervus giganteus (extinct)
a. German Race—Cervus giganteus ruffi
b. Irish Race—Cervus giganteus typicus . :
c. Italian Race—Cervus giganteus italia .
d. French Race—Cervus giganteus belgrandi
e. Forest-bed Race—Cervus giganteus carnutorum
The Rusine or Sambar Group—Sub-Genus Rusa
1. The Sambar—Cervus unicolor
a. Indian Race—Cervus unicolor typicus
b. Malayan Race—Cervus unicolor equinus :
c. Formosan Race—Cervus unicolor swinhoet
d. Szechuan Race—Cervus unicolor dejeani
e. Luzon Race—Cervus unicolor philippinus ‘
Jf. Basilan Race—Cervus unicolor nigricans
2. The Rusa, or Javan Sambar—Cervus hippelaphus
a. Javan Race—Cervus hippelaphus typicus
b. Moluccan Race—Cervus hippelaphus moluccensis

c. Timorese Race—Cervus hippelaphus timoriensis

Contents X11

Ill. THe Typicat DEER—continued BAGH
3. The Philippine Spotted Deer—Cervus alfredi . ; 172

4. The Calamianes Deer—Cervus culionensis : : EF

5- The Bavian Deer—Cervus kuhli : : : : : 174

6. The Para, or Hog-Deer—Cervus porcinus : ‘ ‘ 175

7. The Chital, or Indian Spotted Deer—Cervus axis : : : 179
Incertz Sedis—Cervus lepidus . : . : : ‘ 185

Cervus caspicus . ; , : 3 186

v. The Rucervine Group—Sub-Genus Poe : : 188

. The Swamp-Deer—Cervus duvauceli. : : ; ; 189

2. Se aaa Deer—Cervus schomburgki : : ; : 193

3. The Thameng—Cervus eldi ; : i : ; 195

a. Burmese Race—Cervus eldi typicus . ; ; ‘ 2.00

b. Siamese Race—Cervus eldi platyceros . : 5 : : 200

4. The Siwalik Swamp-Deer—Cervus sivalensis (extinct) . ; ; 201

IV. THe Munryacs—Genus Cervulus . : ; : ; : 202
1. The Indian Muntjac —Cervulus muntjac : : : : 203

2. The Tibetan Muntjac—Cervulus lachrymans . ; é : 208

3. The Chinese Muntjac—Cervulus reevesi : : ; : 208

4. The Tenasserim Muntjac—Cervulus fee ; t 209

5. The Hairy-fronted Muntjac—Cervulus crinifrons : : 3 210

V. Tue Turrep DEER—Genus Elaphodus ; : : 21D,
. The Tibetan Tufted Deer—Elaphodus Pea: . 5 ; Di

2. Michie’s Tufted Deer—Elaphodus michianus . : Ag : 214

VI. THe Miocene Munryacs—Genus Dremotherium (extinct) . : : 215
1. Hensel’s Muntjac—Dremotherium furcatum : : 3 ‘ 2A

2. Kaup’s Muntjac—Dremotherium anoceros : : : ‘ 207

VII. THe Tuskress Munryacs—Genus Dicroceros (extinct) : : ; 218
The Sansan Muntjac—Dzicroceros elegans . : : é 219

VII. THe Warer-DEER—Genus Hydrelaphus . ] ‘ : 219
The Chinese Water-Deer—Hydrelaphus inermis . : : : 221

IX. THe Rors—Genus Capreolus : : : E : 2 222,
1. The European Roe— Capreolus vulgaris : ; : : 223

2. The Siberian Roe—Capreolus pygargus . : : : 227

3. The Manchurian Roe—Capreolus manchuricus . : : : 231

4. The Pliocene Roe—Capreolus cusanus (extinct) . : : F 232

X. THe Mitrou Deer—Genus Elaphurus : ; : : ; 233
Pere David’s Milou Deer—Elaphurus davidianus . ~ . , , : 236

G

XIV Contents

PAGE

XI. Genus AnoGtocuis (extinct) ; : : : : ; 238

1. The Ardé Deer—Anoglochis ardeus : ‘ : 5 ; 239

2. Sedgwick’s Deer—Anoglochis sedgwicki . : : : E 240

3- The Five-tined Deer—Anoglochis tetraceros : . Oe hat 242

XII. con AMERICAN DeEER—Genus Mazama . ‘ : : 243

. The Dorcelaphine Group—Sub-Genus Dao é ‘ : 248

. The Common Americ: Mazama americana. ‘ : 249

a. Virginian Rac ricana typica . : : : 252

b. Western Race—Mazama americana macrura . , : : 257

c. Florida Race—Mazama americana osceola : ; ‘ : 259

d. Sonoran Race—Mazama americana couesi : ; j : 261

e. Texan Race--Mazama americana texana . , : : 261

J Mexican Race—Mazama americana mexicana ‘ ; ; 261

g. Yucatan Race—Mazama americana tolteca . ‘ 5 ‘ 263

h, Central American Race—Mazama americana nemoralis : : 264

i. Colombian Race—Mazama americana gymnotis F ; : 266

J. Savanna Race—Mazama americana savannarum g : : 267

k, Peruvian Race—Mazama americana peruviana : : : 268
2. True’s Deer—Mazama truei . ; , t : : 268 |

3. Crooke’s Black-tailed Deer—Mazama crookei . ; : ; 268

4. The Mule-Deer—Mazama hemionus . : : : : 269

a. Vypical Race—Mazama hemionus typica " ; : A 275

b. Californian Race—Mazama hemionus californica ; ; ; 276

c. La Paz Race—Mazama hemionus peninsule . : ‘ ; 276

d. Western Desert Race—Mazama hemionus eremica . , . 277

4. The Black-tailed Deer—Mazama columbiana . : : ‘ 278

Incerta Sedis—Mazama spinosa . : ; ; d ; 280

Mazama similis . : ; , ‘ é 281

Mazama leptocephala. : : : ; 281

Mazama pusilla . ; , : 5 ; 281

Mazama sp. : ; , j ‘ 281

ii. The Guazu Group—Sub-Genus Blastoceros : ; : : 282

The Marsh-Deer—Mazama dichotoma . ; ‘ , : 283

The Pampas Deer—Mazama bezoartica ; ; ; . 287

3. The Lujan Marsh-Deer—Mazama brachyceros (extinct) : ; 291

4. The Buenos. Aires Marsh-Deer—Mazama ultra (extinct) , : 292

5. Ameghino’s Marsh-Deer—-Mazama fragilis (extinct). : : 2.93

iii. “(he Guemal Group—Sub-Genus Xenelaphus , 7 ; ; 293

. The Peruvian Guemal—Mazama antisiensis . ‘ ; : 295

The Chilian Guemal—Mazama bisulca . 7 ; , ; 2.96

3. The Plistocene Guemal—Mazama senelitica (extinct) . ; 2 298

The Typical, or Brocket Group—Sub-Genus Mazama_. ; / 298

The Red Brocket—Mazama rufa . ’ ; ; : 300

Contents

XII. THe American DEER—continued
2. The Streak-eyed Brocket—Mazama superciliaris
. The Black-faced Brocket—Mazama tema
. The Central American Brocket—MJazama sartorii
. The Wood Brocket—Mazama nemorivaga
. The Peruvian Brocket—MJazama tschudii

WAN FW

The Pigmy Brocket—dazama nana
Incertz Sedis—azama inornata

Mazama aurita .

XIII. THe Pupus—Genus Pudua ;
1. The Chilian Pudu—Pudua pudu
2. The Ecuador Pudu—Pudua mephistopheles

B. THe ABerrant DeeR—Sub-Family Moschine

XIV. THe Musxs—Genus Moschus 2
1. The Himalayan Musk—Aoschus moschiferus
2. The Kansu Musk—Jdoschus sifanicus

APPENDIX— Bibliography of the Cervide

INDEX

XV

301
302
528
393
395
395
306
306

3o7,
37)
308

399
310
201
355
3u/

325

Es Or Piva TES

PLATE

I.

2.

SCANDINAVIAN REINDEER. Froma male in the London Zoological Gardens To face page 38

Erk. From a male in the London Zoological Gardens

Duke or Beprorp’s Deer. From an immature male in summer pelage,
recently living at Woburn Abbey

KasHmir Hancut. From a male specimen in the British Museum

5. THorotp’s Deer. From a male in the British Museum

Eee

IO.

Tie

Tr

1
I4.
15.

16.

iF

18.

Avtrat Wapiri. From a male in winter pelage in the Menagerie at

Woburn Abbey

. Mancuurian Sika. From males in winter pelage in the Menagerie at

Woburn Abbey

. Formosan Sika. From a male in winter pelage in the Menagerie at

Woburn Abbey

PEKIN Sitka. From immature specimens in very early winter pelage at

Woburn Abbey

InpIAN SamBAR. From a male and female in winter pelage at Woburn

Abbey

Maayan SAMBAR. From a young male in winter pelage recently living
at Woburn Abbey

Motuccan Rusa. From a male in winter pelage in the British Museum,
formerly living at Woburn Abbey

CuiraL. From males at Woburn Abbey .
Swamp-DeEeER. From a male specimen in the British Museum

S1aAMESE THAMENG. From a Hainan male in early winter pelage in the

London Zoological Gardens

‘TENASSERIM AND Hairy-FRONTED Munryacs. From the figures of the
type specimens
Micnte’s Turr—Ep DEER AND CHINESE Warer-DeEER. From male

specimens in the British Museum

European Ror. From male specimens in the British Museum, in summer

and winter pelage

»”

»”

»”

”

”

»”

”

»”

”

»”

»”

”

9

”

”

52

200

209

214

224

X

Vill List of Plates

PLATE

. Prkre Davin’s Mitou Deer. From a male living in the Menagerie at

19

Woburn Abbey

. Virointan Deer. From a male in the London Zoological Gardens

. Mure-Deer. Froma male in the London Zoological Gardens

. Pampas Deer. From a young male (the antlers added)
formerly living at Woburn Abbey

. PeruviAN GuEMAL. From a male in the British Museum

. CENTRAL AMERICAN Brocker AND Ecuapor Pupu.
in the British Museum

from Sante Fe,

From examples

»”

”

»

To face page 236

252

270

288
295

oh)

ra)
(9)

COM Dn PWD HE

ie FIGURES

. Group of Deer at Woburn Abbey .
. Another group
. Antlers of Caspian Red be

Map of Distribution of Deer

. Head of Chital 5

. Antlers of Woodland Reindeer

. Skull and Antlers of Woodland Reindeer

. Head of Newfoundland Reindeer

. Antlers of Barren-Ground Reindeer

. Frontlet and Antlers of Elk

. Skull and Antlers of Elk

. Antlers of Elk

. Elk at Woburn Abbey

. Skull and Antlers of Scott’s Elk

. Antlers of Red Deer from Germany

. Antlers of Red Deer from Scotland

. Group of Red Deer at Woburn Abbey

. Antlers of Red Deer from Moritzburg Castle
. Frontlet and Antlers of Red Deer from Exmoor
. A Red Deer at Woburn Abbey

. Head of Caspian Red Deer

. Head of Hangul 5

. Skull and Antlers of Shou .

. Head of East American Wapiti

. East American Wapiti

. Manchurian Wapiti Hind .

. Altai Wapiti

. Bokhara Deer

. Antlers of Japanese Sika :

. Buck and Doe of Manchurian Sika

. Head of Pekin Sika

. Fallow Deer at Woburn Abbey

. Head of Common Fallow Deer

. Head of Mesopotamian Fallow Deer :
. Skull and Antlers of Irish Race of Giant Fallow Dee

. Skull and Antlers of variety of German Race of Giant Fallow Deer

XX Text Figures

. Antlers of Irish Race of Giant Fallow Deer at various ages
. Frontlet and Antlers of Indian Sambar

. Frontlet and Antlers of Malayan Sambar

. Formosan Sambar . ;

. Skull and Antlers of Javan Rusa

. Javan Rusa

. Moluccan Rusa :

. Antlers of Moluccan Rusa at various ages .
. Undetermined Rusine Deer

. Head of Hog-Deer

. Hog-Deer in Summer Pelage

. Antlers of Chital at various ages

. Skull and Antlers of Chital

. Male and Female Chital

. Frontlet and Antlers of Swamp-Deer

. Skull and Antlers of Swamp-Deer .

. Frontlet and Antlers of Schomburgk’s Deer
. Frontlet and Antlers of Burmese ‘Thameng
. Group of Burmese Thameng

. Frontlet and Antlers of Indian Muntjac

. Head of Hairy-Fronted Muntjac

. Head of Michie’s Tufted Deer :
. Frontlet and Antlers of Hensel’s Muntjac .
. Head of Siberian Roe

. Antlers of Siberian Roe :

. Abnormal Antlers of Siberian Roe .

. Pere David’s Milou Deer

. Series of Antlers of Milou Deer

. Antlers of Ardé Deer

. Antlers of Sedgwick’s Deer

. Antlers of Five-Tined Deer

. Antlers of Virginian Deer .

. Frontlet and Antlers of Virginian Deer

. Florida Deer

. Head of Mule-Deer

. Another specimen of the same

. Antlers of Marsh-Deer

. Young Marsh-Deer ,

. Malformed Antlers of Marsh-Deer .

. Young Male Pampas Deer

. Antler of Lujan Marsh-Deer

. Antler of Ameghino’s Marsh-Deer .

. Musk Deer ; ‘

. Skull and Antlers of Caspian Red Deer

fae DE bk JOnSALE. BANDS

Aeris iOKYs OFS tik PAMILY CERVID As

PVR NP RODUC TORY
DEFINITION OF DEER

PoPuLaR and scientific zoology are by no means always in accord, and an
instance of this want of harmony is afforded by the different limitations of
the popular term “deer” on the one hand, and the scientific title Cervzde on
the other. Both, indeed, relate to the same group of animals, and are
therefore to a great extent synonymous, but whereas the chevrotains, or
mouse-deer, are popularly ranked as members of the deer tribe, they are
excluded by naturalists from the family Cervide. There is evidently,
therefore, some character, or characters, not apparent to the popular mind,
which the scientific naturalist regards of primary importance in the defini-
tion of the family Cervide. And it is accordingly necessary to come to
a definite conclusion as to the limits of that family. In other words, we
have to seek a definition of the term “deer” in its scientific sense.

This is by no means such an easy matter as might on first thoughts
seem probable. The males of the more typical members of the group,
such as the red and fallow deer, are furnished with the graceful branch-
ing and deciduous appendages now generally known as antlers. And if
these were present in all (as they are in most) deer, there would be no
difficulty at all in formulating a concise and distinctive definition of the
family. These appendages are, however, lacking in the Chinese water-
deer, which is in every respect a true deer, and they are also wanting in the
more aberrant musk, as they likewise are in a number of species now

B

2 Introductory

extinct. It is manifest, therefore, that antlers, although very important,
are not alone sufhcient to define the Cervide. More especially is this the
case when extinct types have to be reckoned with (and without these there
is no possibility of arriving at the true affinities of a group) ; for not only
are there antlerless deer, but there are other kinds in which these append-
ages were never shed, and seem to have been permanently invested with
the skin, or “velvet.” Nor is this all, for there were certain extinct
ruminants more or less intimately allied to the giraffes, whose heads were
crowned with branching bony appendages apparently agreeing in all
respects with the antlers of the modern deer, except that they were never
shed, and may have remained in the velvet throughout life.

As every one knows, deer are members of the great group, or order, of
hoofed mammals, or Ungulata. The more typical representatives of this
great order are divided into two minor divisions, or sub-orders, distinguished
from one another most readily by the structure of the feet. In the one
sub-order, or Perissodactyla, the toe corresponding to the middle (third)
finger or toe of the human hand or foot is symmetrical in itself, and, as in
the horse, may be the only one remaining. The living representatives of
this sub-order are the horses, zebras, asses, tapirs, and rhinoceroses ; none of
which are at all likely to be confused with deer.

On the other hand, in the second great sub-ordinal group, or Artiodactyla,
the two toes respectively representing the third and fourth fingers and the
corresponding toes of man are symmetrical to a line drawn between them,
and unsymmetrical in themselves. This type of foot-structure when most
fully developed constitutes the familiar ‘“‘cloven hoof.” In its extreme
development, as in the giraffes and many antelopes, only one pair of large
hoofs remain to each foot ; but in the majority of the group a smaller pair
of lateral hoofs exist, which, as in the reindeer and the pigs, may be
functional, or, as in the roes, may be very minute and quite useless.

Having established the fact that deer are members of the drtiodactyla,
we have to find their position in that extensive group, which likewise
includes hippopotami, pigs, peccaries, chevrotains, camels, giraffes, the
prong-buck, antelopes, goats, sheep, and oxen, together with a whole host
of extinct forms. Neglecting the latter, this can be best effected by a
process of elimination.

First, we may eliminate the hippopotami, pigs, and peccaries by the

Definition of Deer 3

structure of their teeth ; all these animals having incisor teeth in the front
of the upper jaw, while their molars, or cheek-teeth, are surmounted by low,
more or less conical, or trefoil-shaped tubercles. On the other hand, the
members of the other groups mentioned above resemble one another in that
the grinding surfaces of the crowns of their hinder cheek-teeth culminate in
half-cylinders, of which there are usually four in each upper, and two in
~ each lower tooth, with the exception of the last, where there are generally
three. With this type of cheek-tooth is associated a complex stomach,
with at least three distinct compartments, and the function of chewing the
cud, or ruminating. Consequently all the members of the assemblage
are commonly known as ruminants.

From this group may be eliminated the camels and Ilamas on account of
the cushion-like structure of their feet, the presence of at least one pair of
incisor teeth in the upper jaw, and the circumstance that the stomach has
but three compartments. In all the others there are no upper incisor teeth,
although the canines, or tusks, frequently persist, and may attain large
dimensions. The chevrotains, or mouse-deer, which resemble the camels
in the absence of either horns or antlers, are externally very like small
antlerless deer ; but they differ from all the existing members of that group
in that the stomach has but three compartments, and also by the circum-
stance that the lateral hoofs of the fore-feet have complete supporting bones
(the metacarpals) extending throughout the whole length of each side of the
cannon-bone, or the main bone of the lower segment of the leg.

There now remain for elimination the giraffes, forming the family
Girafide ; the prong-buck, which alone represents the Axtilocapride ; and
the antelopes, goats, sheep, and oxen, collectively constituting the Bovide,
or hollow-horned ruminants. Together with the deer, these three family
groups are characterised by the absence of upper incisor teeth, the presence
of four distinct compartments to the stomach, and the incompleteness or
absence of the lateral metacarpal bones supporting the front lateral hoofs and
their basal joints. The whole group of four families, which have many
other characters in common, collectively constitute the true ruminants,
or Pecora. It is here that the real difficulty in defining the deer tribe
begins.

The Gvrafida, so far as their living representatives are concerned, are
very Closely allied to the deer, from which they differ by the appendages of

4 Introductory

the skull taking the form of unbranched bony pedicles, which are at first
separate from the bones supporting them, and are permanently covered with
skin. Lateral hoofs, which are present in all deer, are completely absent,
as are also upper canine teeth. In the structure of the skull, and the
general absence of a gall-bladder to the liver, giraffes closely resemble deer.
Moreover, many extinct members of the giraffe family had large branching
appendages simulating the antlers of the elk, but these were not periodically
shed, and may have been covered either with skin or with deciduous
horny sheaths. Several of these animals had two pairs of horns, either
rising separately from the skull or springing from a common base ; and in
this respect they were unlike any known deer. Nevertheless, the kinship
between the deer and the giraffes is evidently very close.

The prong-buck— Antilocapra—is easily differentiated from the deer
by the appendages of the skull forming simple bony cores clothed with a
forked horny sheath which is shed annually. A very similar type of
appendage characterises the Bovid@, the difference being that the horny
sheath is not forked, and is never shed. With the exception of certain
domesticated races, and some extinct forms, such horns are present in the
males, at least, of all species ; and thus at once serve to distinguish between
this family and the Cervide. In nearly all cases the liver has a gall-
bladder ; upper canine teeth are invariably absent ; and when any traces of
the lateral metacarpal bones remain, their lower extremities are always
wanting. The muzzle may be either naked or hairy ; and a gland on each
side of the face below the eye may or may not be present.

Coming to the Cervide, we find that in all existing forms, with the
exception of the Chinese water-deer and the musks, antlers (of which the
nature is explained below) are present in the males, but are absent in the
females of all the species except the reindeer. Save in the musk, there is a
face-gland below the eye, and in the skull a vacuity in this region between
the lachrymal and nasal bones ; both these being absent in a large number
of the Bovide. The lateral hoofs are invariably retained ; and, except in
the muntjacs, these lateral hoofs are always supported by the usual three
joints, or phalanges, of which the first and second are always wanting in the
Bovide. Moreover, the lateral metacarpal bones are always represented by
either their upper or lower ends, the latter of which are never found in the
Bovide. With the exception of the reindeer, some portion of the muzzle

Antlers ie

is devoid of hair, although in the elk the naked area is reduced to a very
small patch. Upper canine teeth are frequently present, and in those
species devoid of antlers attain large dimensions. Lastly, with the exception
of the musks, the liver has no gall-bladder. Certain characters connected
with the internal reproductive organs of the female need not be referred to
on this occasion.

The deer tribe may accordingly be concisely defined as Pecora presenting
the following characters. Either antlers are present in the male, or when
these are absent the upper canines are large and sabre-like, and the lateral
metacarpal bones are represented by their lower extremities. This defini-
tion will include not only the living, but also most of the extinct forms,
although in some of the latter the lateral metacarpal bones not only retain
their lower ends, but are complete in their entire length. At the same
time it must be mentioned that when we have to deal with the earliest and
most generalised forms we find some of these presenting characters common
to the Bovide, Girafide, and Tragulide (chevrotains) ; but this is only
what we should expect to be the case in the common ancestral forms of
all these groups.

Antlers.—As the most characteristic feature of the deer tribe, special
attention must be directed to the appendages which are best termed antlers.
Many writers call these appendages horns, and probably this is etymologi-
cally correct, since it appears that the word “antler” is derived from the
Old French anfoi/ler, a derivative of the Late Latin antoculorum; this term
having been originally applied to what is now called the brow-tine. Still
it is so convenient to have a distinctive term for the cranial appendages of
the deer, as distinct from the sheath-covered horns of the Bovide, that the
use of the word “‘ antler” in this sense is most desirable.

Antlers are supported on a pair of solid bony processes, or pedicles,
arising from the frontal bones of the skull, of which they form an inseparable
portion ; and if in a fully adult deer these pedicles be sawn through, they
will generally be found to consist of solid, ivory-like bone, devoid of per-
ceptible channels for the passage of blood-vessels. The pedicles are always
covered with skin well supplied with blood-vessels ; and in young deer, or
those in which the antlers have been comparatively recently shed, the
covering of skin extends over their summits, when they appear as longer
or shorter projections on the forehead, according to the species. When the

6 Introductory

first or a new antler is about to be formed, the summits of these pedicles
become tender, and bear small velvet-like knobs, which have a high tempera-
ture, and are supplied by an extra quantity of blood, which commences to
deposit bony matter. This deposition of bony matter progresses very
rapidly, and although in young deer and the adults of some species the
resulting antler merely forms a simple spike, or a single fork, in full-grown
individuals of the majority it assumes a more or less complexly branched
structure. All this time the growing antler is invested with a skin clothed
with exceedingly fine short hairs, and is most liberally supplied with
blood-vessels ; this sensitive skin being called the velvet. Towards the
completion of its growth a more or less prominent ring of bone, termed
the burr or coronet, is deposited at its base just above the junction with
the pedicle ; this ring tending to constrict the blood-vessels, and thus cut
off the supply of blood from the antlers.

As some diversity of opinion exists with regard to the state of the antler
after the cutting off of the supply of blood to the velvet, two some-
what opposing statements may be quoted. On page 431 of the article
“Mammalia” in the Encyclopedia Britannica (gth edition) Sir W. H. Flower
writes as follows :—‘*‘ When the growth of the antler is complete, the
supply of blood to it ceases, the skin dies and peels off, leaving the bone
bare and insensible, and after a time, by a process of absorption near the
base, it becomes detached from the skull and is shed. A more or less
elongated portion, or pedicle, always remains on the skull, from the summit
of which a new antler is developed.”

On the other hand, Mr. A. Gordon Cameron, writing in the Fie/d
newspaper of 22nd August 1896, makes the following observations :—‘“‘ A
stag’s antlers, though commonly described as horns, are not horns in the
physiological sense of the word, but bones of similar texture to the flat
bones of the skull, and developed by a similar process. Nor are they dead
bones, though commonly described as such, but living bones composed of a
hardened surface and a porous centre with blood circulating through it.”
And the same gentleman and other writers have adduced instances where
the antlers of a deer have bled when sawn off above the pedicles. It is
quite true that internally antlers display a honey-comb or cancellous
structure ; but the fact that in some instances the pedicles are formed of
impermeable ivory-like bone, seems to preclude the possibility of there

Fic. 1.—Group in one of the Paddocks at Woburn Abbey, including Pekin Sika, Red Deer, Caspian
Red Deer, Chital, and Virginian Deer, together with Mouflon. From a photograph by the
Duchess of Bedford,

Fic. 2.—A second group in the same Paddock, including Caspian Red Deer, Chital, and Virginian Deer.
From a photograph by the Duchess of Bedford.

8 Introductory

being any important supply of blood from the skull to the core of the
antler ; and the description of antlers as dead bone seems therefore justified.
On the other hand, I have seen recently sawn fallow deer antlers which
bled freely from the interior.

When the antlers are freed from the velvet—a process usually assisted
by the animal rubbing them against tree-stems or boughs—they have a
more or less rugose surface, owing to the grooves formed in them by the
nutrient blood-vessels. Although a few living species have the antlers in
the form of simple spikes in the adult male, in the great majority of species
they are more or less branched ; while in some, like the elk and fallow
deer, they expand into broad palmated plates, with tines, or snags, on one
or both margins. In the antlers of the red deer group, which form the
type of the whole series, the following names have been applied to their
different component parts and branches. ‘The main shaft is termed the
beam ; the first or lowest tine the brow-tine ; the second the bez-tine ; the
third the trez-tine, or royal ; and the branched portion forming the summit
the crown, or surroyals. But the antlers of all deer by no means conform
to this type; and in certain groups other names have to be adopted for the
branches.

As already said, the antlers of young deer are in the form of simple
spikes ; and this form is retained in the South American brockets, although,
as shown later on, the simple antlers of these deer appear due to degenera-
tion, and are not primitive types. Indeed, no living deer shows such
primitive spike-like antlers in the adult, and it is doubtful whether such a
type is displayed by any known extinct form, although many have a simple
fork. And, on the whole, there is a regular evolution of complexity in the
antlers of the deer as they are traced upwards from the middle portion of the
Tertiary period towards the present and immediately antecedent epochs.
In the deer of the sambar group, where the antlers never advance beyond
a three-tined type, the shedding is frequently, if not invariably, very
irregular ; but in the majority at least of the species with complex antlers
the replacement is annual, the new appendages attaining their full develop-
ment immediately before the pairing-season. In such species there is a
more or less regular annual increase in the complexity of the antlers up to a
certain period of life, after which they begin to degenerate, or “ go back.”
Of four well-fed red deer kept in captivity at Blair-Athole, the following

Antlers )

record of their antler-development is quoted by Mr. Gordon Cameron :—
“One of the stags continued to improve till he was nine years old, after
which he remained stationary till fifteen, when he became diseased and was
shot. Two of them went on improving till they were eleven years old,
and then remained stationary till they were sixteen, when they became
diseased and were shot ; the fourth went on improving till he was thirteen
years old, when he was killed in a fight.”

Peculiarities in antlers are reproduced year after year in the same stag
with remarkable regularity ; red deer or wapiti with badly formed brow-

Fic. 3.—-Antler of Caspian Red Deer, from a Hungarian specimen in the collection of Viscount
Powerscourt. Counting from the skull upwards, the first tine is the brow, the second the bez,
and the third the trez, above which come the surroyals, or crown.

tines, or fallow deer with an extra tine, reproducing such singularities with
extreme constancy. This, among many other more convincing reasons,
indicates the importance of antlers as a basis of classification of the deer
tribe. And although these appendages were to a certain extent put in the
background by the late Sir Victor Brooke, Mr. A. Gordon Cameron
seems to have successfully proved that they have the right to a very fore-
most place in the scheme of classification. As they are the most easily
recognised feature in a deer’s organisation, their adoption as a basis of
g

ae) In troductory

systematic classification largely simplifies the study of what is in many
respects an exceedingly difficult group of animals.

With regard to the origin of antlers, it has already been. mentioned
that the earliest known deer were devoid of these appendages, and that
when they first made their appearance they were of the simply forked type.
In some at least of these primitive types the entire antler seems to have
been very similar to the pedicle, from which it was not separated by a
distinct burr, and the whole structure was probably covered permanently
with skin, as are the horns of the giraffe at the present day. Such a skin-
clothed structure would, however, obviously have been extremely exposed
to injury ; and the development of a burr led to the cutting off of the
supply of blood from the portion above the ring, and the evolution of the
antler in its present form. But such a structure is incapable of any repara-
tion of an injury, and the next stage in the development was in all prob-
ability the periodical shedding and reproduction. As this reproduction in
the species where the shedding is annual is, as already said, always com-
pleted at the commencement of the pairing-season, it is a natural inference
that, at the present time at least, antlers are largely connected with the
sexual function. And further evidence of this is afforded by their almost
universal restriction to the males. It has indeed been very generally con-
sidered that stags with the largest antlers were those which succeeded in
obtaining the leadership of the herd and the mastery of the females. But
some years ago Mr. Caton remarked that among a herd of wapiti in
captivity in Ottawa the mastership was held by a large stag with small
antlers, while a smaller stag with much larger antlers was kept at a distance.
And more recently Mr. A. Gordon Cameron has stated that among Scotch
red deer antlerless or “ bald” stags, though wielding no weapons and dis-
playing no ornament, prove to be in all respects a match for their armed
and theoretically more attractive rivals, so that they are usually master-
stags, and sometimes acknowledged monarchs of large herds.

This is certainly very strong evidence against the theory that sexual
selection can be regarded as sufficient to account for the origin and purpose
of the frontal weapons of this group of ruminants. And in the opinion of
the writer last quoted their original purpose was as weapons of offence
against the attacks of beasts of prey ; such weapons having at first been
common to both sexes, but subsequently lost, as a normal feature, in the

Pelage II

females of all species of the deer tribe except the reindeer. As this subject
is one which obviously admits of a large amount of argument, and as the
object of the present volume is to record facts rather than to discuss
theories, it must be dismissed with this brief mention.

Other Structural Features of Deer.—The general external appearance
of deer is too well known to require any special mention, although it
‘may be observed that when at rest they all carry the head well above
the line of the back, that the ears are always well developed and
frequently large, and that the tail never reaches below the hocks, 1s
frequently very short, and may be practically wanting. Very generally
the epithet ‘“‘ graceful” is applied to these animals collectively ; but although
its application is perfectly justified in the case of such species as the
Virginian and the fallow, it is certainly inapplicable to formis like the
reindeer, the elk, and Thorold’s deer, the second of which is one of the
most ungainly of all animals, while the third is decidedly a clumsily built
creature. And, as a whole, it seems that in respect of general gracefulness
of form and appearance the deer must give place to the antelopes and
gazelles, although the size and beauty of their antlers will always render
them a most attractive group. In point of bodily size there is every grada-
tion between the maximum height, as exemplified by the elk (65 feet),
and the minimum, as represented by the Chilian pudu (134 inches).

In respect to their coat and colour several interesting peculiarities are
presented by the deer tribe. All are well clothed with hair, which is
generally more or less coarse and stiff, although frequently finer in summer
than in winter; except in the reindeer, it has no resemblance to wool,
and, save in that species, there is little or no true under-fur. In most, if
not all, species two annual changes of the pelage take place, the one in
spring and the other in early autumn. In the spring change the process
may be either sudden or protracted ; and in the wapiti, while this change
is rapid, the autumnal one is a slow process. Writing of this species, Mr.
Caton says: “The winter coat is all detached so nearly together that if
the hairs were dropped off so soon as they are loosened, the animal would
appear almost naked, so short would be the new coat. But the inner coat
of fur has during the winter become felted together, embracing and confin-
ing the long coarse hairs, so that they cover the animal as with a blanket,
after considerable portions have become loosened, thus allowing the young

12 Introductory

hairs to obtain some length before their predecessors are gone. Indeed,
the old coat does not in fact drop off, as in ordinary cases, but is torn away
in large patches, by contact with the shrubbery.” A precisely similar
process is observed in the shedding of the coat of the American bison.
When the process of changing is a slow one, in many species the red coat
of summer may be seen showing through the gray one of winter, or vice
versa, giving to the animal a curiously mixed type of coloration. Not only
is the summer coat generally finer than that of winter, but in many species
the hairs themselves are different in structure, being thin, straight, and
compact, instead of thick, waved, and more or less pithy. In the deer of
temperate and cold climates the summer coat is frequently retained for a
very short period, often not more than three months, or even less.

Judging from the number of species which are spotted, either per-
manently or during the period that they are in summer pelage, and likewise
from the still more frequent occurrence of white spots in the pelage of the
fawns, as well as from the circumstance that the young of so many other
Ungulates are either spotted or striped with white, it may be inferred that
spotting was the original type of coloration in all the members of the group.
And as such spotting is generally associated with a ground-colour of some
shade of tawny or reddish, it may further be inferred that the same tints
characterised the ancestral forms. Asa rule, those species which are per-
manently spotted, such as the Indian chital, show but little change of colour
between the summer and winter coats; but in all the species which are
spotted only in summer, and in many of those which are uniformly coloured
at all seasons of the year, the summer coat retains the original fawn or
reddish tint, whereas the winter pelage tends to some shade of gray or dark
brown. In the former condition, as is well exemplified by the case of the
Virginian deer, the animal is said by hunters to be in the red, and in the
latter in the blue.

It is thus evident that in such species as show a marked difference between
the colour of the summer and winter pelage, it is the red summer coat that
retains the ancestral type of coloration, and that the gray or brown winter
coat is an acquired character of comparatively late development. And it is
obvious that those species, like the Japanese sika, in which the summer
coat is spotted, are one stage nearer the original type than those, like the
Virginian deer, in which the coloration is uniform at all seasons. A further

Coloration 13

step in the development of what may be termed the modern or specialised
type of coloration is presented by forms like the Malayan sambar, in which
the colour of the pelage is some shade of uniform dark brown at all seasons,
although spots are still retained by the fawns. And the culminating stage
is presented by the Indian sambar and the elk, in which the coat is uniformly
dark-coloured at all seasons and all ages. It is not a little remarkable that

-while this highly specialised type of coloration is presented by so many

members of the sambar group, all retain antlers of a comparatively simple type.

The general darkness of coloration noticeable in the sambar group, which
is as remarkable in the spotted Philippine deer as in the typical forms, appears
to be an instance of that tendency towards blackness, either universal or
individual, observable in many animals inhabiting moist and generally warm
districts." But, as exemplified by the case of the chital and swamp-deer,
this melanistic tendency need not affect all the members of a group or family
inhabiting the same or similar areas. Another example of the same tendency
is well exemplified by the gradual darkening of the coloration of the forms
allied to the Virginian deer as they approach the hot humid districts of
tropical America, where this dark pelage is retained throughout the year.
The dark coat of the elk is also developed in an animal essentially an
inhabitant of humid, although colder, districts. What led to the permanent
‘“‘melanism” of the dark-coated British variety of the fallow deer does not at
present admit of determination.

In those species of deer presenting a marked difference between the
coloration of the summer and the winter coat, there is not only a distinct
change in this respect at the time of assumption of each pelage, but minor
changes are going on almost continuously throughout the year. In con-
sequence of this the same animal looks different almost every month, if not
every week ; and the task of describing its true coloration becomes almost a
matter of impossibility. Indeed, in such cases a coloured portrait of the
animal is a true representation only for an extremely limited period ; and to
obtain an adequate idea of the gradual changes the creature ought to stand
for its portrait at least every fortnight. Fading of the colour of the hair,
especially on certain portions of the body, seems to play an important part
in such gradual changes; the light-coloured patches on the rump of most
members of the red deer group and of the roes exemplifying this fading.

1 See a paper on “ Melanism ” by Dr. E. Lénnberg, published in the Zoo/. Fahrbuch for 1897.

14 Introductory

These aforesaid light-coloured or white patches on the hind-quarters of
many species, such as the members of the red deer group, the sikas, the
fallow deer, and the roes during the winter, come under the category of
recognition marks ; and are in all probability for the purpose of enabling
the individuals of a herd to follow their leaders, when seeking safety from
danger in flight, with as much ease as possible. The case of the roes, in
which the white patch on the rump is fully developed in winter, but is
quite wanting at the first assumption of the red summer pelage, is a very
curious one, and one decidedly difficult of explanation. The does, however,
drop their fawns about May, and at this time keep apart from the bucks.
Now the fawns for a considerable period do not flee with the mother at
the approach of danger, but are concealed by her in covert while she makes
her escape alone. By the time the fawns are sufficiently grown to be able
to flee from danger by the side of their dams, the white caudal patch is
assumed, and the two sexes go about in company. ‘These circumstances
may afford an explanation of the disappearance of the white patch with the
assumption of the summer coat. In the common sika the hairs of the
caudal patch are capable of erection and expansion, thus increasing the size
of the white area; and the fallow deer, when alarmed, produces the same
effect, by raising the tail. In the Virginian deer and its allies there is no
such recognition mark when the animals are feeding ; but when starting off
in alarm these deer elevate the tail, the white under surface of which, with
the white on the inner sides of the buttocks, forms a most conspicuous
lozenge-shaped recognition mark. Although such marks are most con-
spicuous in the groups mentioned, they occur to a more or less marked
degree in many other members of the family, even the chestnut streaks on
the buttocks of the Indian sambar coming under the same category. It
will be obvious that these recognition markings, by rendering it more
conspicuous to enemies, are a disadvantage to the individual, and that they
are developed for the good of the species as a whole.

Colour-markings are, however, by no means the only aids with which
deer are provided for the purpose of assisting them in finding the where-
abouts of their companions. Of the “calling” or roaring of the adult
stags during the pairing-season mention will be made in the sequel. The
aids to recognition now to be referred to are glands. It has been stated
that almost every species of deer is provided with a pair of glands situated

Glands 15

on the face immediately below the eyes. In some kinds, like the muntjacs
and the Virginian deer, these glands are comparatively small, forming little
more than folds in the skin, and having only very small and shallow
depressions in the skull for their reception. In other kinds, however, and
especially in the various races of sambar, they attain an enormous develop-
ment, and rest in large and deep cavities in the bones of the skull. The
upper surface, or lid, of such a gland is capable of complete eversion,
leaving exposed the large cavity in the skull lined with glistening pink
mucous membrane ; and so peculiar is the appearance of the animals when
the glands are thus opened, that the Chinese have conferred upon the
Formosan race the designation of the four-eyed deer. As these glands are
opened most fully and most frequently during the pairing-season by the
stags, it is evident that they are largely connected with the sexual function.
The muntjacs have also a single gland situated on the forehead, and hence
termed the frontal gland.

In addition to these face-glands, most deer have interdigital or foot-
glands, situated between the two main hoofs of each foot, and doubtless
serving to leave a strong scent upon the ground over which their owner has
passed. There may also be either one or two glands situated on each hind
leg, the position of which is indicated by a tuft of hair longer than that
clothing the rest of the limb, and frequently also differing more or less
markedly in colour. The more commonly developed of these glands is
situated on the outer surface of the lower segment of the hind limb above
the foot, this segment being the one containing the cannon-bone, and
technically known as the metatarsus. Hence the gland and tuft are spoken
of as the metatarsal gland and tuft. In the red deer group this tuft forms
a patch situated generally high up on the cannon-bone, and not differing
markedly in colour from the hair of the rest of the leg. In the mule-deer
it forms an extremely elongated strip extending along the greater part of
the outer side of the cannon-bone, and of the same general coloration as
the shorter hair adjacent. On the other hand, in the Virginian deer the
gland forms a very small circular black patch near the lower extremity
of the cannon-bone, surrounded by a conspicuous fringe of long white
hairs. Not a little remarkable is the circumstance that in some smaller
southern forms closely allied to the Virginian deer this gland is totally
absent. In the sequel it is suggested that its presence or absence may

16 Introductory

be dependent upon whether the animals live in herds or merely consort
in pairs.

Less common is a gland situated on the inner surface of the hock, which
is anatomically equivalent to the ankle, or tarsus, of other animals. Hence
this gland and its tuft are termed the tarsal gland and tuft; its position
being well shown in the coloured plate of the reindeer, where it stands out
in conspicuous whiteness from the dark colour of the rest of the limb.

Lastly, the musks are peculiar in the possession by the males of a large
glandular pouch situated on the abdomen, and secreting the substance from
which these animals take their name.

As non-scientific persons frequently have very hazy ideas as to the
nature and function of glands (which in truth are not fully known even to
the scientific), the following extract on this subject, taken from Sir William
Flower’s above-mentioned article on the ‘“ Mammalia,” may be quoted im
extenso. ‘* Most mammals,” writes Sir William, “have special glands
situated in modified portions of the integument, often involuted to form a
shallow recess or a deep sac with a narrow opening, situated in various
parts of the surface of the body, and secreting odorous substances, by the
aid of which individuals appear to recognise one another, and probably
affording the principal means by which wild animals are able to become
aware of the presence of other members of the species, even at great
distances. Although the commencement of the modifications of portions
of the external covering for the formation of special secretions may be at
present difficult to understand, the principle of natural selection will readily
explain how such organs become fixed and gradually increase in develop-
ment in any species, especially as there would probably be a corresponding
modification and increased sensibility of the olfactory organs. Such in-
dividuals as by the intensity and peculiarity of their scent had greater power
of attracting the opposite sex would certainly be those most likely to leave
descendants to inherit, and in their turn propagate, the modification.”

Classification.—Very different views as to the best way of classifying the
members of the Cervide have obtained, and still obtain, among systematists ;
and the question, like others of a similar nature, depends largely upon the
individual bias of the particular writer. To some minds a system appears
to be more lucid when a number of divisions, based upon more or less
trivial characters, are adopted ; while to others greater clearness is obtained

Classification ny

when leading characteristics are alone taken as the ground of important
divisions, and minor differences are reckoned of comparatively small moment.
Neglecting earlier writers, three important syllabic classifications of the
family have been published in comparatively modern times ; the first two
of which followed the plan of making as many divisions as possible, while the
third relegated a large number of such divisions to a much lower grade of
importance. The first of these is one by the late Dr. J. E. Gray of the
British Museum, which appeared in the Scientific Proceedings of the Zoologi-
cal Society of London for the year 1850, and was expanded and otherwise
modified in two Catalogues subsequently published by the British Museum.
The second is that of the late Professor L. Fitzinger, which was published
in the Sitzungsberichte of the Vienna Academy of Sciences at intervals
ranging from 1873 to 1879. The third was written by the late Sir Victor
Brooke, and appeared in the Proceedings of the London Zoological Society
for 1878. Although it was published after the appearance of most of
Fitzinger’s memoir, the author does not appear to have seen the latter ; and
hence while it professes to give a complete synonymy of the various genera
and species, many names are omitted which ought to have appeared. It
also contains a rather high average of serious errors; and as it has been
copied by nearly all subsequent writers, such errors have not only been
perpetuated, but widely disseminated. Nevertheless, in spite of these
deficiencies, the paper is one of a high order of merit ; and its general
scheme of classification is the one followed in the present work, with such
modifications and amendments as seemed necessary or advisable. In the
case of genera, sub-genera, and species, numerous alterations in nomenclature
have been rendered necessary by the revolution which has of late years
taken place in this branch of zoological science ; and, in the opinion of
advanced advocates for such changes, there are doubtless others that might
have been made. The author is, however, a strict conservative in such
matters, and very loath to insert changes in names now in current use.

The following table exhibits the generic and sub-generic groups into
which the existing and some nearly allied extinct deer may be divided, with
their leading distinctive characters :—

A. Sus-Famity Cervin&.—Antlers, with one existing and some extinct exceptions,
present in the male; liver without a gall-bladder ; a face-gland and gland-pit.
D

18 Introductory

I. Genus Rancirer.—Lateral metacarpal bones represented only by their lower
extremities ; antlers present in both sexes, complex. Northern part of
Holarctic region.

I]. Genus Atces.— Lateral metacarpals as in preceding ; antlers (as in the
following genera) present only in the male, arising at right angles to the
median longitudinal line of the skull, and extending at first in the plane of
the forehead, after which they expand into a broad palmation margined
with snags. Northern portion of Holarctic region.

III. Genus Cervus.—Lateral metacarpals represented only by their upper ends.
Antlers arising at acute angles to the median line of the skull (as in the
following genera), at first projecting from the plane of the forehead, and
then continued upwards nearly in that plane, supported on short pedicles,
and furnished with a brow-tine, never regularly forked at first division,
but generally of large size, and with not less than three tines ; skull with-
out ridges on the frontals forming the bases of the pedicles of the antlers.
Upper canine teeth small, or wanting. Holarctic and Oriental regions.

1. Sub-Genus Cervus.—Antlers rounded, usually with five or more tines,
generally including a bez (second), and always a trez (third) ; pelage of
adult generally unspotted, with a large light-coloured disk surrounding
the tail ; young spotted. Hlarctic region.

i. Sub-Genus Pseudaxis.—Antlers smaller and simpler, four-tined, with a
trez (third), but no bez (second); pelage of adult spotted, at least
in summer, with a white area bordered by black in the region of the
tail, which is also black and white. South-eastern portion of Eastern
Holarctic region ; Europe in Pliocene.

iil. Sub-Genus Dama.—Antlers without a bez, but with a trez-tine, above
which the beam is more or less palmated, and generally furnished
with numerous snags; pelage of adult spotted in summer, uniform
in winter, with black and white markings in the region of the tail
similar to those of Pseudaxis ; young, spotted. South-western portion
of Eastern Holarctic (Mediterranean) region, but more widely spread in
Europe during the Pliocene epoch.

iv. Sub-Genus Rusa.—Antlers rounded, three-tined, with the bez- and trez-
tines wanting, and the beam simply forked at the summit; pelage
either uniform or spotted at all seasons. Oriental region.

v. Sub-Genus Rucervus.— Antlers flattened or rounded, without bez- or
trez-tine, the beam dichotomously forking, and one or both branches

again forked, so that the number of tines is at least four, brow-tine

RV:

VI.

Nelle

VIII.

TX:

XI.

Classification 19

forming a right angle or a continuous curve with the beam; pelage of
adult generally more or less uniform, of young spotted. Oriental region.

Genus Cervutus.—Lateral metacarpals as in Cervus ; antlers small, with a
brow-tine and an unbranched beam, supported on long bony pedicles, con-
tinued downwards as convergent ridges on the forehead ; upper canines of
male large and tusk-like. Oriental region.

. Genus ErapHopus.—Nearly related to the last, but the antlers still smaller,

with shorter pedicles and divergent frontal ridges ; upper canines of male
not everted at the tips. South-eastern portion of Eastern Holarctic region.

Genus DremoTHERIUM (Extinct).—Lateral metacarpals in some cases com-
plete, and in others apparently wanting ; antlers, when present, mounted
on long bony pedicles, with a very imperfect burr, simply forked, and
non-deciduous ; upper canines generally tusk-like ; cheek-teeth low-
crowned (brachydont). Eastern Holarctic region, from the Oligocene to
the Miocene periods.

Genus Dicroceros (Extinct).—Closely allied to the last, but with the
antlers of a somewhat more advanced type, no tusk-like upper canines,
and the cheek-teeth tall-crowned (hypsodont). Eastern Holarctic region
during the Miocene period.

Genus HypretapuHus.—Lateral metacarpals as in Rangifer; antlers wanting ;
upper canines of males tusk-like and growing from semi-persistent pulps ;
cheek-teeth tall-crowned (hypsodont) ; tail moderate. South-eastern portion
of the Eastern Holarctic region.

Genus Capreotus.—Lateral metacarpals as in Rangifer; antlers rather
small, without a brow-tine or sub-basal snag, dichotomously forked, with
the upper or posterior prong again forking ; tail rudimentary; vomer not
dividing posterior nasal aperture of skull. Eastern Holarctic region.

. Genus ELrapuurus.—Lateral metacarpals as in Cervus ; antlers large, with-

out a brow-tine or sub-basal snag, dichotomously forked, with the upper
prong of the fork curving forwards and dividing, and the lower prong
long, simple, and projected backwards, the beam making a very marked
angle with the plane of the face; tail very long ; vomer as in Capreolus.
Eastern portion of Eastern Holarctic region.

Genus Awnoctocuis (Extinct). — Lateral metacarpals unknown ; antlers
large, without a brow-tine, but with a sub-basal snag, above which the
beam is dichotomously forked, one or both prongs of the fork again
dividing once or more, and the whole antler forming a marked angle with

20 Introductory

the plane of the face; vomer unknown. South-western portion of Eastern

Holarctic region.

XH. Genus Mazama.—Lateral metacarpals as in Rangifer ; antlers very variable
in size, forming a marked angle with the plane of the face, without a brow-
tine ; when consisting of more than a simple prong, dichotomously forked,
frequently with a sub-basal snag, and always with the lower prong of the
fork projected from the front edge of the beam, in some cases the lower,
in others the upper, and in others both prongs again dividing ; tail long ;
tarsal gland generally present; metatarsal gland very variable, both as
regards presence and position ; vomer dividing the inner aperture of the
nostrils in the skull into two distinct chambers. | Western Holarctic,
Sonoran, and Neotropical regions.

1. Sub-Genus Dorcelaphus.—Antlers large and complex, with a sub-basal snag,
and the lower prong more or less developed at the expense of the upper
one; metatarsal gland usually present ; tail long or moderate, and hairy
below ; face very long and narrow ; the face-gland small, and the gland-
pit in the skull of moderate extent ; no upper canines ; size generally
large. Western Holarctic, Sonoran, and northern portion of Neotropical
region.

uu. Sub-Genus Blastoceros.—Antlers large and complex, without a sub-basal
snag, and the upper prong more developed than the lower one; metatarsal
gland absent ; tail short ; face moderately long ; face-gland and gland-
pit well developed ; upper canines usually present in male. Size large
or rather small. Neotropical region.

iii. Sub-Genus Xenelaphus.— Antlers small and simple, forming a single
dichotomous fork ; metatarsal gland absent ; tail short ; face moderately
long ; face-gland and gland-pit well developed ; upper canines present
in both sexes. Size medium. Western and southern portions of
Neotropical region.

iv. Sub-Genus Mazama.—Antlers in the form of simple unbranched spikes ;
metatarsal, and in one case also the tarsal, gland absent ; tail very short ;
face elongated ; face-gland small and gland-pit deep and triangular ; hair
of face radiating from two whorls ; upper canines sometimes present in

old male. Size small. Neotropical region.

XIII. Genus Pupva.—Skull and metacarpals generally as in Mazama ; size very
small; hair coarse and brittle ; antlers in the form of short, simple spikes ;
cannon-bones very short; tail very short or wanting; no whorls in the

hair of the face; face-gland moderately large, and gland-pit deep and

|

Classification 2I

oval ; tarsal and metatarsal glands wanting ; ectocuneiform bone of tarsus
united with the naviculo-cuboid. Neotropical region.
B. Sus-Famity Moscuin.—Antlers wanting in both sexes; liver furnished with
a gall-bladder ; no face-gland or gland-pit.

XIV. Genus Moscnus.—Hair coarse and brittle ; upper canines of male very
long ; no tarsal or metatarsal glands or tufts; lateral metacarpals repre-
sented by their lower extremities ; lateral hoofs very large ; tail very short ;
naked portion of muzzle extensive ; male with a large abdominal gland.

It will be seen from this table that the number of generic groups
adopted is comparatively small, but that many of these are divided into
sub-genera. The possession of one important common feature, such as the
presence of a brow-tine to the antlers in the members of Cervus, and of the
division of the hinder aperture of the nostrils in the skull of the American
deer by the vertical plate of bone known as the vomer into two chambers,
is regarded as entitling most or all of the forms in which it occurs to be in-
cluded in one genus. Minor differences, on the other hand, constitute the
basis of division into sub-generic groups.

Very similar considerations govern the grouping into species and sub-
species, or races. For instance, when we have two members of a genus as
distinct from one another as is the red deer from the wapiti, they are
regarded as forming separate species. But the wapitis of Asia differ from
the American forms, and if these were regarded as species, a distinction
higher than specific would be required between the red deer and the
wapiti. Consequently the Asiatic and American wapitis are regarded as
sub-species, or races, of a single widely spread and variable species. In like
manner, the various deer more or less nearly resembling the Indian sambar
are Classed as races of one species; as are those akin to the Virginian deer
of another. Should the system here adopted not be found acceptable to
any students of the group, it can readily be modified into a more complex
one. For instance, the Cervine and Moschine may be raised to the rank of
families, when several of the generic groups will form the types of sub-
families. The sub-genera will then stand as genera; and the larger sub-
genera, such as Cervus proper, will have to be divided into groups which
will then possess sub-generic value. ‘The red deer will then represent one
sub-genus and the wapitis a second ; while the Asiatic and American forms
of the latter will represent as many species.

22 Introductory -

Distribution.—In the study of any group of animals their distribution
over the surface of the globe forms nowadays one of the most important
factors. And distribution may be divided into two sections, namely, the
present or geographical distribution, and the past or geological distribution.
As regards the former of these, a few words are necessary to explain the
divisions into which the world may be parcelled out from the distribution
of mammals. First, there are three primary divisions respectively
named the Notogwic, Neogwzic, and Arctogezic realms. The first of these,
with which we have but little concern in this volume, includes New
Zealand, Australia, New Guinea, New Ireland, New Caledonia, etc. The
second contains the Neotropical region, which embraces the southern
portion of Mexico, Central and South America, and the West Indies.
The third, and by far the largest realm, consists of the rest of the world,
and is divided into several regions. These include the Malagasy region, as
represented by Madagascar and the Comoro Islands; the Ethiopian region,
embracing Africa south of the Sahara; the Oriental region, containing
India, Southern China, and the Malayan countries, the islands of Celebes
and Timor, being, on the whole, best placed here, although their faunas
show some affinities with that of Australia; the Holarctic region, embrac-
ing the greater part of Asia, Europe, North America, and Africa north of
the Sahara ; and the Sonoran region, which contains all the southern parts
of North America not included in the Neotropical. The great Holarctic
region may be divided into an eastern and a western division, the former
embracing all the Old World countries constituting the region, and the
latter such part of North America as comes within its limits. From the
eastern division of the Holarctic some writers cut off Northern Africa and
the other Mediterranean countries, together with those forming the
northern boundary of the Oriental region, to constitute a separate Mediter-
ranean region.

As regards the geographical distribution of the deer tribe, the most re-
markable fact is their entire absence from the whole of the Ethiopian region.
By far the greater number of generic and sub-generic types are confined
to the Old World, which is likewise the habitat of the majority of the
species. The occurrence of deer without antlers, and also of forms allied
to the muntjacs in the Oligocene and Miocene Tertiary strata of Europe, and
likewise of others possessing antlers of the sikine and perhaps the rusine

Distribution 28

type in the Pliocene of the same area, points to the conclusion that the Old
World was the original home of the types of deer by which it is now
populated. And it may further be considered certain that the wapiti is a
comparatively recent immigrant into North America by way of Bering
Strait. Not improbably the elks are likewise immigrants into the New
World from the Old, although generalised types occur in the later Tertiary
deposits of both areas. With regard to the place of origin of the reindeer,
there is no decisive evidence available. [The most primitive of the brow-

"e

Ce i inh iat eet a ea as

: ’ Q “ ra —
ty Ory : , ‘, or id
AES op, ales Bie OG AO Ln
© > Tn EER
4, 4)
. .
°

maa ee ee
De:
i

Fic. 4.—Diagrammatic Map of the distribution of Deer. The oblique shading indicates the approximate
range of the Reindeer and Elk ; the vertical that of the sub-genus Cervus, and in the Old World
also of the Sikine and Damine groups, as well as of the Roes ; the cross-shading that of the Rusine
group and Muntjacs ; and the horizontal that of the American Deer.

antlered Old World deer are undoubtedly the muntjacs, sambars, chital, and
sikas, and there is proof that most or all of these existed in the European
Tertiaries. ‘Their existence in the Oriental region, and some adjacent
parts of the Eastern Holarctic region, where species of the red deer group
are wanting, is one out of many instances of the survival of ancient forms of
life in these areas. That the fallow deer group is a specialised off-shoot
from the sikas, may be asserted with considerable confidence; and the red
deer group, characterised by the speciality of generally developing a bez-
tine, may likewise be regarded as descended from the latter. Hence all

24 Introductory

the brow-tined deer (Cervus, Cervulus, and Elaphodus), which agree in the
retention of the upper extremities of the lateral metacarpal bones, are
essentially Old World types. .

Much more difficulty exists with regard to the relationship of the fork-
antlered section, which includes the roes (to which the Chinese water-deer
appears nearly related), the Asiatic milou deer, the extinct Old World genus
Anoglochis, and the modern American deer. These present two distinct
types of metacarpal structure, namely, one in which the upper ends of the
lateral bones are retained, and another in which the lower ends remain ; the
condition existing in the extinct forms being unfortunately unknown.
Assuming Mr. A. Gordon Cameron to be correct in regarding all the antlers
of the forked type as indicative of a common ancestry, it is obvious that
in such ancestors the lateral metacarpals must have been complete.

With regard to the roes, there is no definite information beyond the fact
that they date from the earlier portion of the Pliocene epoch ; but they are
doubtless Old World types, and not improbably trace their ancestry to
some of the extinct hornless muntjacs (Dremothertum), from which both
types of antlers may have originated. ‘Their metacarpal structure is the
same as in the Americandeer. Nothing definite can be said with regard to
the milou deer, in which the opposite type of metacarpal structure obtains,
and the extinct Anoglochis, except that both are of Old World origin.

The American deer are usually regarded as the descendants of the two
extinct North American genera Blastomeryx and Cosoryx, the former of
which is regarded as the descendant of the extinct European Dremotherium.
Even on this view, the American deer are ultimately of Old World origin.
But it is difficult to believe that antlers of such essentially similar general
structure as are those of the modern American deer and the extinct European
Anoglochis have originated independently ; and there seems, therefore, to be
a considerable probability that the American deer have taken origin from
an Old World type allied to or identical with Anoglchis. ‘The question is
again discussed in the sequel, but it may be mentioned that Cosoryx is con-
sidered to be the ancestor of the American prong-buck (Anti/ocapra), and
if this be correct, is it probable that it also gave rise to the American deer ?

Before proceeding further, it may be well to mention that the geo-
logical formations of the Tertiary period, or those newer than the Chalk,

! See Scott and Osborn, Bu//. Mus. Harvard, vol. xx. p. 76 et seq. (1890).

Distribution 2s

are divided, in descending order, into the Plistocene, Pliocene, Miocene,
Oligocene, and Eocene epochs.

Whether they originated in the Old World or in North America,
the American deer passed into South America in the early portion
of the Pliocene epoch, when a land connection was first established wa
the Isthmus of Panama between the northern and southern halves of the
New World, and a large number of animals belonging to an essentially
northern type poured into South America, where they became mingled
with the proper indigenous fauna of that region. In the earlier
Tertiary formations of South America, which contain numerous fossilised
remains of armadilloes, of the allied gigantic extinct glyptodonts, of
ground-sloths, of American monkeys, and of various peculiar groups of
Ungulates, those of deer are entirely wanting ; whereas when we came to
the Pliocene epoch the latter are abundant.

To revert to the Old World, great difficulty exists in giving any
thoroughly satisfactory explanation of the entire absence of deer from
Africa south of the Sahara, more especially as remains of species of an Oriental
type occur in the later Tertiary deposits of India. Typical deer of the red
deer group are, however, wanting in the Indian deposits, as they are in India
at the present day, and as India seems to have been the great distributional
feeder of Ethiopian Africa by way of Syria, it is easy to see why there are
no members of that group south of the Sahara. With regard to the absence
of Oriental types of deer in Ethiopian Africa, it is as difficult to discover any
valid reason why these should have continued to flourish in the Oriental
region since the Pliocene epoch without having entered Africa, as it is to
explain why giraffes, hippopotami, and ostriches, which formerly inhabited
the former area, should have completely disappeared from it to find a per-
manent home in the latter. The North African red deer of course effected
an entrance into the north-western corner of Africa by means of a land-
bridge connecting that area with Spain, as did likewise the fallow deer ; the
Sahara presenting an impassable barrier to their progress southwards.

Temperature seems doubtless to have been a factor of some considerable
importance in the distribution of the various groups of the family. The
members of the red deer group, for instance, have a general distributional
area situated approximately between the fortieth and sixtieth parallels of
north latitude, although passing somewhat to the north of this zone in

E

26 Introductory

Norway, and south in North-Western Africa. The elk and reindeer, although
impinging on the northern limit of the red deer group and in certain
districts overlapping it, are characteristic of a colder zone, where Rangifer is
practically circumpolar. On the other hand, the Oriental deer of the
rusine and rucervine groups are essentially inhabitants of hot countries, and
are abundant within the tropical zone. To a certain extent the sikas are
intermediate in this respect between the rusas and the red deer, occupying
to a large degree a zone between the two latter, but also intruding into the
area ofeach. Their occurrence in the European Pliocene suggests, however,
that comparatively warm regions formed their original habitat. The
American deer range between latitude 40° north and the extreme south of
the continent, and are accordingly able to stand excessive heat and a con-
siderable amount of cold. Among these, the brockets are mainly restricted
to hot climates, although one is found high up on the Cordillera ; while the
guemals are exclusively inhabitants of cold regions. The dorcelaphine
section would seem to have been originally a temperate form, as the southern
races allied to the Virginian deer are all small and apparently degraded
types. On the other hand, from the present distribution of the marsh-deer
and the number of allied extinct species in the Tertiary deposits of Northern
Argentina, the blastocerine group appears to prefer a hot climate.

As already observed, remains of deer more or less closely allied to existing
types are met with throughout the formations of the Plistocene and Pliocene
epochs ; some of those from the former, such as Sedgwick’s deer and the Irish
deer, attaining a development of antler unequalled by any living species. As
we descend in the Pliocene the antlers tend to become smaller and simpler ;
and the distribution of the various groups is also found to differ considerably
from that obtaining at the present day. When we reach the Miocene, all
the deer are small, and their antlers never attain a development beyond that
of a simple fork; while some of them were entirely devoid of these
appendages. In the Upper Oligocene of America there are a few forms
referred to the family in which the antlers, although simply forked, seem
to have been clothed with skin and were permanently retained; but the
Upper Oligocene deer of Europe were mostly devoid of antlers, and apparently
furnished with long upper tusks, or canines, while the lateral metacarpal
bones seem to have been complete. The earliest known form which can be
in any way afhliated to the deer tribe is a small ruminant from the Middle

Habits 27

Oligocene marls of Ronzon, in the department of Puy-en-Velay, France,
known as Ge/cus. In this primitive little Ungulate the skull was devoid of
either horns or antlers, and the hinder cheek-teeth had very short crowns,
with low cusps and shallow depressions, but it is not improbable that upper
incisor teeth had disappeared. The lateral metacarpals and metatarsals
formed complete splints of bone, and in the hind limb the two median
metatarsals had coalesced to form a cannon-bone, although the corresponding
metacarpals remained distinct, as they do in the modern chevrotains. It has
been suggested that in this extinct ruminant we have the ancestral type of
both the chevrotains and the deer; and, in any case, it serves to show that
as we recede in time we depart further and further away from the modern
specialised types of the ruminant group.

Habits.—Since a considerable amount of space is devoted in the sequel
to the habits of the individual species, a few observations will suffice on those
of the group as a whole. Like most large family groups, the deer tribe is
essentially an adaptive one, although in this respect it is decidedly inferior
to the Boviaz, many members of which have established themselves in desert
regions, where the Cervide are almost entirely wanting. With the exception
of such desert regions, deer have succeeded in establishing themselves in
most districts of the countries into which they have been able to effect an
entrance. Members of the family are found from the ice-bound shores of
Greenland and the Arctic tundras of Siberia to the burning plains of Bengal
and Brazil ; while they are common to the dense grass-jungles and forests of
India and the open pampas of Argentina; as they are to the latter and to
the heights of the Andes and the upland forests of the Himalaya. Never-
theless, as a group, they are mainly inhabitants of forests or scrub-covered
districts, especially where there are open tracts of grass near by on which
they can graze ; and many prefer the thin jungle on the borders of forests
to the forests themselves. The near neighbourhood of water seems to be
essential to the majority, if not indeed to all; and many, like the elk, Pére
David’s deer, and the Chinese water-deer, delight to spend a considerable
portion of their time in the water itself. The Virginian deer, too, may
often be seen swimming from island to island in the American lakes.

While some kinds, such as the red deer, the fallow deer, the chital, and
the sikas, congregate in larger or smaller herds led—at least during the
breeding-season—by one or more of the older stags, others, like the elks,

28 Introductory

muntjacs, roes, and musks, go about, either during the breeding-season or
permanently, in pairs, or even singly. During the season of the renewal of
the antlers the stags of the gregarious species, however, become solitary ;
and in the case of woodland species are extremely careful of their move-
ments. As the antlers mature and the velvet dries up, the animals become
bolder. They at first select the stems of yielding saplings as rubbing-posts
for removing the tattered remnants of the velvet, but in a few days endeavour
to clean off the last remaining shreds by rubbing their antlers against the
rugged bark of any tree that may be convenient.

Although in a wild state deer flee from man and their other enemies
with the greatest precipitation, yet they have a strong spice of curiosity in
their disposition, and may frequently be attracted within range by the sight
of any moving object with the nature of which they are unacquainted, such
as a cap or hat mounted on a pole and slowly waved about above a fence.
This spirit of curiosity is much more strongly developed in the hinds than
in the stags, the former being always the first to approach a strange object,
as they are to come up to be fed in a park. The stags seem, indeed, to
be largely dependent on the hinds as to their movements in all cases
of doubt.

At the pairing-season the adult stags of all the larger species, as well as
many of those of the smaller kinds, exchange their habitually peaceful dis-
position and become extremely pugnacious towards one another. Even
this warlike disposition is, however, chiefly the result of mutual jealousy,
for in a wild condition it appears to be only very rarely that they attack
other creatures. With deer kept in confinement, even when their paddocks
are very extensive, the case is very different; and there are few more
spiteful or more dangerous beasts than an old wapiti stag at such seasons.
When in this condition the males of the larger species are by no means
prepossessing in appearance. ‘They stalk about with the face-glands everted,
the head stretched out, and the blood-shot eyes rolling in a peculiarly
wicked-looking manner; the gait at the same time being curiously stiff
and stilted, while at every obstacle to their progress the head is lowered
and vicious sweeps are made with the antlers. Sometimes, apparently out
of pure spitefulness, the antlers are thrust deep into a bank of sand or clay,
and the broken soil tossed about in all directions. Often, too, they wallow

in the mud and thus make themselves still more unsightly objects.

Habits 29

At the commencement of the pairing-season the old stags of the larger
species commence calling, or roaring ; the cry differing very considerably
even in closely allied species, or even races. In the case of the elk the cry
of the stags is reciprocated by the hinds, although in a different key, and
its object would therefore seem to be to bring the two sexes together. But
in the case of the red deer and wapiti it seems to be a challenge from one
male to another. When two adult stags meet in response to the challenge,
they engage in a combat which lasts till one of the pair is completely
vanquished, upon which the hinds, who have watched the conflict from
a safe distance, put themselves under the escort of the victor. In the
case of the smaller species the calling partakes more of the nature of
a bark. Besides this periodical calling, most or all deer have a perfectly
distinct alarm-cry, more or less resembling either a snort, a sneeze, or
a whistle.

Many of the larger and more specialised deer are comparatively slow
breeders, for although the hinds frequently, if not generally, breed annually,
yet it is but seldom that more than a single fawn is produced at a birth,
twins being rare, while triplets are practically unknown. The elk is, how-
ever, a marked exception in this respect, twins being far from uncommon,
while triplets have often been recorded. The American deer, exclusive
of the wapiti, are more rapid breeders, the Virginian deer often giving
birth to a pair of fawns at a time. Muntjacs likewise frequently give
birth to twins ; but the most prolific member of the whole family is the
Chinese water-deer, which commonly produces four. As is the case with
all ruminants, the fawns are produced in an advanced state of development,
and very soon after birth are able to walk by the side of their parents.
Naturally, however, some considerable time must elapse before they attain
anything approaching the speed of the latter ; and during this period they
are, in most cases, concealed on the approach of danger in thick covert by
their dams, who then lose to a great extent their natural timidity, and
display either great boldness in repelling foes, or cunning in attracting
them away.

Like all other ruminants, deer are exclusively vegetable feeders, but, as
a rule, they browse on leaves to a much greater extent than the Bowde,
although not to the same degree as the giraffe. The elk alone browses on
the needles and twigs of coniferous trees ; but, in addition to this species,

30 Introductory

several crop the leaves of water-lilies and other aquatic plants. Sambar
and the chital do much damage to growing rice-crops, as do some of
the South American brockets to plantations of cocoa and nutmeg.
The reindeer alone subsists during winter more or less exclusively on
lichens and moss, which it obtains by scraping away the covering of snow
with its hoofs.

Economic Uses ——From a commercial and economical point of view deer
are of more importance than is often considered the case. The uses of the
reindeer to the Laps and other northern races are familiar to all; these
animals affording meat, milk, and clothing, and likewise serving for draught
and the carrying of burdens. The elk was also formerly trained to go in
harness, and is then capable of performing long journeys at a high rate of
speed. Antlers of various species are manufactured into the handles of
knives, walking-sticks, and hunting-crops, and were formerly used as a
source of ammonia; while they are still extensively employed as medicine
by the Chinese. Much more important is the use of the skins of deer for
leather, as will be apparent from the following figures taken from Mr. H.
Poland’s Fur-bearing Animals. It is there stated that from 80,000 to
100,000 skins of the American wapiti are annually imported for this
purpose ; of elk a few hundred Russian skins from time to time make their
appearance in the London market, and in 1890 over goo skins of this species
were offered for sale. The pelts of Virginian and mule-deer are imported
in large quantities into London, and again exported to Germany, where
they are used in the manufacture of leggings for the Bavarian peasants, in
one year no less than 227,000 skins being imported. The import of reindeer
skins reaches several thousands annually, some of which are made into
leather gloves. The trade in the antlers of this species is also very large,
Denmark importing about 30,000 from Greenland and 8000 from Russia
annually ; they realise from about £13 to £14 per ton. A trade in reindeer
tongues, which formerly attained considerable importance, seems to have
almost ceased on account of the low prices obtained in England. Of
chital skins no less than 53,922 were sold in London during 1891.
These pelts make most excellent leather, and are now almost exclusively
used for this purpose, their value varying from six to seven shillings each.
A few are, however, worked up into foot-muffs and other articles by the
London furriers. Of the antlers of this and perhaps other Indian species

Economic Uses 31

no less than 424 tons weight were sold in London in 1891, the price
ranging from £1 to £1: 6s. per hundredweight. Fallow deer skins,
known in the trade as buckskin, also afford a most excellent soft and pliable
leather, but it is difficult to obtain accurate figures of the number sold.
It is from this species that most of the venison sold in the London shops
is obtained.

Fic. 5.—Head of Chital, from East Berar, in the possession of Lieut.-Colonel F. H. Whitby.
The antlers are the longest on record, measuring 38 inches along the curve.

PART II.—DESCRIPTIVE

A. THe True DEER—Svus-FamMity CERVINZE

In this section are included all the existing members of the family ex-
cept the musks ; an essential feature being the absence of a gall-bladder to
the liver. The brain, too, is characterised by the great development of the
convolutions on its hemispheres; and there are certain other peculiar
features in the internal organs—especially those of the female—which aid
in distinguishing between the two sub-families. A difference in the
structure of the skull is noticed under the heading of the second sub-family.

With the exception of the existing Chinese water-deer and certain
extinct forms, antlers are always developed in the male, and constitute one
of the most convenient, and apparently also one of the most natural, means
of dividing the members of the sub-family into generic and sub-generic
groups. In his well-known synopsis of the deer, the late Sir Victor Brooke
attributed but little or no classificatory value to these sexual appendages,
but attached prime importance to the lateral (2nd and 5th of the typical
series of five) metacarpal bones, which show two distinct types of ossifica-
tion. In the one type, which is found in almost all the Old World deer,
these bones are represented by their upper (proximal) extremities ; whereas
in the other their lower (distal) ends are alone retained, and serve for the
support of the bones of the lateral or spurious hoofs. ‘To this type belong
all the New World deer with the exception of the wapiti. Impressed with
the importance of this distinction, Sir Victor divided the deer into the two
sections Te/emetacarpi and Plestometacarpi; the former including the genera
Rangifer, Alces, Hydrelaphus, Capreolus, Mazama, and Pudua, and the latter all
the rest. This classification has, however, the disadvantage of sundering
from the American deer such a species as Pére David’s deer, which re-

sembles the former in having antlers of a forked type without a brow-tine.

Reindeer oe

This and other objections to Sir Victor Brooke’s classification have attracted
the attention of Mr. Allan Gordon Cameron, who, in an important paper
communicated to the Fre/d newspaper,' has strongly urged the importance
of the antlers as a primary means of classification, and has called attention
to the essential difference between the “ brow-tined” and the “ forked”
types of antler. And as both these types are of very considerable antiquity,
dating from ages long antecedent to the present epoch, it does appear that
antlers should be accorded much higher value in classification than many
writers are disposed to assign to them. Moreover, in the case of the
majority of extinct species reliance has to be placed almost or quite ex-
clusively on the characters of these appendages ; and it is only by taking
them as the basis that it is possible to bring the living and extinct forms
into line. There are many instances among animals where two distinct
characteristics, if taken as bases of classification, afford dissimilar results; and
as in the present case the antlers appear to be of decidedly more importance
than the lateral metacarpals, the divisions proposed by Sir Victor Brooke
are superseded by those advanced by Mr. Gordon Cameron.

I. REINDEER—GENUus RANGIFER

Rangifer, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 304 (1827),
as a sub-genus ; Brooke, Proc. Zool. Soc. 1878, p. 927; Riitimeyer, d/.
schweiz. pal. Ges. vol. viii. p. 51 (1881).

Tarandus, Ogilby, Proc. Zool. Soc. 1836, p. 134; Gray, Proc. Zool. Soc.
Ronee pe 2245 An. Mag. Nat. Est. ser. 2, vol. ix. p. 416 (1852), Caz.
Ungulata Brit. Mus. p. 189 (1852); Fitzinger, SB. Ak. Wien, vol. Ixvii.
abel p..3401 (1873), Ixix. part 1. p. $34 (1874).

Characters.—Lateral metacarpal bones represented only by their lower
(distal) extremities. Antlers present in both sexes, complex, situated close
to the occipital ridge of the skull, and thus far away from the sockets of
the eyes (orbits), with the brow-tines of adult males palmated, laterally
compressed, deflected towards the middle line of the face, and often un-
symmetrically developed. Above the brow-tine is developed a second
palmated tine, which appears to represent the bez; there is no trez, but
some distance above the bez the beam is suddenly bent forwards to form an
“elbow,” on the posterior side of which is usually a short back-tine ; above

1 See Appendix.
F

34 Reindeer

the back-tine the beam is continued for some distance to terminate in a
large palmation. Antlers of female simpler and generally smaller. Muzzle
entirely hairy ; ears and tail short ; throat maned. Pelage unspotted at all
ages, with a whitish area in the region of the tail, which includes its sides,
but not its upper surface. Main hoofs short and rounded, lateral hoofs very
large. A tarsal, but no metatarsal gland and tuft. In the skull the gland-
pit shallow, and the vacuity of moderate size ; nasal bones well developed,
and much expanded at the upper end. Upper canines wanting ; cheek-

Fic. 6.—Antlers of Male Woodland Reindeer. From a specimen in the British Museum.
(Rowland Ward, Records of Big Game.)

teeth small and low-crowned, with the third lobe of the last in the lower
jaw minute. Bodily size large.

That reindeer are very different from other Cervide@ is apparent not only
from the presence of antlers in both sexes, but likewise from their peculiar
form and backward situation on the skull. With regard to whether the
possession of these appendages is a primitive or an acquired feature, more
evidence than is at present available seems necessary. Especially is this the
case with regard to the ancestry of the group, which is totally unknown,
fossil reindeer remains not antedating the Plistocene, and then being indis-
tinguishable from those of the living European form. Mr. Gordon Cameron
writes as follows on the question :—“ Since there is every reason to believe

SO a

Characters aus

that the possession of defensive weapons, whether cranial or dental, by both
sexes was a primitive character, we have no difficulty in fixing limitation to
sex as the first point of divergence in the history of the antlered deer. We
conclude, further, that Rangifer represents the oldest line of existing deer
with antlers, and the conclusion may be pressed home with the facts noted
by Darwin relative to the time of life at which the antlers first appear. In
seven species, of which the males alone carried antlers, belonging to distinct
sections of the family, and inhabiting different regions, these appendages
were found to develop at periods varying from nine months after birth in
the roebuck, to ten, twelve, or even more months in the stags of the six
other and larger species. In the case of the reindeer, Professor Nilsson found
that the antlers appeared in the young animals within four or five weeks after
birth, and at the same time in both sexes, so that we have here a structure
developed at an unusually early age in one species of the family, and likewise
common to both sexes in this one species only.” A fawn born at Woburn
Abbey also showed antlers in a few weeks.

Against the view that reindeer are a primitive type may be urged the
loss of the spotted coat in the young, which is evidently a specialised feature,
although one which may possibly be of late acquisition. The absence of
remains of reindeer from formations older than the Plistocene period has
been already mentioned. With regard to the argument from the presence
of antlers in both sexes, it is rather difficult to see on what foundation this
rests. In the Pliocene and Miocene cervine genus Dremotherium some
individuals were provided with antlers which were lacking in others, and it
has yet to be demonstrated that these latter are not the females of the former.
Again, in the Bovide, or hollow-horned ruminants, it is mostly in the
specialised and modern forms that the females, if horned at all, carry large
horns. For instance, in the modern oxen the females have these appendages
very largely developed, whereas in some of the Pliocene species of the genus
this sex was hornless. On the other hand, in the kudus (Strepsiceros), har-
nessed antelopes (77age/aphus), and elands (Orias), in which the cheek-teeth
are of a low-crowned primitive type, it is only in the last-named genus that
the females are normally horned. Again, in the gazelles and their allies,
which are known to be an ancient group dating as far back as the Miocene
period, it is generally only the males that are horned.

Although they present an enormous amount of racial and individual

36 Reindeer

variation, yet, as Mr. Caton observes, the antlers of reindeer “ possess features
never to be mistaken by the careful observer, who will at once recognise the
caribou’s antler, no matter what its form.” In some types the beam is long
and rounded and the palmation slight, while in others the former is short
and flattened, and the latter greatly developed. Then, again, whereas in
some antlers the back-tine is placed far above the level of the bez, and the
terminal expansion separated by a long interval from the back-tine, in
others the bez is very much expanded, and situated high up on the short
beam, only just below the line of the back-tine, so as to form a dichotomous
fork with the other half of the beam. Asymmetry is likewise a very
frequent feature. In a Norwegian reindeer in the British Museum with a
simple brow-tine on one side, the corresponding tine of the opposite side is
wanting. In the American woodland reindeer, one brow-tine, generally the
left, is enormously palmated, while the other forms a simple prong.

A magnificent series of reindeer antlers is preserved in the British
Museum, but, unfortunately, in a large proportion of these the locality is not
recorded, in many cases there being no evidence even as to the hemisphere
from which they were obtained. Consequently it is no easy matter in this
country to obtain a good idea of the characters distinguishing the antlers of
the various geographical sub-species. On account of the different curvatures
of the beam, reindeer antlers are extremely difficult to measure accurately ;
the following measurements of some fine examples are taken from Mr.
Rowland Ward’s work, but in several cases the locality is unknown :—

Extreme Length. Circumference. Maximum Span. Locality.
62 53 494 Canada
5 1
60 58 413 ;
593 7 44 ?
57% 54 ? Arctic regions
ral ]
574 54 44% »
542 46 ? North America
54 6% 324 7
54 5 414 Norway
50% 46 29 ?
1 ‘
49 4% 32% e
48 63 254 Newfoundland
47% 54 ? North America

47 4% 29 Norway

Distribution a7

Distribution.—The northern portion of the Holarctic region, but
descending much farther south (as far as France) in the Plistocene epoch
than at the present day. Whether reindeer are ancient or modern inhabitants
of the Arctic regions is a moot point. And whereas some writers consider
that they are comparatively recent immigrants, Dr. Scharff! is of opinion
that their original home is in the north, whence they wandered southwards.
In the absence of decisive evidence in support of one or the other view, it
may be pointed out that if these deer are considered to have originated in
the north, it presupposes the existence of some earlier unknown member of
the family in those regions, and of this we have no present cognisance.

THE REINDEER, OR CArRIBOU--RANGIFER TARANDUS

Cervus tarandus, Linn. Syst. Nat. ed. 12, vol. i. p. 93 (1766) ; Caton,
Antelope and Deer of America, p. 86 (1877) ; Nehring, Tundren und Steppen,
p- 108 (1890).

Cervus guettardi, Desmarest, Mammalogie, vol. 11. p. 447 (1822).

Cervus (Rangifer) tarandus, H. Smith, in Griffith’s Animal Kingdom, vol.
IEP uO 351V.0P. 304 (1627).

Cervus tarandus schottini, Sternberg, Iss, 1828, p. 482.

Tarandus rangifer, Ogilby, Proc. Zool. Soc. 1836, p. 134; Gray, Cat.
Ungulata Brit. Mus. p. 189 (1852), Cat. Ruminants Brit. Mus. p. 66 (1872) ;
Fitzinger, SB. 4k. Wien, vol. lxix. parti. p. 534 (1874).

Rangifer tarandus, Gray, List Mamm. Brit. Mus. p. 181 (1843); Brooke,
Proc. Zool. Soc. 1878, p. 928 ; Ward, Records of Big Game, p. 47 (1896) ;
Scharff, Proc. Irish Acad. ser. 3, vol. iv. p. 473 (1897).

Characters—In spite of the existence of more or less well-marked
geographical races, reindeer from all parts of the northern hemisphere
present such a marked similarity in general appearance that it seems pre-
ferable to regard them as all belonging to a single wide-spread species, of
which most of the characters will be the same as those of the genus. The
pelage is remarkable for its density and compactness ; the general colour of
the head and upper-parts being clove-brown, with more or less white or
whitish gray on the under-parts and inner surfaces of the limbs, while
there is also more or less white above the hoofs and on the muzzle, and

1 Proc. Irish Acad. ser. 3, vol. iv. p. 473 (1897).

38 Reindeer

there may be whitish rings round the eyes ; there is a white area in the
region of the tail, which includes the sides but not the upper surface of the
latter ; and the tarsal tuft is generally white. The antlers are smooth, and
brownish white in colour, but the hoofs are jet black. Albino varieties
occasionally occur in the wild state. A height of 4 feet ro inches at the
shoulder has been recorded in the Newfoundland race.

Distribution —The same as that of the genus.

a. SCANDINAVIAN RacE—RANGIFER TARANDUS TYPICUS
Plate I

The wild Scandinavian reindeer, which may doubtless be regarded as
the typical form of the species, is a smaller animal than the American
woodland race, with antlers approximating to those of the barren-ground
race, but less elongated, and with a distinct back-tine in the male, the
brow-tines moderately palmated and frequently nearly symmetrical, and the
bez not excessively expanded. Female antlers generally much smaller than
those of the male, although occasionally as large, but with much fewer
points. Pasterns of feet very short and broad, without any distinct cleft
between the two component phalangeal bones of each segment, and a tuft
of hair immediately above the cleft of the hoof. Tail very short. ‘Tarsal
tuft very long. The white ring above the hoofs passing imperceptibly
into the fawn of the limbs without any distinctly marked line of division
in the male, and, in some cases at least, almost absent in the female. A
distinct light ring round the eye in the male, but not in the female.
Upper surface of tail but little darker than the sides and under surface.
Some white on the muzzle.

The above description is taken from a mounted male and female from
Norway in the British Museum, supplemented by Dr. J. A. Allen’s notes.
Seen side by side with the male woodland reindeer mounted in the same
case, they confirm the latter writer’s statement that the Scandinavian rein-
deer has the appearance of being quite a different animal from either of the
New World races. Most marked is the difference in the conformation of
the feet, although other specimens are required to show how far this is
constant. In their shortness and expansion, the feet of the European rein-
deer seem more adapted for marshy districts than are those of the American

THe DEER or Att LAnps. Pruate |.

SCANDINAVIAN REINDEER.

Published. by Rowland, Ward) rd .

Scandinavian Race 39

forms. In the female specimen referred to above, the antlers are as large as
those of the male, but simpler and without the back-tine, but in other
Norwegian examples they are much smaller.

Distribution—The Scandinavian reindeer inhabits a large part of
Norway, Sweden, and Lapland, and extends into Russia, although its
eastern limits in Asia cannot yet be ascertained. In the Ural district the
southern limits are about the 52nd parallel of latitude in the Kirghiz
Steppes, but in Kazan reindeer are not known south of latitude 54°. The
domestic race has been introduced into Iceland. In the extreme north,
reindeer, although unknown in Franz Josef Land, are found at Cape
Chelyuskin, and also in Novaia Zemlia, Spitzbergen, and Phipps and
Parry Islands, lying between the 8oth and 81st parallels. I have, however,
no means of knowing to what race these belong. Even in the historic
period the southern range of the reindeer seems to have been more ex-
tensive than at present, as these animals are stated to have existed in the
Black Forest during Czsar’s time, although their alleged occurrence in
Caithness is probably incorrect. Be this as it may, there is abundant
evidence of the existence of reindeer in Britain during the Plistocene epoch,
and they even extended as far south as the valleys of the Dordogne
and Garonne in France. In the memoir already cited, Dr. Scharff states
that two races of reindeer, the one representing the American barren-
ground and the other the woodland type, are met with in Europe, while
only the latter occurs in Asia. And he proceeds to draw the inference
that while the woodland type reached Asia by way of Bering Strait,
the barren-ground race has migrated by way of Greenland, and that this
race alone reached Ireland. With regard to the occurrence of the wood-
land form in Eastern Asia, I agree with the author, but I cannot see that
the Scandinavian reindeer can be identified with the barren-ground race,
neither can I find evidence of the existence of the woodland type in
Europe at the present day, whatever may have been the case in past epochs.
Whether the wild Kazan reindeer are the same as the Scandinavian, may
perhaps be doubtful. Eversmann! states that the females of the former are
generally without antlers, and that the males are considerably larger than
those from Norway and Lapland.

Habits.—In treating of the habits of the Scandinavian reindeer, those

1 Bull. Soc. Moscou, 1840, p. 58.

40 Reindeer

of the other Old World races may be included, as they appear to be very
similar. In Norway, wild reindeer inhabit the high fjelds, where almost
the only other inhabitants are the ptarmigan and the black and yellow
lemming. These high plateaus contain numerous hollows in which snow
remains throughout the summer, and on these snow-beds the reindeer are
fond of lying, so that in stalking a telescope is frequently of little use in
discovering the whereabouts of a herd. Although essentially gregarious
animals, it appears from the account given by Mr. E. N. Buxton! that
during the summer the herds on the high fjelds of Norway comprise nearly
g5 per cent of females, and it is suggested that this may be due
to the males having thicker hides, and thus being less tormented by
mosquitoes, so that they are able to remain in the valleys below, where
the food is richer. The same writer was also told that the hinds carry
their antlers for some months after those of the bucks are shed, and during
this time are able to keep their lords and masters in complete subjection.
At the present day the numbers of the wild Norwegian reindeer are
greatly reduced. But it appears from Mr. J. Lloyd’s Scandinavian
Adventures that in the early part of the century these animals were as
abundant as blesbok in Africa. In that work it is stated that one day in
June 1826 the fjeld, for a breadth of seven miles, was covered with reindeer
as thickly as an English field by sheep when feeding ; the herd extending
so far that the eye could not embrace the whole at once. Subsequently
this immense herd broke up into three divisions.

In Spitzbergen, according to the narrative of Baron Nordenskidld, the
wild reindeer resort in summer to the grassy plains in the valleys free from
ice; late in the autumn they are reported to descend to the coast, where
they pick up a subsistence on the sea-weed thrown by the waves on the
beach, but in winter they once more return to the inland heights, where, in
spite of the intense cold, they thrive well on the reindeer-moss, or lichen,
growing beneath the snow. Indeed, when they return to the coast in
spring they are in excellent condition, but shortly afterwards, when the
melting snow cakes into ice on the mountain-sides, they are much put to
it to find food, and consequently become very thin. With the return of
summer they speedily recover, and by the end of autumn are quite fat. Even
in such a remote district as Spitzbergen the numbers of these animals have

1 Short Stalks, p. 330 (1892).

Spitzbergen Race AI

been greatly diminished, and in the north-western districts they have been
completely extirpated, although they are numerous in Ice-Fjord. Whether
the stock on this comparatively small island is from time to time replenished
by immigration across the ice from some unknown land to the north-east,
is not definitely known, although it is suggested that such may be the case.

Apparently in all the continental parts of their habitat reindeer under-
take periodical migrations, and in Eastern Siberia herds numbering thousands
of individuals have been observed descending from the northern hills and
crossing the open plains to reach the forests where they pass the winter.
Such herds split up into minor parties, which move slowly and regularly,
and are led by an old deer of large size. One reason that these animals
migrate to high grounds in summer is to escape the attacks of mosquitoes
and gad-flies, by which they are much tormented in the valleys,
their skins being sometimes full of bots. The breeding habits appear
similar to those of the American races. Reindeer have been tried at Woburn
Abbey, but with no success, as they always die after a comparatively short
sojourn. This and other instances may be taken as a convincing proof that
the climate of England has undergone very considerable alteration since the
time when the country was inhabited by these animals.

5. SPITZBERGEN RaAcE—RANGIFER TARANDUS SPETZBERGENSIS

Cervus tarandus spetzbergensis, Andersen, Ofvers. Vet. Ak. Forhandl.
1862, p. 457; Nitsche, fahresb. Ver. Nat. Wiirtt. 1893, p. 111.

Characters.—This form is described as differing very markedly from
both the Scandinavian and Greenland races, but there are no mounted
examples in English collections. The most marked characteristic appears
to be afforded by the nasal bones of the skull, as figured by Dr. Nitsche.
In the Scandinavian reindeer these bones increase regularly in width from the
lower end to the maximum diameter on the line of the lachrymal vacuities,
but in the Spitzbergen form they are much expanded at both extremities
and deeply constricted in the middle, and there is also a decided difference
in the form of their superior border. The same type of nasal bones is
exhibited in a male skull in the British Museum purchased in Spitzbergen
by Dr. J. W. Gregory ; andI am informed by Mr. Trevor-Battye that the
same feature is constant in all the specimens that have come under his

G

42 Reindeer

notice. In the British Museum skull the antlers approximate to the

Scandinavian type, although they are smaller and the beam is relatively

shorter ; both the brow- and bez-tines are expanded, the right brow being

much larger than the left. In the two skulls figured by Dr. Nitsche the

antlers are simple, and are probably either immature or those of females.

The whole animal is inferior in size to the Scandinavian reindeer.
Distribution —Spitzbergen.

c. WoopLanp RacE—RANGIFER TARANDUS CARIBOU

Cervus tarandus caribou, Gmelin, Syst. Nat. vol. i. p. 177 (1788).

Cervus hastalis, Agassiz, Silliman’s “fournal, 1847, p. 436.

Cervus tarandus sylvestris, Richardson, Fauna Bor. Amer. p. 251 (1829).

Rangifer caribou, Audubon and Bachman, Quadr. N. Amer. vol. iii. p.
111 (1853) ; Baird, N. Amer. Mamm. p. 633 (1857); J. A. Allen B77
Amer, Mus. vol. viii. p. 234 (1896); Miller, P. Boston Soc. vol. xxviii. p.
40 (1897).

Tarandus hastalis, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 349
(1873), xix. part 1. p. 542 (1874).

Characters.—This form is the one commonly known in America as the
woodland caribou, and is of large size, with the antlers stout, flattened,
much palmated, and not of excessive length ; one of the brow-tines being
much expanded, while the other is simple ; the bez-tine being also more
palmated than in the Scandinavian form, and the back-tine well developed.
Female antlers are proportionately smaller than in the Scandinavian race.
From the evidence of a mounted specimen in the British Museum obtained
from Hudson Bay, and Dr. Allen’s notes (which are based on three
examples from New Brunswick in the autumn pelage), the following
features seem distinctive of this form, especially as compared with the New-
foundland race. The pasterns of both fore and hind feet are very long and
slender, with the cleft of the hoofs continued upwards on the front line
as far as the lower end of the cannon-bone, and there is no distinct tuft of
hair above the cleft of the hoof. The colour of the body and limbs is
much darker than in the Newfoundland race, the dark area extending over
the anterior half of the lower surface of the body ; and, except the extremity
of the upper lip, the muzzle is as dark as the face, while there is no light

Woodland Race 43

ring round the eye. On the limbs the white is restricted to a sharply-
defined band of about half an inch in width just above the hoofs, but
ascending behind to enclose the lateral hoofs. In the British Museum
example the tarsal tufts are much less conspicuous than in the Scandinavian
reindeer; and the muzzle has rather more white than Dr. Allen describes.

Regarding the variations in colour displayed by the woodland race, Mr.
Caton writes that while the head and legs “are always distinctly coloured,
in a large majority of individuals white
predominates, especially on the neck,
which is almost universally the whitest
portion of the animal. The long white
mane of the old buck is a very striking
Gmaracteristic, . . . Dhere is less uni-
formity in the colour of the body than
of the head, neck, and legs. While the

head and legs are tawny brown of varying

1
iy
S 4
-_ o
f

intensity, and the neck white, in some
much more pronounced than in others,
the body is sometimes nearly all white ;
but others are a rich rufous brown on
the back as well as the legs, and only
the tail and rump are white above, the
belly and inside of the legs being also
white. Like all the others, the early
winter coat, which replaces the summer

Evabeia September, is or the deepest 4, 7 side view of Skull and-Antlers, of
colour, is finer, softer, and more brilliant Male Woodland Reindeer. From a Nova
than later, when the clove-brown shade nace
which first prevailed has given place to the dirty white of mid-winter.”

A fine pair of antlers from Siberia in the British Museum show all the
characteristics of the American form, and lead to the conclusion that the
range of the latter extends into Eastern Asia. If the Siberian form prove
distinct, it will apparently require a new sub-specific name, as it cannot be
identified with the Scandinavian race. The antlers from Nova Scotia
represented in Fig. 7 exhibit the extreme development of the woodland race,
approximating in their massiveness to the Newfoundland race, but have

44 Reindeer

fewer points on the upper surface of the extremity of the beam than usual ;
those shown in Fig. 6 are more elongated, and have more points.
Distribution —Upper North America as far north as the limits of forests,
but the boundary varying somewhat owing to seasonal migrations ; and
apparently North-Eastern Asia as far as Siberia. In America the range
includes Labrador and Northern Canada, and thence south to Nova Scotia,
New Brunswick, Northern Maine, and Lower Canada on both sides of the
river St. Lawrence, whence it passes westwards through the districts north
of Quebec to the neighbourhood of Lake Superior, to the south of which
reindeer are unknown. It is curious that Mr. Caton identified this form
with the typical Scandinavian reindeer, although the latter appears to be
more nearly related to the American barren-ground race.
Habits.—Although its habits are doubtless essentially the same as those
of the Scandinavian reindeer, yet the woodland race has certain distinctive
ways of its own, which serve to differentiate it from the barren-ground
reindeer. In the first place, it is essentially a forest-dwelling animal, and
to this may be due the comparative shortness and thickness of its antlers.
In the second place, while the woodland race generally associates in small
parties of not more than a dozen head, the barren-ground form collects in
herds numbering thousands of individuals. The Newfoundland race is,
however, somewhat intermediate between the other two in this respect, as
it collects in considerably larger droves than its woodland relative of the
continent. More important are the differences in the migratory habits of
the two, the woodland race travelling northwards and the barren-ground
southwards in the autumn. On this subject Sir John Richardson wrote as
follows :—‘‘ Contrary to the practice of the barren-ground caribou, the
woodland variety travels to the southward in the spring. They cross the
Nelson and Severn rivers in numerous herds in the month of May, and
pass the summer on the low marshy shores of James Bay, and return to the
northward, and at the same time retire more inland in the month of
September.” Commenting on this passage, Mr. Caton has the following
observations :—‘ Here, then, we find the woodland caribou migrating to
the northward, on the west coast of Hudson Bay, and west of it as high as
55 to 57° of north latitude, or within one degree of Churchill, which is
near the southern limit of the range of the barren-ground caribou in that
longitude as given by Richardson, though I have authentic evidence that

Newfoundland Race 45

they sometimes come considerably farther south in exceptional seasons.”
After stating that they have been known to wander as far south as Lake
Huron, in about lat. 47°, he concludes that the two forms must not unfre-
quently meet in the breeding-season, and yet there is no evidence that they
ever cross. Assuming its correctness, this statement is very remarkable,
bearing in mind that such totally distinct forms as the red deer, wapiti, and
Japanese sika will readily interbreed.

Writing of the woodland reindeer, Mr. Caton observes that it ‘“‘is
fond of arboreous food, grasses, and aquatic plants, but its great resource is
lichens. It frequents marshy and swampy grounds more than any other of
the deer family, for which it is admirably adapted, and where it is well
protected from pursuit. In the winter it resorts to the dense forests on
high ground. The breeding-season is in September, and the fawns, which
may be one or two in number, are born in the following May. The males
shed their antlers in December, although those of the does remain till the
spring.” Mr. Caton describes the woodland reindeer as an exceedingly
wild and restless animal, of a highly wary and suspicious nature, and one
much given to constant change of feeding-grounds. An alarm from which
a wapiti will be caused to flee but a few miles, will drive a caribou clean
out of the country. Hence the animal does not promise well for domestica-
tion; and, as a matter of fact, no thorough attempt seems to have been
made to domesticate either of the American races, although tame reindeer
from Lapland have been imported into Alaska. In summer these reindeer
graze on the rich grass of the valleys, keeping quite clear of the thickets.

d. NEWFOUNDLAND RAcE—RANGIFER TARANDUS TERRA-NOVZE

Rangifer terra-nove, Bangs, Description of Newfoundland Caribou, Boston,
1896; J. A. Allen, Bu//. Amer. Mus. vol. viii. p. 233 (1896).

Rangifer tarandus terre-nove, J. A. Allen, MS., see op. cit. p. 235.

Characters.—Dr. J. A. Allen, by whom this sub-species has been
described, gives the following characteristics as distinctive. The bodily
size is large, and the antlers are very massive and much _ palmated, with
numerous points on all the branches and especially on the hinder border of
the beam. The adult male in the autumn pelage has the following colora-
tion: upper-parts grayish brown, becoming lighter on the flanks, and

46 Reindeer

passing into nearly pure white on the under surface of the body ; neck
dirty white, somewhat purer in front ; a broad, ill-defined light ring round
the eye, and the entire muzzle and lower portion of the face, as. well as the
extremity of the lower jaw, grayish white ; rest of head like the back ;

edges and lower surface of tail and buttocks white ; front and outer surface

Fic. 8.—Head of Male Newfoundland Reindeer. From Dr. J. A. Allen,
Bull. Amer. Mus. vol. viii. (1896).

of limbs brownish gray ; feet and terminal third of the metacarpal and
metatarsal segments white, passing gradually into the general colour of the
limbs above. The female shows rather less white ; and the young is still
darker, with a dusky line on the flanks, and a blackish streak running down
the back and expanding over the shoulders.

By previous writers the Newfoundland reindeer has been considered

identical with the mainland race. Indeed, after stating that it is found in

Barren-Ground Race 47

numbers on the island, Mr. Caton mentions that it is reported frequently
fOucioOsssonmtne ice fo the continent. If this be true, the right to sub-
specific distinction would be more than doubtful. The chief claims to dis-
tinction appear to be the generally lighter colour, the much greater extent
of white on the feet and muzzle, the light rings round the eyes, and the
shorter and more massive antlers. From a series of specimens that I have
lately seen, the characters of the present form appear to be very constant.
Distribution —-Newfoundland.

e. GREENLAND RAcE—RANGIFER TARANDUS GRCENLANDICUS

Cervus tarandus grenlandicus, Gmelin, Syst. Nat. vol. i. p. 177 (1788).

Rangifer grenlandicus, Baird, N. Amer. Mamm. p. 634 (1857) ; Caton,
Antelope and Deer of America, p. 105 (1877); J. A. Allen, Bull. Amer. Mus.
vol. vill. p. 234 (1896).

Characters—The Greenland reindeer appears nearly allied to the
barren-ground race, having long, slender, rounded antlers, with few points,
and showing great variability in form. There is a broad, sharply defined
white ring round the eye; and a distinct, deep white band immediately
above the hoofs. Not having seen specimens, I am unable to give further
particulars about its coloration. In the antler figured by Dr. J. A. Allen
there is a back-tine, but the bez has only two points.

Distribution —Greenland.

J. BARREN-GRoundD RAcE—RANGIFER TARANDUS ARCTICUS

Cervus tarandus arcticus, Richardson, Fauna Bor. Amer. p. 239 (1829).
Tarandus arcticus, Baird, U.S. Patt. Off. Rep. Agric. 1851, p. 105.
Rangifer arcticus, J. A. Allen, Bull. Amer. Mus. vol. viii. p. 234 (1896).
Characters.—Size small. Antlers of male very long, slender, and
rounded, with but few points on the expanded portion of the beam, which
is separated by a long interval from the bez-tine; the latter tine, and
generally one of the brows, more or less expanded; back-tine generally, if
not always, wanting ; female antlers much smaller, simpler, and scarcely
curved at all.1_ As I have never seen a skin of this form, I can say nothing

1 See figure on p. 104 of Mr. Caton’s work.

48

Reindeer

as to the structure of the feet, and I am compelled to rely on Dr. Allen’s

notes as regards coloration.

He states that instead of the narrow white

ring above the hoofs found in the woodland reindeer, the whole foot and

leg are white, with the exception of a tawny brown stripe running down

Fic. 9.—Side view of Antlers of Male
3arren-Ground Reindeer. From a
specimen in the British Museum.

the front of each, which narrows till it reaches
the lateral hoofs. There is no distinct white
ring round the eye; the amount of white on
the muzzle is not stated.

Contrasting the present with the wood-
land race, Mr Caton observes that the former
has a foot similarly provided with coarse stiff
hairs, but the white is much more extended.
Instead of being confined to a narrow band
surrounding the hoofs, while the leg is of a
nut-brown shade, the whole foot and leg are
white, except a tawny brown stripe extending
down the front of each leg, with white
hairs interspersed, diminishing in extent as it
proceeds downwards till it terminates in front
of the accessory hoofs.

The antlers herewith figured are fair
average examples of those of this race; a
much larger and still more characteristic pair
are now exhibited on the top of the front of
the reindeer case in the British Museum, but
as one of the bez and both of the brow-tines
are wanting, the specimen is not well adapted
for figuring. These antlers are much longer
than any examples of the woodland reindeer,
and indeed this is very generally the case

with this form, although its bodily size is so much inferior to that of the

latter.

Distribution.—The barren districts lying to the northward of the forest

zone in Arctic America, and extending to the confines of the Polar Sea.

Habits —The gregarious nature and long migrations of this reindeer

have been already alluded to under the head of the woodland race.

Elk 49

Although, unlike the latter, the present race is normally an inhabitant of
open country, during its southern winter migration it enters the forest
districts normally tenanted by the woodland caribou. According to Sir
J. Richardson, the majority of the males and females herd together
separately for the greater part of the year ; in the winter the former retire
for some distance into the woods, while the latter linger on the skirts of the
barren-grounds and proceed to the neighbourhood of the coast quite early
in the spring. Both the woodland and barren-ground races enjoy a
plentiful supply of winter provender, since alike in Labrador and the
districts farther north the reindeer-moss forms a carpet of a yard in depth,
rendering it possible to traverse with ease boulder-clad districts which
without this covering would be almost impassable. The pairing-season is
stated to be later than that of the woodland race.

II. Etx—Genvus ALces

Alce, H. Smith, in Grifith’s Animal Kingdom, vol. v. p. 303 (1827), as a
sub-genus, zec Blumenbach, 1803.

Alces, Jardine, Naturalists Library—Mamm. vol. iii. p. 125 (1835) ;
Ogilby, Proc. Zool. Soc. 1836, p. 135; Gray, List Mamm. Brit. Mus. p. 182
(ics ai Pitzineer, Sb. 72: Wren, vol. Ixvi. patt 1. p. 348 (1873), lxix.
Paton ps 521 (0674): Brooke, Proc. Zool. Soc. 1878, p. 915; Rutimeyer,
Abh. schweiz. pal. Ges. vol. vill. p. 54. (1881).

Alcelaphus, Gloger, Handbuch Naturgeschichte, p. 143 (1841), nec de
Blainville, 1816.

Cervalces, Scott, Proc. Ac. Philadelphia, 1885, p. 181.

Characters.—Lateral metacarpals as in Rangifer. Antlers present only
in the males (as in all the following genera), situated low down on the
skull, from which they arise at right angles to the median longitudinal
line, extending at first directly outwards in the plane of the forehead, and
finally expanding into a broad palmation margined with snags. As shown
by immature—especially European—specimens, the antlers are essentially
dichotomous, both main branches being palmated, but the upper much
superior in size to the lower. Muzzle broad, long, and overhanging, with
a very small triangular naked area between the lower angles of the nostrils ;
head and limbs very long, neck and body short ; tail very short, main hoofs

H

50 Elk

long and pointed, lateral hoofs large. A small metatarsal gland and
tarsal tuft, the former situated high up. Pelage, uniformly coloured at
all ages and all seasons,' long and coarse ; males provided with a peculiar
pear-shaped pendulous expansion of skin covered with long hairs on the
throat. In the skull, the nasal bones very short, and the nasal aperture
consequently of great extent; gland-pit and vacuity between the bones of
the face moderate. Upper molar teeth broad, low-crowned, and approxi-
mating to those of the giraffe. Size very large, and build clumsy.

With the exception of a certain similarity in the conformation of their

Fic. 10.—Front view of Frontlet and Antlers of Elk. From an immature specimen in the
British Museum.

antlers to those of the American deer (with which they also agree in the
structure of the lateral metacarpal bones), elk seem fully as isolated a type
as reindeer. Although there is evidence of the existence of the genus
since the Pliocene epoch, there is nothing definitely known as to its
relationship with other members of the family. Essentially, as seen in
immature specimens (Fig. 10), the antlers are of the forked, or dichotomous,
type; the anterior prong of the fork simply dividing again, while the
posterior prong splits into a fork of which the front prong is simple but the
hinder branch expands to form the main palmation. In very old animals

' Young American elk are stated to show a faint dappling.

Characters ot

almost all the tines may unite to form a continuous palmation, although a
more or less distinct cleft remains to mark the position of the first fork.
The entire antler in immature specimens is not unlike that of the marsh-
deer in general plan, with the exception of the different direction of the
beam ; and it is remarkable that abnormal antlers of several of the American
deer frequently exhibit a tendency to palmation. Whether this structural
similarity between the antlers of d/ces and Mazama does indicate a more or
less intimate relationship, I am not prepared to say. Mr. Gordon Cameron,
from the different direction of the beam, considers that it does not ; and the
structure of the molars is decidedly different in the two, but we have yet to
learn the morphological value of these characters. That the shortness of
the nasal bones and the large size of the narial aperture of the skull are
specialised features of comparatively modern acquisition, is evident from the
fact that the skull of the young elk is much less different in this respect from
that of an ordinary deer than is the case in the adult. This inference is also
confirmed by the circumstance that the extinct North American form
described as Cerva/ces has simpler antlers and longer nasal bones, thus
indicating a more generalised type. And, in my opinion, this type does
not seem worthy of more than specific distinction from the existing
form.

Since the generic name 4/ce, from which the later 4/ces can scarcely be
regarded as distinct, was first applied to the gigantic extinct fallow deer,
it evidently properly belongs to the damine group of Cervus, and should
thus supersede Dama. Ifsuch a change be thought desirable, Cervalces will
apparently have to stand for the present genus, the existing species then
taking the name of C. a/ces.

Some degree of uncertainty has existed as to the presence of a metatarsal
gland, but the question has been answered in the affirmative by Dr. H.
Nitsche,' who points out that this gland is similar to the corresponding one
in Mazama, being naked with a marginal fringe of hair, instead of fully
haired, as in Cervus. It must, however, apparently be very minute, as
it is not visible in the mounted specimens in the British Museum, which
exhibit the tarsal tuft very distinctly.

Distribution —The more northern districts of the Holarctic region.

1 Zool. Anzeiger, vol. xiv. p. 181 (1891).

Elk

oa
yw)

1. THE ELK, or MoosrE—ALCES MACHLIS

Cervus alces, Linn. Syst. Nat. ed. 12, vol. i. p. g2 (1766); Caton,
Antelope and Deer of America, p. 69 (1877); Nehring, Tundren und Steppen,
p. 107 (1890); Nitsche, Zoo/. Anzeiger, vol. xiv. p. 181 (1891).

Cervus alce, Boddaert, Elenchus Animalium, p. 135 (1785).

Cervus coronatus, Lesson, Man. Mammalogie, p. 356 (1827).

Cervus (Alce) alces, H. Smith, in Griffith’s Animal Kingdom, vol. iv.
p. 72,.¥s ‘pe sea; (me27)

Cervus (Alce) coronatus, H. Smith, /.c. pp. 95 and 304.

Alces americanus, Jardine, Naturalist’s Library—Mamm. vol. ili. p. 125
(1835) ; Merrick, Mammals of Minnesota, p. 270 (1892).

Alcelaphus alce, Gloger, Handbuch Naturgeschichte, p. 143 (1841).

Alces machlis, Ogilby, Proc. Zool. Soc. 1836; Brooke, Proc. Zool. Soc.
1878, p. 916; Lydekker, Cat. Foss. Mamm. Brit. Mus. part iu. p. 78
(1885); Grevé, Zool. Garten, vol. xxxv. p. 267 (1895); Ward, Records of
Big Game, p. 50 (1896).

Alces palmatus, Gray, List Mamm. Brit. Mus. p. 182 (1843) ; Blasius,
Sdugeth. Deutschl. p. 434 (1857).

Alces (Cervus) alces, Sundevall, K. Svensk. Vet. Ak. Hand. for 1844, p. 176
(1846).

Cervus lobatus, Agassiz, Proc. Boston Soc. vol. 11. p. 188 (1846).

Alces malchis, Gray, Knowsley Menagerie, p. 56 (1850), Cat. Ungulata
British Museum, p. 187 (1852), Cat. Ruminants British Museum, p. 66 (1872).

Alces muswa, Richardson, Zool. of Herald—Mamm. p. tor (1852).

Alces jubata, Fitzinger, Naturges. Saugethiere, vol. iv. p. 86 (1864),
SB. Ak. Wren, vol. \xix. part i. p. 521 (1874).

Alce alces, Gilpin, Mamm. Nova Scotia, p. 119 (1871).

Alces lobata, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 348
(1873), Ixix. part i. p. 528 (1874).

Alces lobata coronata, Fitzinger, op. cit. Ixix. p. 528 (1874).

Alce americanus, Merriam, Mammat/s of the Adirondacks, p. 138 (1884) ;
Miller, Proc. Boston Soc. vol. xxviii. p. 40 (1897).

Plate Ll

Characters.—The type species. Largest of living deer, the height at

Tae Derr or Ati Lanps PLATE Il.

MALE ELK.

Published. by Rowland. Ward. hid

Characters 3

the withers varying from 5 feet g inches to 6 feet 6 inches.'’ Antlers with
a very short beam and the palmation so developed as frequently to obliterate
almost all traces of the primitive form, with the exception of a remnant of
the cleft of the first fork. Nasal bones of skull extremely short, and the
premaxille not extending sufficiently far upwards on the front edges of the
maxille to articulate with the nasals. Hair long, coarse, and rather brittle,
longest about the neck. General colour varying from yellowish gray to
deep blackish brown, generally darker in American than in European

Fic. 11.—Upper view ot Skull and Antlers of Elk. From an American specimen in the possession of
the Duke of Westminster. This appears to be the largest on record, the maximum span being
6 feet 14 inches.

examples ; lower portion of limbs whitish ; forehead dark chestnut ; face
below the eyes nearly black, but reddish gray near the muzzle.

The winter coat, which in America first makes its appearance in
September, is darker than the summer pelage ; and it is when first assumed
that it is darkest, its colour gradually fading till the spring change. More-
over, it is only in animals of the second or third year that the winter coat
attains its deepest sable, growing gradually lighter each succeeding year,

1 The late Prof. Garrod, Casse//’s Natural History, vol. iii. p. 51, gives the height at 8 feet ; the tallest
specimen definitely recorded is a Canadian one shot by General R. S. Dashwood, which measured 64 ft.

54 Elk

till in old males it becomes more or less grizzly. In regard to this senile
fading, Mr. Caton writes as follows: ‘That this is much more the case
with some than with others, we may not question, any more than that
individuals of all ages differ very appreciably in colour, which is admitted
by all. It is by far the darkest coloured of all our deer, and it is probably
the darkest of any known deer of any part of the world. It has always
been recognised as much darker than the Swedish elk, with which, I am
entirely satisfied by critical examination, it is specifically identical.” Not
only does this last conclusion appear well founded, but it seems impossible
to regard the Old World and New World elks as even representing dis-
tinct sub-species, although many zoologists refuse to believe that one form
can be common to two separate continents. In weight an elk will scale
from goo to 1400 pounds, and the antlers may weigh as much as 60 pounds.

At the present day, European antlers never attain dimensions equal to
those of the largest American examples, but it is possible that this may be
due to the greatly diminished numbers of the former. As in the case of
the reindeer, the locality of many of the elk antlers in the British
Museum is unknown. The following are the seven largest specimens
recorded by Mr. Rowland Ward, all being American :—

Length to Longest Point and Circumference Tip to Tip Greatest Breadth of
on Inside Curve. above Burr. Span. Width. Palmation.
1
44 ? ? 614 ?
1 1 1 al!
434 83 40 594, 142
3 3 JL
423 10 354 56 134
? : ? 67 ?
41 gh ; ; 213
if
41 P P 545 P
41 r P 65 24

With regard to nomenclature, it is quite certain that, according to the
strict rules of priority, 4/ces machlis is not the proper title for the elk.
But the difficulty of arriving at a satisfactory conclusion as to what this
should be is so great, that it seems preferable to allow the name commonly
in use to remain. It is true this name is antedated by Aéces americanus ;
but many naturalists would replace this by <A/ces alces, or Alce alces—
combinations which do not meet with approval on my own part, and prob-
ably on that of many of my readers. Moreover, as mentioned above,

Distribution 55

Alce properly belongs to the giant extinct fallow deer, and I have great
hesitation in admitting that /ces is sufficiently distinct from A/e to be
allowed to stand as a separate generic name. I have already suggested that
Cervalces should be adopted for the genus, and a/ces for the species, but as
it is probable many zoologists would refuse to admit that the type of the
former is generically identical with the living elk, this sweeping change
has not been made.

Distribution —The forest and marshy districts of Norway and Sweden,
Eastern Prussia, Livonia, Northern Russia, and thence eastwards through
Siberia north of about latitude 50° to Amurland. In America the elk is
found in Alaska, Montana, Nova Scotia, and New Brunswick. Baird gives

Fic. 12.—Antlers of Elk viewed obliquely. (Rowland Ward, Records of Big Game.)

the distribution in his time as ‘‘ west coasts of America from the shores of
the Arctic Ocean nearly to the Columbia River. Farther east the northern
limit is about 65°, and thence through Canada to Maine, New Hampshire,
Vermont, and the northern parts of New York.” From the Adirondack
region of New York it was exterminated about 1861. It has been
commonly stated that the elk occurs in the Caucasus, but this, according
to Dr. K. Satunin,' is an error, although it may not improbably range to
the forests on the northern flank. In Europe the range of the elk has
been steadily contracting for centuries, while everywhere its numbers are
rapidly diminishing. In Saxony the last example was slain in 1746, and
in Silesia in 1776. In Julius Czsar’s time it was abundant in the Black
Forest, and even in the third century seems to have been spread over all

1 Zool. Fahrbuch Syst. vol. ix. p. 309 (1896).

56 Elk

the forest-clad portions of Germany, while its range also included Britain.
Although so much diminished in numbers in Europe, elk are said to be
still abundant in the valley of the Lena, in the neighbourhood of Lake
Baikal, in Amurland, Mongolia, and Tungusia. In Norway they are
chiefly found in the eastern provinces of the south, while in Sweden their
home is the primeval forests clothing the chief mountain range. Antlers
and bones referable to this species have been obtained from the superficial
deposits of Cleveland, Walthamstow, and other parts of England, some of
which are preserved in the British Museum. The following paragraph
taken from the Times of 2nd January 18g1 affords interesting informa-
tion as to the number of elk and reindeer in Scandinavia :—‘‘ The total
number of elk killed in Norway last year was 850, of which 515 were
male and 335 female, and of the eleven provinces in which they were
killed, Northern Trondhjem contributed 303. The number of reindeer
killed during the same period was 468, and 143 of them were killed in the
province of Romsdal. The number of elk killed in Sweden during the
same period was 1782, as against 2097 and 2178 in the two preceding
years, and it has been remarked for some time that the number of elk in
Sweden has been diminishing.”

Habits.—The long limbs and deeply cleft hoofs of the elk show that it
is adapted to dwelling in marshy ground ; while the short neck and the
long, fleshy, prehensile muzzle indicate as clearly that it is a browsing
animal and unable to graze like other deer. In both hemispheres elk are,
indeed, essentially forest-dwelling animals, their favourite haunts in the
Scandinavian peninsula being the boundless forests of spruce and birch
clothing the central plateau. According to Mr. E. N. Buxton, in Norway
they are, however, much more abundant in the forests of deciduous trees
than in those composed of conifers, their food in summer consisting to a
great extent of the leaves of the mountain-ash.

At this season their feeding time is in the early morning, and again in the
evening ; and by about ten o’clock in the morning they betake themselves
to their resting-places, from which, if unmolested, they do not stir till
about three in the afternoon. Elk are remarkable for their acuteness of
hearing, and are consequently exceedingly difficult to approach. In
districts where they are much hunted one of their favourite devices to avoid

successful pursuit is to return for a longer or shorter distance now and

Habits $7

again on the leeward side of their path, so that they are sure to receive
the scent of any person or animal following their trail. Whether the
Scandinavian elk is as fond of wading after aquatic plants as its western
brother, I have been unable to ascertain.

As a rule, elk in both hemispheres do not associate in herds, although a
few may collect together during winter. For a considerable part of the
year the stags and hinds keep more or less apart, but at the beginning of
the hunting season in Norway a male and female, together with one or two

SW hye

Fic. 13.—Elk in the Park at Woburn Abbey. From a photograph by the Duchess of Bedford.

calves, may frequently be found in company, although at the same time
other individuals of both sexes roam about alone. The pairing-season in
Norway commences about the latter part of September, and at this time
the stags become extremely pugnacious. During this season both sexes
“call,” the cry of the female having been compared to the roar of an
angry bear. Many hunters, however, have stated that they have constantly
hunted elk in Norway without ever hearing them call; and Sir Henry
Pottinger in the Badminton Library suggests that elk may possibly be
less noisy in Scandinavia than they are in America. During the season
I

58 Elk

in question the male elk eats but little, and consequently gets into very
poor condition. The fawns, which may be either one or two in number,
and very rarely three, are produced in the spring.

In America, where the elk is known as the moose, and the former
name transferred to the wapiti, the general habits of the animal are very
similar to those of its European brother. Its favourite haunts are the dense
thickets situated round the swamps and shallow lakes near the sources of
the North American streams. In winter they resort to higher ground,
where boughs are abundant; and at this season, as the snow increases in
depth and hardens, one or more individuals form what is known as “ moose
yards,” which may be of very considerable extent, and are kept open by
constant trampling. It is commonly said that the males and females “ yard ”
by themselves, but this is expressly denied by Mr. C. C. Ward,! although
it seems to be ascertained that very old males will often winter by them-
selves. The ‘‘yard” is situated in a district where white poplar, maple,
and mountain-ash are abundant ; and although the latter of these form the
staple food of the imprisoned elk, yet the needles and young twigs of
juniper, balsam-fir, and other conifers are also consumed. ‘ During their
writes Mr. Herrick,? “‘ the
accessible shrubs are very closely cropped, but ordinarily the tree is not

’

confinement in yards at the height of winter,’

killed, since only one side is stripped. The bark is removed to a height of
ten feet, as the animal rears upon its hind legs and peels the bark. The
direction a herd is moving may be ascertained by one familiar with their
habits, since the bushes browsed are pulled towards the animal. The fir
trees are browsed but the bark is not eaten, yet hunters state that young
firs suffer more than other species from the habit of the males of rubbing
their heads upon them in such a way as to apply the balsam to the abraded
skin about the horns.”

Although the union is only for a season, the male elk in America, as
in Scandinavia, selects a single female at the commencement of the pairing-
season, with whom he retires to the densest thickets for a period of three
weeks or a month. Here the two remain peaceably, unless disturbed by
the intrusion of another male, when a fierce struggle immediately takes
place. The antlers of the males are shed during January, and the new pair

1 In Mayer’s Sport with Gun and Rod, p. 159.
* Mammals of Minnesota, p. 273.

Johnson’s Elk 59

attain their full development by August. While the antlers are in velvet
elk are extremely fond of wading deep into the water, where they lazily
crop the spreading leaves of the yellow water-lily, or pull up its succulent
roots from the mud. An elk which lived for some time at Woburn Abbey
displayed this habit very frequently, and was often to be seen standing in
one of the lakes in the park up to its belly. American elk seem to be less

_ given to repose throughout the greater part of the day than are those of

Norway, and are stated to feed during the mid-day hours. As soon as the
antlers are free from velvet, the males leave the denser thickets to take to
more open ground, where they commence calling. This call may be
answered either by the challenge of a rival male, or by the softer note of
an expectant female. When about to give birth to her fawn, the latter
seeks a secluded situation fairly safe from the intrusion of wolves or bears.
Elk generally run with a long swinging trot, which can be maintained
without any perceptible slackening for an incredible length of time; but
if forced into a gallop, they soon tire. ‘* Formerly,” writes Mr. Lloyd,
‘these animals were made use of to draw sledges ; but owing, as it was
said, to their speed frequently accelerating the escape of people who had
been guilty of murder or other crimes, their use was prohibited under great
penalties. Though I apprehend these ordinances, if not abrogated, are
obsolete, I am not aware that elk are ever made use of in that kingdom at
the present day either to draw a sledge or for other domestic purposes.”
In a later work (Scandinavian Adventures) the same author mentions that he
has known several instances of Swedish elk which were taken young becoming
as tame as domestic cattle; and Mr. Caton bears similar testimony with

regard to the American elk.

2. JOHNsoN’s ELK—ALCES LATIFRONS (Extinct)

Cervus l/atifrons, Johnson, Ann. Mag. Nat. Hist. ser. 4, vol. xu. p. 2
(1874).

Alces latifrons, Lydekker, Cat. Foss. Mamm. Brit. Mus. part 11. p. 78
(1885) ; Dawkins, Brit. Pleist. Mamm. part vi. Cervida, p. 1 (Pal. Soc. 1887).

Cervus alces latifrontis, Pohlig, Paleontographica, vol. xxxix. p. 238
(1892).

Characters.—Probably closely allied to the next species, with which,

1 Field Sports of the North of Europe, p. 336.

60 Elk

when compared with the living species, it agrees in the greater relative
length of the beam and the simpler palmation of the antlers, of which
some good examples are figured by Professor Dawkins in the: monograph
cited.

Distribution —Europe, in the Lower Plistocene epoch, the known
remains having been obtained from the Norfolk forest-bed and Thuringia.

3. Scorr’s Erx—Atces scorti (Exzict)

Cervus americanus, Harlan, Fauna Americana, p. 245 (1825), mec
Erxleben, 1777.

Cervalces americanus,’ Scott, Proc. Ac. Philadelphia, 1885, p. 181.

Characters.—Beam of antlers relatively much longer than in 4. machivs,
and palmation smaller and more distinctly divided into an anterior and

= /
Beet | | g
eo ay
« — =
ae, ee e
\ ee | t es

Fic. 14.—Skull and Antlers of Scott’s Extinct Elk from the Plistocene of North America.
After Dr. W. B. Scott.

posterior branch, with two outwardly directed snags near their line of
division. Nasals longer, and the premaxille extending upwards to unite
with them.

Distribution.—North America, in the Plistocene period. Although this
is undoubtedly a more generalised form, I see no reason for separating it
generically from the existing elk. The outward direction of the beam of
the antlers shows that there is no approximation to Cervus.

| This species requires a new name, Cervus americanus being preoccupied.

Typical Deer 61

III]. Tue Typitcat DEER—GeENus CERVUS

Gerurs Wann. Syst. Nat. ed, 12, vol.1. p. g2 (1766).

Lateral metacarpals represented only by their upper (proximal) ends.
Antlers arising at acute angles to the median line of the skull (as in the
following genera), at first projecting from the plane of the forehead, and
then continued upwards nearly in that plane, supported on short pedicles,
and furnished with a brow-tine arising close to or a short distance above

Fic. 15.—Antlers of Red Deer. From a German specimen in the collection
of Viscount Powerscourt.

the burr, never regularly forked at first division, but generally of large size
and more or less complex structure ; skull without frontal ridges forming
the bases of the pedicles of the antlers. Muzzle with a considerable naked
area surrounding the nostrils; ears generally large ; tail moderately or
very short; face long. Pelage variable ; main hoofs long and pointed ;
lateral hoofs moderate. A metatarsal gland and tuft,’ situated (except in
C. albirostris) high up on the cannon-bone, present, but no tarsal gland or
tuft. In the skull the nasal bones (as in the following genera) well

1 Absent in C. duvaucel.

62 Elaphine Group

developed, the gland-pit large and deep, and the unossified vacuity
extensive ; face-gland variable. | Upper canines small or wanting; cheek-
teeth large and more or less high-crowned. Stature generally large or
medium, and rump low.

The question of the limitations of the genus is one on which very
different views are entertained. Taking the antlers as a basis of classifica-
tion, the forms without a brow-tine, such as Pére David’s deer, may be
eliminated. With regard to the others, the sikine deer are related so
closely to the red deer group on the one hand, and to the fallow deer
on the other, that sub-generic division seems preferable in their case. The
Oriental deer, forming the rusine and rucervine groups, are more distinct,
but, on the whole, as they possess the brow-tine, it seems more convenient
to include them in the same genus, which will thus embrace all the larger
Old World deer with brow-tines to the antlers of the males, and the
antlers themselves bearing at least three tines.

Distribution —The Holarctic and Oriental regions ; represented in the
Western Holarctic only by the wapiti.

i. THE Rep Deer, or ELApHINE Group—Sus-GeEnus CERvus

Elaphus, H. Smith, in Grifiith’s Ammal Kingdom, vol. v. p. 307 (1827).
Pseudocervus, Hodgson, Ff ourn. As. Soc. Bengal, vol. x. p. 904 (1841).
Strongyloceros, Owen, Brit. Foss. Mamm. and Birds, p. 470 (1846) ;
Fitzinger, SB. Ak. Wien, vol. Ixviii. part 1. p. 850 (1873).
Characters.—Antlers rounded, complex, usually with five or more tines,
generally including a bez (second), and always a trez (third). Pelage of
adult, at least in winter, uniformly coloured, generally with a large pale-
coloured area surrounding and including the tail, below which it usually
passes into white; in the young, spotted with white ;_ metatarsal tuft
coloured like rest of leg. Throat maned ; ears large; tail short. Naked
portion of muzzle not extending much on to the upper surface of the face,
with its superior border forming an acute angle, and the portion below the
nostrils constricted to a narrow bar, and then expanding to join the upper
lip. Upper canines present ; upper molars moderately tall, with a flattened
additional column on the inner side. The face-gland, or tear-pit, always

of moderate development. Size large.

Characters 63

As regards the number of their tines, the antlers of this group are
evidently the most specialised of all the existing members of the genus, no
other group having brow, bez, and trez all normally developed. Cupping
in the crown of the antler, as a normal feature, seems to be in the main
confined to the red deer, where it is most developed in its typical or
western form, although occasionally found in the wapiti; and as we
proceed eastwards the type of antler gradually simplifies. It has been
suggested that “cupping” is a modern feature solely due to high feeding ;
but it is observable in the antlers preserved in several of the old German
castles, and also in many prehis-
toric specimens from the British

fens. This is finely exemplified |
in a magnificent pair of antlers
from an Irish bog preserved in the
British Museum, one of which is
figured on page 472 of Owen’s
British Fossil Mammals and Birds.
All the members of the red deer
group from Central Asia have

comparatively simple uncupped
antlers, the most primitive type
being displayed by Thorold’s

deer, in which the antlers are

flattened and have no bez-tine. Fic, 16.—Antlers of Red Deer from Scotland, with fully
This type of antler approximates ote cups. (Rowland Ward, Records of Big
closely to that of the sikine group, -:
from which the elaphines are doubtless derived. Central Asia may, indeed,
be regarded as the original home of the present group, where it probably
developed from the sikines. One branch from this primitive stock seems
to have spread westward to culminate in the modern red deer, and suc-
ceeded in reaching as far south as North Africa, at a time when that area
was in immediate connection with Europe. A second branch is formed
by the wapitis, of which two races are still inhabitants of Central and
North-Eastern Asia, while the third and fourth migrated by way of what
is now Bering Strait to North America.

The generally uniform coloration of the adult pelage, and the develop-

64 Elaphine Group

ment of the large light area in the caudal region, must likewise be regarded
as specialised features ; but it is interesting to note that the caudal disk is
least developed in some of the Central Asiatic species, in’ which the
markings are very like those of the sikas. Evidence of affinity with that
group is further proclaimed by the dappled coat of the fawns. And
although it is generally stated that the adult pelage is entirely unspotted,
one or more rows of such spots may frequently be detected in English red
deer hinds in the summer coat, and a herd of Caspian red deer in the

same pelage living at Woburn Abbey in the summer of 1897 were

Swaytynn

Fic. 17.—Group of Red Deer at Woburn Abbey. From a photograph by the Duchess of Bedford.

almost as much spotted as many sikas at the same time of year. In all
members of the present group the velvet of the antlers is of a grayish
fawn colour.

Distribution.—The Holarctic region, dating from the Plistocene epoch.

1. THE Rep DEER—CERVUS ELAPHUS

Cervus elaphus, Linn. Syst. Nat. ed. 12, vol. i. p. 93 (1766) ; Jenyns,
Brit. Vert. Animals, p. 37 (1835); Bell, Brit. Quadrupeds, p. 394 (1837) ;
Keyserling and Blasius, Wirbelth. Europ. p. 26 (1840); P. L. Sclater,
Trans. Zool. Soc. vol. vii. p. 342 (1871); Fitzinger, SB. 4k. Wien, vol.
Ixix. part i. p. 565 (1874); Ward, Records of Big Game, p. 22 (1896) ;
Satunin, Zool. fahrb. Syst. vol. ix. p. 309 (1896) ; Biichner, Ann. Mus.
Zool. St. Petersburg, 1896, p. 387.

Red Deer 65

Cervus (Elaphus) elaphus, H. Smith, in Grithth’s Animal Kingdom, vol.
IMO. Vv. pe 307 (1827).

Cervus primigenius, Kaup, “fahrb. Mineral. 1839, p. 168.

Cervus priscus, Kaup, op. cit. p. 297 (1839).

Cervus (Strongyloceros) elaphus, Owen, Brit. Foss. Mamm. and Birds,
p. 472 (1846).

Characters—Height reaching to 4 feet, or 4 feet 6 inches, at the
shoulder. Antlers rounded throughout, normally with a bez-tine and more
than five points; when fully developed, forming a more or less distinct
cup at the crown, the brow-tine rising close to the burr, and frequently
longer than the bez, and the fourth tine not specially enlarged and not
situated in the same plane as the portion above, but in fully developed
specimens more or less completely included in the terminal cup. Tail
rather long and pointed, and the light caudal disk of moderate dimensions ;
ear longer than half the head; general colour of adult summer pelage
typically reddish brown, of winter pelage grayish brown; mane not
markedly darker than rest of coat, and under-parts lighter than the back ;
no white on the muzzle. Cry of male in the breeding-season a distinct
roar, somewhat like that of a leopard.

Although the great cupping and complexity of the crown of the antlers
of the typical European red deer indicate great specialisation, yet the red
summer pelage, which is evidently a retention of the colour of the fawn
and of that of the sikas, as well as the comparatively long tail, shows that
in some respects the species is more generalised than several of the Asiatic
forms and the wapiti, in which the red tinge is more or less completely
lost, spots never show themselves in the adults, and the tail is shorter.

Distribution Europe, North Africa, Asia Minor, and Northern Persia.

a. EUROPEAN RAcE—CERVUS ELAPHUS TYPICUS

Cervus elaphus germanicus, Desmarest, Mammalogie, vol. uu. p. 434
(1822) ; Fitzinger, Bestr. Landesk. Osterreichs, vol. i. p. 317 (1832), SB.
Ak. Wren, vol. \xix. part i. p. 573 (1874).

Characters —Height at shoulder reaching to about 4 feet (often less in
Scotch examples) ; face comparatively short and blunt; general colour of
pelage of upper-parts reddish brown in summer and grayish brown in

K

66 Elaphine Group

winter,’ the caudal disk being pale straw, passing into white on the inner
side of the thighs, and in adults bordered by a blackish band ; face, throat,
legs, and under-parts more or less slaty gray ; antlers, when fully developed,
attaining great complexity in the crown, the number of points on each
frequently reaching twelve or fourteen.

As in the other varieties, the summer pelage is short and glistening,
but that of winter is longer and rougher. During the summer the head
and legs are grayer than the body, and the throat of a still paler gray, the
fringe of long hair on the throat attaining its greatest development in the
breeding-season. Regarding the colour of the European form, Mr. A.
Gordon Cameron writes as follows: ‘‘It is the fashion nowadays to say
that red deer are not red, but the present writer is not of this opinion.
Place a band of deer, whether stags or hinds, among green bracken under
an August sun, and it will be seen at once that red deer are rightly named.
While red is the prevailing colour, deer are met with in all shades, from
dark brown, verging on black, to pale cream or dun, verging on white.
Black deer and white deer are on record. Attempts to establish any fixed
rule or relation between colour and condition—e.g. that good stags are
iron gray, or that good hinds (to shoot) are blue—seem scarcely justified
by facts, for plenty of good stags and hinds are red or yellow by nature,
and the pink of condition will do no more than put an extra gloss on their
coats. Wasting deer tend to grow paler in colour, and look ragged
from loss of bloom.” In park deer, spots, especially along the middle of
the back, are occasionally visible in the summer pelage.

The typical red deer varies greatly in point of bodily size and in the
relative development of its antlers. The largest examples are met with in
the forests of the Continent, and the smallest in islands, such as Harris.
The antlers of Scotch hill deer, although comparatively small, yield to none
in symmetry of form and gracefulness. By far the finest series of antlers
in existence are those preserved in the King of Saxony’s castle at Moritz-
burg, near Dresden, many of which have been figured by Dr. A. B. Meyer
in the work cited in the Appendix. Among these, specimens with twelve,
thirteen, and fourteen, or even more points on each side are by no means
uncommon. Sir Victor Brooke observes that ‘‘a comparison of the gigantic
antlers of the red deer of the fifteenth, sixteenth, and seventeenth centuries

1 Park deer frequently remain more or less red throughout the year.

Red Deer 67

preserved in the old hunting schloss at Moritzburg (built 1540), near
Dresden, with antlers from the islands of Harris and Sardinia shows in a most
striking manner the great variations to which this species is subject. Some
of the antlers at Moritzburg measure 50 inches along the outside curve, are
1o inches in circumference round the smallest part of the beam, and the two
antlers carry from twenty-four to fifty points. The spread between the
coronal tines of one specimen is 74 inches. Antlers from Harris and
Sardinia rarely exceed 30 inches in length, their circumference being about

SWipvre

Fic. 18.—Antlers of Red Deer. From a specimen in the Castle at Moritzburg, Saxony.
After Dr. A. B. Meyer.

4 inches ; they very rarely carry a larger number than twelve points, and
their span seldom exceeds 28 inches.”

A pair of the Moritzburg antlers are shown in Fig.18. Nothing like these
are to be found at the present day, unless it be occasionally in Hungary,
and as these fine specimens are not recorded in Mr. Ward’s book, it is use-
less to quote the smaller measurements given there. The contrast between
the Moritzburg antlers and those of the Caspian red deer is very marked ;
and it is very rarely that the short bez-tine so often found in the latter is
noticeable in the former. Next to the Moritzburg collection comes one

belonging to Count Arco-Zinneberg, now preserved in his house at Wittels-

68 Elaphine Group

bacher Platz, Munich. Among British collections, that of Viscount
Powerscourt, at Powerscourt, Enniskerry, Ireland, claims a very prominent
position, one of the specimens from this series being represented in Fig. 15.
The great hall at Hampton Court contains a small but unusually fine series
of red deer antlers, all of which appear to be of great antiquity, and doubt-
less came from the Continent. Several of them appear equal in size and
number of points to some of the Moritzburg specimens, and the collection
is probably unrivalled in Britain. It also includes a few examples of
wapiti antlers, and some remarkably fine specimens of those of the elk.

Fig. 19 shows the antlers of a wild red deer from Exmoor. It has
six tines, and serves to exhibit the difference between such red deer antlers
and those of the wapiti (Fig. 24, p. 95), the fourth tine being com-
paratively small, and projecting externally to the beam, quite out of the
plane of the two terminal tines, whereas in the latter the fourth tine is
very large, and projects inwardly in the plane of the two terminal tines.
In the island of Jura there exists a remarkable breed of red deer, locally
known as ‘‘ cromies,” and characterised by the peculiar downward curvature
of their antlers. Although deer with these appendages normally developed
also live in the island, the two are stated never to interbreed, but to keep
apart.

Distribution —The greater part of Europe, probably as far east as the
Don and Volga, but exclusive of much of the region in the neighbourhood
of the Black Sea and the Eastern Carpathians ; farther northwards there is
a want of definite information with regard to the eastern range.’ Sir
Victor Brooke gives the following list of countries and districts now
inhabited by wild red deer of the typical variety, viz. Ireland, County
Kerry ; Great Britain, Devonshire and Somerset, Highlands of Scotland, and
the islands of Harris, Skye, Rum, Mull, Jura, and Arran ; Isle of Hitteren,
Norway ; the south of Sweden, France, Spain, Austria-Hungary, Turkey,
and Greece. Italy, it will be noticed, is omitted from this list, although
it is included in the habitat of the race in Bell’s British Quadrupeds ;
and from Switzerland the red deer has long since completely disappeared.
From a series of heads belonging to the Duc d’Orléans, which I have lately
had the opportunity of seeing, it appears that the Spanish red deer has a very

' Dr. E. Biichner has published a paper on this subject, but does not clearly distinguish between the
typical and Caspian races.

Red Deer 69

red summer pelage (the specimens were shot towards the end of March),
and that the antlers are comparatively weak, markedly curved, and with
the bez-tine generally, if not invariably, wanting. Not impossibly the
Spanish deer may prove to be closely allied to, or perhaps identical with,
the North African race of the species. As already mentioned, antlers in-
distinguishable from those of the typical race are of common occurrence in
the fens and some of the other more superficial deposits in the British

Fic. 19.—Frontlet and Antlers of Red Deer. From an Exmoor specimen in the possession of
Lord Elphinstone.

Islands, but those from some at least of the Plistocene beds of both Britain
and the Continent appear referable to the Caspian variety.

Habits —So much has been written on the mode of life of the red deer,
especially in Scotland, that a comparatively short notice will suffice on this
occasion. For the following paragraphs I am indebted to Mr. Allan
Gordon Cameron, who has had special opportunities of observing Scottish
red deer in their native haunts. In the Highlands red deer are found
on the open hills, seeking food and shelter in storm-swept glens, and in
this respect living under conditions quite different from those natural to
the species in other lands. Isolated for centuries from their continental

70 Elaphine Group

relatives, they exhibit in their relatively small dimensions the character-
istic of an island race, retaining, nevertheless, amid the sterile grandeur of
their mountain home an elegance and dignity of appearance unsurpassed
by any of their congeners. The hinds breed in their third year and calve
when three years old, producing, with rare exceptions, but one at a birth,
and remaining fertile, under favourable conditions, so long as they live.
The fawn is dropped in early summer on the heather, wherein for some
days it will lie closely concealed till able to run with its mother, whom it
will accompany thereafter, if a male, for a year, and if a female, for two
years. The sexes occur at birth in nearly equal proportions, but probably
not more than half those born survive as yearlings, and the mortality of
males seems to exceed that of females from an early age. Hill stags attain
maturity at twelve years old, and appear to decline, as a rule, some five or
six years after, but their maturity is sometimes prolonged, for a Jura stag,
which had reached its prime when first observed, was still in best condition
when killed twelve years later. The calibre of a stag’s head, like the
condition of its body, has its periods of advance, maturity, and decline,
which vary with individuals and cannot be precisely determined ; but,
excepting disease or accident, the same style of head will throughout life
infallibly distinguish the same stag. A Royal stag in Cromarty carried a
twelve-point head for thirteen seasons, and put out a thirteenth point in
his fourteenth season, though his antlers then appeared to be losing weight.
Old stags lose their front teeth, fall off in condition, and deteriorate gener-
ally, and the extreme term of their natural life may be placed at thirty
years. Hinds mature earlier, and are probably, on an average, longer lived
than stags. A Jura hind, which was full grown, with a big fawn, when
first observed, retained all her teeth and was still fertile twenty-one years
later, when accidentally killed, having reared twenty fawns during the
period of observation. With both sexes, and at all ages, the chief cause of
death is parasitic disease—lung-worms, intestinal worms, and liver-flukes,
the first named being by far the most destructive, and probably responsible
for great periodical mortality among fawns. Wet seasons make the highest
death-rate, and spring is the deadly time.

Stags shed their antlers in spring (March to May), earlier or later
according to age and condition, and attain their prime for the year in

autumn (September), shortly after the new antlers are hard and clean, and

Red Deer ia

a little before the pairing-season, which about the middle of October is at
its height. Owing to the large numbers of deer gathered on the bare hills,
the pairing-season in the Highlands presents a combination of scene and
sound without parallel in continental forests. The hills re-echo with the
roaring of rival stags, which roam to and fro, restless and defiant, now roll-
ing in the peat-pools, now rounding up their hinds, and now fighting
desperate battles. With individual deer the spur of sexual excitement will
sometimes totally dispel the fear of man. Thus a wild stag in Jura once

Fic. 20.—A Red Deer in the Park at Woburn Abbey. From a photograph by the
Duchess of Bedford.

took possession of some tame hinds that were accustomed to be hand-fed,
and for the time being resolutely guarded them from all interference.
Contrary to the general opinion, mastership in the herd goes to the
weightiest and most vigorous stag without respect to the calibre of his
antlers, so-called “bald” stags being not unfrequently masters of large
herds. Stags lose condition rapidly during the pairing-season, at the close
of which they are completely exhausted.

Except in the pairing-season, adult stags and hinds do not associate
freely, but lead to some extent independent lives in large or small bands,

72 Elaphine Group

the numbers and distribution of which are determined chiefly by conditions
of food and weather. In summer, with fine weather and abundant food,
deer are scattered all over the ground in small parties and at. high eleva-
tions ; in winter, when food is scarce and the weather severe, they shift to
lower levels, are more concentrated, and seek sheltered places. Hinds have
a strong homing instinct and seldom leave the ground where they are bred ;
but stags are great travellers and exhibit a twofold migratory impulse,
namely, a general shift from summer to winter quarters, and a special shift
from summer quarters to the accustomed pairing-ground, the same places,
however, being sought at the same season year by year. Deer take the sea
fearlessly and swim strongly, stags introduced into Hoy Island, Orkney,
having swum southwards to the coast of Sutherland, and northwards as far
as the Bay of Skail, in the Mainland of Orkney, in either case a distance,
as the crow flies, of eight miles. Hill deer, like wood deer, feed in the
morning and evening and rest during the day. They are dainty feeders,
requiring a wide range for their support, since they neither bite closely on
a limited area like sheep, nor tear coarse herbage with the aid of a pliant
tongue like cattle, but pick here and there, nipping the tender tops of
plants that suit them, and moving forward all the time to fresh places.
The appetite of both sexes for bones and antlers, which they grind with
the cheek-teeth, is a familiar fact, as is also their taste for sea-weed, of
which sea-coast deer partake freely, while not a few stags frequenting
inland districts make periodical journeys to the coast for its sake. High
ground affords the sweetest pasturage, and deer that dwell habitually on
the hill-tops make the best venison. It is a habit of deer, though probably
not an invariable one, to drink before feeding. Stags accustomed to herd
together exhibit a strong affection for their companions, living in perfect
harmony, and being always under a recognised leadership, but whether of
the oldest or heaviest stag cannot be determined. Hinds are less cere-
monious, more independent, and occasionally quarrelsome, often fighting
with their fore feet, and rearing straight up on their hind legs.
Although not specially keen-sighted, red deer have their senses of smell
and hearing acute. Hence, when disturbed or travelling, they always
move to windward, and when resting their favourite position is on the
lee side of sloping or rising ground, just below the ridge or crest, facing
the landscape with the wind in their backs. Stags that have been shot at,

Red Deer re

or otherwise alarmed, have often been observed to travel some distance and
then deliberately lie down in such a position that danger approaching
from in front is seen, and danger from behind scented. Hence they are
most readily approached after feeding. As mentioned by Mr. Cameron in
the foregoing notes, the red deer hind attends to the needs of her offspring
with remarkable assiduity ; the fawn being generally born among heather
or other suitable covert, where it is left alone during the day, and visited
by its parent at nightfall. To make it lie down, according to Mr. Scrope,
the mother presses the fawn with her nose ; and when once settled com-
fortably, remains curled up throughout the day. The hind does not, how-
ever, depart far from her offspring, which she is always ready to defend
from danger. This affectionate care of the hind for her fawn——-whether
the species referred to be the red deer or the fallow deer—did not escape
the notice of Xenophon. As translated by Mr. H. G. Dakyns, he has the
following observations in his essay on hunting. ‘‘ As day breaks, he [the
hunter] will espy the hinds leading their fawns to the places where they
will lay them severally to rest. Having made them lie down and suckled
them, they will cast anxious glances this way and that to see that no one
watches them ; and then they will severally withdraw to the side opposite
and mount guard, each over her own offspring. . . . When his eye has
lit upon the object of his search, he will approach quite close. The fawn
will keep perfectly still, glued as it were to earth, and with loud bleats
suffer itself to be picked up; unless it happen to be drenched with rain ;
in which case, it will not stay quiet in one place. .. . The huntsman
having seized the fawn, will hand it to the keeper. The bleating will
continue ; and the hind, partly seeing and partly hearing, will bear down
full tilt upon the man who has got her young, in her desire to rescue it.
. . . Young fawns may be captured in the way described. Those that are
already big will give more trouble, since they graze with their mothers and
the other deer, and when pursued retire in the middle of the herd or
occasionally in front, but very seldom in the rear. The deer, moreover, in
order to protect their young will do battle with the hounds and trample
them under foot ; so that capture is not easy, unless you come at once to
close quarters and scatter the herd, with the result that one or another of
the fawns is isolated.” All this is almost absolutely true to nature.

L

7 4 Elaphine Group

5. CorsicAN Racr—CERVUS ELAPHUS CORSICANUS

Cervus elaphus corsicanus, Erxleben, Syst. Regn. Animal. vol. i. p. 304
(1777).

Cervus mediterraneus, de Blainville, ‘fourn. Physique, vol. xciv. p. 262
(1822) ; Gervais, Hist. Nat. Mamm. vol. ii. p. 216 (1855).

Cervus elaphus minor, Fitzinger, SB. Ak. Wien, vol. |xix. part i. p. 575
(1874).

Characters—The smallest race of the species, and according to Sir Victor
Brooke connecting the European with the North African race, although
Gervais regards it as closely allied to the latter, with which it is identified
by M. Lataste. If this latter view be correct, the name corsicanus, as the
earlier, must supersede darbarus. General colour of the pelage of the upper-
parts dark brown, becoming blackish in winter.

Distribution.—Corsica and Sardinia. Fitzinger, on the authority of
Polybius, states that deer are not indigenous to either of these islands, and
were introduced by human agency, but this requires confirmation.

c. NorTH AFRICAN RACE—CERVUS ELAPHUS BARBARUS

Cervus barbarus, Bennett, List Anim. Gardens Zool. Soc. p. 31 (1837) ;
Gray, Cat. Ungulata Brit. Mus. p. 197 (1852), Cat. Ruminants Brit. Mus.
p. 68 (1872) ; Sclater, Trans. Zool. Soc. vol. vil. p. 344 (1871) ; Fitzinger,
SB. Ak. Wien, vol. |xix. part 1. p. 577 (1874).

Cervus corsicanus, Lataste, Act. Soc. Linn. Bordeaux, vol. xxxix. p. 286
(1885).

Characters —Smaller than the typical, but larger than the Corsican race ;
generally lacking the bez-tine to the antlers ; pelage (? summer) dark brown,
with a grayish brown streak down the middle of the back, and small
irregular whitish spots on the flanks, and sometimes on the back.

Although formerly living in the menagerie at Knowsley, and in the
London and Paris Zoological Gardens, this deer appears to be now very
rarely imported, and I have never seen an example. The British Museum
possesses several pairs of antlers which are curved forward near the middle

of their length somewhat after the manner of those of the shou. Since,

ed: Deer as

however, all these very probably belong to the same animal, this may be
merely an individual peculiarity. According to Sir Victor Brooke, the
still smaller red deer from Sardinia and Corsica “‘completely bridge over
the characters which have been advanced as distinctive of the African and
European races.” The red deer being essentially a northern form, the loss
of the bez-tine in the southern race may be due to degeneracy owing to the
unsuitability of a warmer climate to full development.

Distribution North-Western Africa, in Tunisia and Algeria, more
especially in the forests of Constantine between Bone and Calle and also in
the district of Tabessa. On the plains in the neighbourhood of Douirat a
correspondent of M. Lataste states that this deer is found in open country
where there is not a vestige of a tree. To the Moors it is known as the
alwassi. The possibility of the Spanish red deer belonging to this race has
been already mentioned.

I have no information as to the habits of this race, which are, however,
doubtless generally similar to those of the typical form.

d. CasPp1AN RacrE—CERVUS ELAPHUS MARAL

Cervus maral, Ogilby, Rep. Council Zool. Soc. 1840, p. 22; Sclater,
Trans. Zool. Soc. vol. vil. p. 336 (1871) ; Gray, Cat. Ruminants Brit. Mus.
pCon(to72) jsbitzinger, SB. Ak. Wien. vol. |xix. part i. p. 597 (1874) ;
Brooke, Proc. Zool. Soc. 1878, p. 812; Satunin, Zool. “fahrb. Syst. vol. ix.
p- 309 (1896) ; Ward, Records of Big Game, p. 30 (1896).

(?) Strongyloceros speleus, Owen, Brit. Foss. Mamm. and Birds, p. 469
(1846).

(?) Cervus elaphus varius, Fitzinger, SB. dk. Wien, vol. \xix. part i.
P- 574 (1874).

Cervus elaphus antiqui, Pohlig, Paleontographica, vol. xxxix. p. 246
(1892).

Characters.—Size somewhat larger (the height at the shoulder reaching
to 4 feet 6 inches) than in the typical race of the species, the build stouter,
the neck thicker, and the head longer and more pointed. General colour
of summer pelage red, in immature animals frequently, if not invariably,
marked with numerous yellowish spots ; winter pelage dark slaty gray, with
the caudal disk a deeper yellow than in the typical race, and the shoulders,

76 Elaphine Group

thighs, and under-parts nearly black. Antlers large and _ less complex

than those of the latter, the number of points seldom exceeding eight,
and being often only six ; the bez-tine, which may be absent, frequently
much shorter than the brow-tine, which is long and much curved

upwards ; and the fourth tine generally more distinct from the crown.

Head of Caspian Red Deer, from a specimen shot in Asia Minor by Mr. F. C. Selous.
(Rowland Ward, Records of Big Game.)

That this form is not entitled to rank as a distinct species seems fairly

certain. It appears impossible to formulate a precise definition, but the

race may be roughly described as large red deer with relatively long faces,
spotted summer coats when immature, and with large antlers of a simpler
type than equally large and fully adult specimens of the typical race, the
bez-tine being often proportionately much shorter than in the latter. This

Red Deer 77

last feature is, however, by no means constant, and in the handsome head
represented in Fig. 21, the brow- and bez-tine are of approximately equal
length. Out of four fine specimens obtained by Mr. E. N. Buxton, of
Knighton, Buckhurst Hill, from Tartarow, in the Galician Carpathians,
two have the bez-tine very short, in the third there is a very short bez on
one side only, and in the fourth this tine is absent on both sides. On the
other hand, in the seven-pointed Hungarian antlers from the collection of
Viscount Powerscourt shown in Fig. 3, the brow- and bez-tine are of
approximately equal length, so that it is evident that the shortness of the
latter can in no wise be regarded as invariably distinctive of the race. At
the same time it must be remembered that Hungary is on the borderland
between the habitats of the two races; and it is therefore to be expected
that antlers from this district would exhibit intermediate characters.
Indeed, it is only provisionally that Lord Powerscourt’s specimen is
assigned to the eastern race.

Antlers seldom, if ever, attain the extreme complexity of the typical red
deer as exemplified by old German examples (Fig. 18) or specimens from
the English peat or Irish bogs ; and Fitzinger gives the number of points
on the crown as not exceeding two or three. But a specimen from the
North-Western Caucasus, presented by Mr. St. George Littledale to the
British Museum, has eight points, and Mr. Ward records another with
eleven from the same range. Mr. Buxton also has a many-pointed specimen,
although the palmation of the crown indicates that it is somewhat abnormal.
In the Caucasus this deer is known as the ollen, and Mr. St. George Little-
dale ' has published the following remarks regarding its characters and antlers.
“When I hunted the ollen,” he writes, ‘I had no notion that I should ever
be called upon to carefully discriminate between them and their kin in
other countries, so that I am obliged to rely upon my memory for any
points of difference, and memory only suggests that whereas the wapiti
rarely (if ever) has cups on his antlers, the ollen royal has the peculiar cup-
formation as often as the red deer. Again, the call of the Caucasian stag
in the rutting-season (September) is similar to that of the Scotch stag, and
does not resemble the weird whistle of the wapiti. In size both of body and
antler the ollen comes very near to the great American stag.” With regard
to the cupping of the antlers, the plate annexed to Mr. Littledale’s note

1 Badminton Library, Big Game Shooting, vol. i. p. 36.

78 Elaphine Group

shows seven heads of his shooting, but only two of these appear to show the
perfect cup comparable to those of old continental examples of the typical
red deer, and then only on one side. One of these heads is figured at the
end of the present volume.

Antlers of this race from Asia Minor and the Crimea not unfrequently
exhibit a great paucity of tines, with a corresponding development of the
beam. In the Proceedings of the Zoological Society for the year 18go0 I
have given a figure’ of an abnormal antler from Asia Minor doubtless
belonging to this race, in which there is no tine above the brow till the
crown, which is much expanded and bears a number of points. Another
antler from the Crimea, represented in the same plate, consists merely of a
beam with a couple of tines at the summit ; and antlers of such abnormal
type seem by no means uncommon in the Crimea and Asia Minor, although
I have seen none precisely comparable from the habitat of the typical race.

In the summer of 1897 a herd of red deer from some part of the
Caucasus or Transcaucasia was added to the collection at Woburn Abbey.
It comprised a sub-adult stag and several hinds, all in summer pelage. The
hinds were somewhat larger than ordinary red deer, and the stag had the
long brow and short bez-tine which seem very characteristic of this race ;
all showing the long, slender, maral face. The coat in both sexes was
distinctly red, marked with a large number of yellow spots ; so much spotted,
indeed, that these deer had almost the appearance of large sikas. Although
a few spots may be seen on the back of some typical red deer hinds in the
summer pelage, I have never seen any comparable in this respect to the
Caucasian herd, neither have I ever seen the stags so spotted. Possibly the
C. elaphus varius of Fitzinger is based on the Caspian race. The specimens
are important in showing that the summer pelage of the hinds and young
stags is reddish. In winter these deer turned very dark gray, becoming
nearly black on the shoulders, thighs, and under-parts ; they had a dark
stripe down the middle of the back, and a dark patch in front of the caudal
disk, which was bordered with black. ‘This type of coloration, as well as
the spots in summer, are stated by Fitzinger to be characteristic of all
specimens not fully adult, but the blackness is well shown in the old stag in
the British Museum; and Mr. Sclater tells me that he saw a spotted herd
at St. Petersburg. Several of the Woburn specimens are shown in Figs. 1

1 Plate xxx.

Red» Deer 79

and 2, where they are the largest members of the groups. I have no
information as to the summer colouring of Carpathian examples.

In its generally simpler antlers this race apparently forms a partial
transition from the typical race in the direction of the three under-mentioned
Asiatic species; and it is noteworthy that examples in which these
appendages attain unusual complexity are recorded from the Carpathians and
Caucasus, where this race must impinge very closely on the habitat of the
typical red deer, with which it not improbably intergrades. The name
““maral” being the Persian equivalent for deer in general, it is applied
throughout a great portion of Asia to all the larger members of the genus.
Hence much confusion as to what constitutes the true maral in the zoological
sense of the word, the Altai wapiti being commonly designated by this term.

Distribution —The typical locality of this race of the red deer is the
Caspian provinces of Northern Persia, where it is the only representative of
the present group of the genus, as it also is in the Crimea and probably in
Asia Minor. Thence the range extends into Transcaucasia, the Caucasus,
probably Circassia, and the Galician Carpathians. With regard to the
Caucasus, Dr. Satunin speaks of the red deer as being common in many parts
of that range and in Transcaucasia, but doubts if the “ maral” occurs there,
although he rightly says that the latter is the only stag in the Crimea.
What distinction he finds between the Crimean and Caucasian deer I have
no means of knowing, but both must apparently be referred to the present
race, although the typical race may extend into the Northern Caucasus.
The Carpathian deer are referred to the present race in Mr. Rowland
Ward’s book, and Mr. Buxton’s specimens indicate the apparent correctness
of this reference. His specimens, as already mentioned, are from the
Galician or Eastern Carpathians, and information is urgently needed as to
how far we have to travel westward along the range before entering the
habitat of the typical red deer. Mr. Ward uses the name of Carpathian
deer for the present animal, but this is obviously inadvisable, as Persia is its
typical habitat, and in the Carpathians we may expect to find it inter-
grading with the western race. Information is also required with regard
to the red deer of Turkey.

Some years ago Dr. A. Nehring’ suggested that the large red deer
antlers from the Plistocene deposits of Europe belonged to the maral, and it

1 SB. Ges. Nat. Berlin, 1887, p. 67.

So Elaphine Group

seems not improbable that the large fragments from Kent’s Cavern, near
Torquay, figured by the late Sir R. Owen under the name of Strongy/oceros
speleus, may be referable to the same race. At a later date, however,
Dr. Hans Pohlig? identified the latter form with Kaup’s C. primigenius, which
he regarded as indicating a race more nearly allied to the typical red deer.
At the same time he described and figured certain antlers from the Middle
Plistocene deposits of Thuringia, under the name of C. e/aphus antiqui. One
of the specimens figured in plate xxv of his memoir shows the short brow-
tine and simple crown so frequently found in the maral, with which race
I have little hesitation in identifying the extinct form. If the C. primigenius
of Kaup be likewise identical, that name must supersede C. maral. The
Plistocene red deer being of the maral type, the highly specialised antlers
of the typical red deer are evidently a late western development which did
not appear till about the date of the fen-deposits.

Habits.—Sir Victor Brooke writes that a pair of maral “ which lived in
one of my parks for some years, kept entirely apart from the red deer
inhabiting the same park. They bred together ; and during the rutting-
season the two species never showed the faintest desire to cross. This
was the more remarkable, as the old stag maral, though considerably
larger in size, lived in great fear of the red deer stags, which during
that season roamed incessantly through the park in search of hinds,
but at all times treated the female maral with sovereign disdain,
although at any time they could have taken possession of her had they
so desired.”

The late Sir O. B. St. John, as quoted in Mr. Blanford’s Zoology and
Geology of Persia, wrote as follows :—‘“ The maral is very numerous in the
forests of the Caspian provinces, but does not occur elsewhere [in Persia].
It is often brought alive to Tehran, and, before the famine, the Shah’s
zoological gardens contained seven or eight specimens, which died of
starvation or were killed and eaten by the keepers.” The roar of the stags
in the Woburn herd, although not quite like that of the European race,
has no wapiti element in its composition.

The following dimensions of antlers of this form are taken from Mr.
Rowland Ward’s book :—

| Palaontographica, vol. xxxix. p. 246 (1892). ‘The recent Siberian skull figured on p. 254 as that

of a maral belongs to one of the Asiatic wapitis.

THe DEER or Att Lanvs. Puarte lil.

BEDFORD’S DEER IN SUMMER PELAGE.

Publighed. by Rowland Ward. Ltd.

Duke of Bedford’s Deer SI

er cpemeeeere TROT Wakhinie Points. Soy
484 6 30 414 5-6 Crimea
48 74 ? is Caucasus
478 2 38 : 6-5 "
47 3 36 374 6-7 :
47 63 213 363 5-5 Crimea
463 52 18 33 8-8 Caucasus
454 Wes 255 358 8-8 s
45% St 42 353 6-7 Crimea
45 8 36 42 TI-I0 Caucasus
454 73 g-6 Asia Minor
444 63 33% 358 5 Caucasus

2. THe DuKeE or BEpForD’s DEER—CERVUS XANTHOPYGUS

Cervus xanthopygus, Milne-Edwards, Ann. Sci. Nat. ser. 5, vol. viii.
p- 376 (1867) ; Rech. Mamm. p. 181, plate xxi (1868-74) ; Fitzinger, SB.
Ak. Wien, vol. \xix. part i. p. 590 (1874) ; Brooke, Proc. Zool. Soc. 1878,
p. 68 ; Ward, Records of Big Game, p. 17 (1896).

Cervus bedfordianus, Lydekker, Proc. Zool. Soc. 1896, p. 932.

Plate III

Characters.—Size about the same as a large red deer. Antlers rounded,
with the brow-tine greatly elongated, and about three times the length of
the very short bez-tine, which is separated by a long interval from the
former ; five tines in those of the third year; the curvature of the beam
regular ; the fourth tine not greatly enlarged, and probably no cup formed at
the crown. ‘Tail shorter than in the red deer, and the caudal disk, when
developed, relatively large. Summer pelage uniformly bright foxy red on
the upper-parts and outer side of limbs; face, throat, and inner side of
limbs bluish gray ; in winter the upper-parts brownish gray, with a large
bright orange caudal disk, the mane being blackish, with a considerable
amount of black on the tips of the hairs. No white on muzzle.

This species was originally described on the evidence of a male from the
Imperial gardens at Pekin, which appears to be an old animal with the
antlers retrograding. In the figure the colour of the pelage is reddish, and
there is a distinct orange caudal disk, with a dark front margin. From the

M

8 2 Elaphine Group

evidence of the next specimen, it would seem that the figure was taken just
before the complete replacement of the summer by the winter coat, at
which time the caudal disk makes its first appearance, apparently through
fading of the original colour. The tail is represented as very long. The
left antler has three, and the right four tines; the brow-tine in the latter
being very long and the bez very short.

The second definitely known example is a young male from near Pekin,
received at Woburn Abbey in the summer of 1896, and provisionally
discribed as a distinct species under the name of C. bedfordianus. The
antlers had four tines, with a distinct excess of the brow- over the bez-tine
in length. At that time the pelage was short, smooth, and glossy, and of a
bright foxy red colour, without the slightest trace of a light disk on the
buttocks round the short tail. There was no throat-mane. By the middle
of September, the summer coat was being replaced by the winter one. The
most extraordinary change was the development of a large yellowish disk
on the buttocks, including all the tail. ‘This disk was clearly produced by a
change in the colour of the hairs of the summer coat ; but it appeared to
be also developing in the winter coat. The general colour of the latter
seemed to be bluish gray, or brown, with a tendency to fawn on the neck.
A distinct fringe had also developed on the throat. This was very thin,
with bands of black, and white tips to the hairs : thus being quite different
from the thick, uniformly-coloured fringe of the wapiti. Still later, the
general colour of the coat became more wapiti-like, and the caudal disk
more distinct and brighter. By the middle of June 1897 the animal had
again assumed its summer pelage, which was precisely similar to that of the
previous year, thus proving that the absence of the light caudal disk was
not a feature due to immaturity. The new antlers had acquired five points,
and were remarkable for the excessive proportionate length of the brow-tine,
which was nearly a foot long, whereas the bez-tine was not more than one-
third the length. In the plate the animal is represented bearing these third
antlers. In general form the antlers come nearest to those of the Caspian
variety of the red deer. The animal, after a long period of ill-health, died
in October 1897, and its head is now mounted in the British Museum.

The large brow- and small bez-tine of the antlers appear sufficient to
justify the identification of the specimen in question with C. xanthopygus,
in spite of the circumstance that the type of the latter is depicted with a

Tae DEER or Att Lanps. Plate Dy.

HANGUL

wbinghed. by Rowland Ward lid

Hangul 8 3

tail of great relative length, which must accordingly be regarded as an error
on the part of the artist.

The red colour and the absence of the caudal disk in the early summer
pelage, as well as the simple antlers, are distinctly generalised characters,
although the extremely short tail is a specialised feature sharply distinguish-
ing this species from the red deer. The appearance of the caudal disk by
fading of the summer pelage is paralleled by the bleaching of this area in the
wapiti, and may indicate the original mode of formation.

Distribution —Manchuria, and probably other parts of Eastern Asia.
How far west this species extends, is a point for future investigation ; the
range given by Sir Victor Brooke includes the areas of other species.

Habits —Although nothing is known of this deer in a wild state, the
specimen at Woburn Abbey lived almost entirely on leaves during the
summer, whereas both red deer and wapiti are to a great extent grazing
animals.

3. THe Hancur—Cervus CAsHMIRIANUS

Cervus cashmerensis, Gray, List Osteol. Brit. Mus. p. 65 (1847),—no
description.

Cervus cashmeriensis, Adams, Proc. Zool. Soc. 1858, p. 529.

Cervus cashmeertanus, Falconer’s Paleontological Memoirs, vol. 1. p. 576
(1868) ; Sclater, Trans. Zool. Soc. vol. vii. p. 339 (1871) ; Gray, Caz.
Ruminants Brit. Mus. p. 68 (1872) ; Brooke, Proc. Zool. Soc. 1878, p. 912;
W.L. Sclater, Cat. Mamm. Ind. Mus. part ii. p. 184 (1891).

Cervus wallichu, Kinloch, Large Game Shooting, part i. p. 44 (1869).

Cervus cashmirianus, Fitzinger, SB. Ak. Wien, vol. \xix. part i. p. 586
(1874) ; Sterndale, Mamm. Ind. p. 512 (1884); Blanford, Fauna Brit.
Ind.—Mamm. p. 535 (1891) ; Ward, Records of Big Game, p. 38 (1896).

Plate IV

Characters—Height from 4 feet to 4 feet 4 inches at the shoulder.
Antlers rounded throughout, with a bez-tine, and usually only five points,
so that no cup is formed at the crown ; the brow-tine generally arising at a
considerable distance above the burr, and usually shorter than the bez ; and
the fourth tine shorter than the fifth, or terminal tine ; the beam much curved
towards the middle line, so that the summits of the terminal tines of the

84 Elaphine Group

opposite antlers are more or less closely approximated. Tail short, and the
light caudal disk, at least frequently, very small, and not extending on to the
upper surface of the buttocks, thus excluding the tail. Ear equal to half
the head in length. Hair on back not reversed, and no white on the
muzzle above the lips. Metatarsal tuft lower down than in the red deer.
General colour of pelage in winter brown, brownish ash, or liver-colour,

Fic. 22.—Head of Hangul, from a Kashmir specimen. (Rowland Ward, Records of Big Game.)

with the individual hairs speckled ; the light area on the inner sides of the
buttocks dirty white, with a dark line on its outer border running down
the inner side of the thigh ; upper surface of tail black ; flanks and limbs
paler than the back ; lips and chin white, and ears whitish internally. In
summer, the general colour lighter and more rufous, with most of the under-
parts whitish, although brown posteriorly in the male. Throat-fringe small
and not markedly darker than the rest of the pelage. Spotting of fawn

Hanegul 85

stated to persist till the third or fourth year, instead of more or less completely
disappearing during the first year, as in the typical race of the red deer.
Cry of male in the breeding-season a loud prolonged squeal, like that of the
Wwapitl.

Although differing by its short tail and the slight development of the
rufous tinge in the summer pelage, this very well-marked species presents
a decided approach to the sikas in the comparatively simple form of the
antlers, the long persistence of the spots, and the, at least frequent, slight
extent of the light caudal area, in which the dark marginal streak down the
thigh is precisely similar to the sikine type. The distance by which the
brow-tine is separated from the burr does not seem to have been previously
noticed as a point of distinction between this species and the red deer, although
it is a very marked one, the interval being sometimes fully an inch and a
half in length.

Distribution.—Kashmir and Yarkand.

a. KasumMiR RAcE—CERVUS CASHMIRIANUS TYPICUS

Characters —The terminal or fifth tines of the antlers so much curved
inwards as to be separated only by a very small interval. The coloration
of the pelage is that described above.

Further information is required as to the degree of development of the
light caudal disk. Mr. Blanford, who apparently describes it as well
developed, writes as follows :—‘‘In Sclater’s figure, from an animal in the
Zoological Gardens, there is no caudal disk; the tail is dark brown above,
pale below, and only the buttocks pale rufous. Whether this is due to
variation in colouring or to age, it is impossible to say, but a skin from the
Zoological Gardens, now in the British Museum, agrees with the figure.”
It is this skin which forms the subject of plate iv. In the original de-
scription given on page 577 of the first volume of Falconer’s Pa/eonto-
logical Memoirs the following words are used, viz. :—‘‘ Hind aspect of
the buttocks marked by a very well-defined patch of white, running up to
and terminating at the angle of junction of the tail, and pursuing down on
the inside of the buttocks towards the thighs; no disk of white on the
rump.” As this description accords precisely with the two specimens
noticed above, it is evident that the slight development of the light caudal

86 Elaphine Group

area 1s at least a very frequent characteristic of the species. It may be
added that the passage quoted refers to a female, but on page 576 of the
same work the absence of a caudal disk is also noted in the male, of which
several examples are described. .

The following are the dimensions of some of the largest antlers re-
corded by Mr. Rowland Ward :—

oisnceae: cium: Tip to Tip, Widest inside. ars

48 ? ? ?

47 ? P ?

47 78 21 36 To
47 64 30 358 8-8
458 8 35 41 6-6
458 6 254 36 8-8
45 65 19 34 6-6
445 7 20 43 5-5
443 83 313 444 ba
44 74 30% 408 5-5
44 6 27 36 sn

Colonel J. G. Allan sends me the following dimensions of a remarkably
symmetrical pair of antlers, with six tines on each side, shot by himself in
Kashmir. Outside length of right antler 45 inches, of left 441; extreme
inside width, 37 ; outside span, 42 ; maximum distance between points, 52
inches.

Distribution—The valley of Kashmir and some of the neighbouring
valleys, such as Maru-Wardwan, Kishtwar, Badrawar, and Tilel; west-
ward limits unknown. In summer hangul are found at elevations of from
gooo to 12,000 feet, but in winter descend to the level of the Walar
Lake, at about 5000 feet above the sea.

Habits.—After stating that by the end of March all the old stags and
most of the young ones have shed their antlers, Colonel R. J. Percy!
writes as follows concerning the migrations of the hangul :—‘‘' The deer
collect into large herds and begin moving off to their summer quarters,
those in the western corner of the valley (of Kashmir) going to the banks

' Badminton Library, Big Game Shooting, vol. ii. p. 277. Some emendations in the spelling of
place-names are made in the quotation.

Haneul 87

of the Kishanganga river (Tilel). The herds which strike the river at its
nearest point below Gurez cross it, and retire to the range of hills on the
southern border of Astor. Only a very few stags cross this range, the bulk
of the deer remaining on the Kashmir side. The deer on the northern
and eastern sides of the valley retire to the slopes of Haramuk and the
high ground south of the range which separates Kashmir from Dras and
Suru, but do not appear to cross it. The farther east one goes from
Srinagar the less the deer appear to migrate, merely retiring to the heads
of the valleys. The altitude of the birch copses just above the limit of the
pines is what they seek, and this they can find close at hand on the north
and east of the valley, but they have to travel some distance to it on the
west.” None are found on the Pir Panjal range, forming the southern
barrier of the Kashmir valley.

In June I have seen hangul close to the summit of the Zogi-la, the pass
between Kashmir and Dras, at an elevation of 11,000 feet ; but by the time
the antlers are free from velvet they descend to much lower levels. By
September the antlers of the stags are clean, and about the 2oth of that
month calling begins, although the date of its commencement depends
somewhat upon the season and the weather. Although the older writers
on Himalayan sport, like Colonel Markham, speak of hangul calling
throughout the day, at the present time they only do so in the mornings
and evenings, commencing in the late afternoon. During the calling-season
stags may frequently be seen escorting a party of hinds into the open glades
between the patches of forest, and it is then that the sportsman finds his
best opportunity. These deer seem by no means to confine themselves to
a single forest, but are much given to wander from one belt to another.
The calling-season generally ends by the latter part of October, after
which the stags are exceedingly difficult to find, until the snow drives
them down to the open country of the Kashmir valley. The fawns are
born in April.

Although the term Barasingha (12-tined), or occasionally Burrasingha
(big-horned), is applied to these deer by Kashmiri shikaris, they never use
it when conversing among themselves ; and it has been imported from the
plains of India, where it properly belongs to the swamp-deer.

88 Elaphine Group

6. YARKAND RacE—CERVUS CASHMIRIANUS YARCANDENSIS

Cervus cashmirianus yarkandensis, Blanford, Proc. Zool. Soc. 1892, p. 117.

Cervus yarkandensis, Blanford, op. cit. 1893, p. 447.

Cervus yarcandensis, Lydekker, ibid. 1896, p. 933.

Characters.—The antlers less spreading, with their terminal (fifth)
tines less inclined inwards, and therefore separated by a wider interval than
in the typical race. The pelage is stated to be generally pale-coloured,
with a well-marked caudal disk. Several skulls in the British Museum
show the normal five tines, but in one specimen presented by Mr. H.
Leonard an extra tine is developed on the crown of each side so as to form
an imperfect cup.

In his original description Mr. Blanford was careful to state that the
name yarkandensis was to be used in a sub-specific sense, and that this form
must not be regarded as specifically distinct from the Kashmir stag. In the
type specimen the two antlers are respectively 35 and 34 inches in length,
measured along the beam and round the curves, the basal circumfer-
ence above the burr being 632 inches.

Distribution Eastern Turkestan, in the woods on the Yarkand or
Tarim river.

I have not met with any account of the habits of this deer, which are
probably, however, very similar to those of the Kashmir race.

4. THE SHou——-CERVUS AFFINIS

(?) Cervus wallichi, Cuvier, Ossemens Fossiles, ed. 4, vol. vi. p. 89
(1835).

Cervus affinis, Hodgson, ‘fourn. As. Soc. Bengal, vol. x. p. 721 (1841) ;
Sclater, Trans. Zool. Soc. vol. vii. p. 343 (1871) ; Brooke, Proc. Zool. Soc.
1878, p. 913; Sterndale, Mamm. Ind. p. 514 (1884) ; W. L. Sclater, Caz.
Mamm. Ind. Mus. part ii. p. 184 (1891); Blanford, Fauna Brit. Ind.—
Mamm. p. 537 (1891) ; Ward, Records of Big Game, p. 41 (1896).

(?) Pseudocervus wallichu, Hodgson, ‘fourn. As. Soc. Bengal, vol. x.
p. 904 (1841) ; Gray, List Mamm. Brit. Mus. p. 180 (1843).

Characters.—Size probably equal to that of the wapiti. Antlers of the

Shou 8g

general type of those of the hangul, but larger, and with the beam bent
suddenly forwards at the trez (third) tine, so that its upper half overhangs
to a great extent the face ; number of points usually five, the brow-tine less
constantly longer than the bez, and separated by a shorter interval from the
burr, than is the case in the hangul. With regard to coloration, which
Mr. Blanford states resembles that of C. cashmirianus, with the caudal disk
well marked, more information is required. In a mounted head in the

Fic. 23.—Front view of Skull and Antlers of Shou. From a specimen in the British Museum.
(Rowland Ward, Records of Big Game.)

British Museum the colour is pale rufous brown ; and this is the colour of
the entire body, including the buttocks in the coloured figure of a stag in
Brian Hodgson’s sketches. A second sketch, representing a stag and hind,
in the same series, shows, however, the colour darker and a large white
caudal disk; this being probably the winter coat. In the aforesaid
mounted specimen the ear is not more than one-third the length of the
head, and is not distinctly white internally, but it appears longer in the
sketches, and its small size in the stuffed head may be due to shrinking.
N

go Elaphine Group

In one of the two sketches the tail is relatively long, whereas in the other
it is comparatively short.

The following dimensions of antlers of this species are taken from Mr.
Rowland Ward’s book; the first two in his list being omitted, as it is
doubtful whether they are rightly named.

oe h ae Se as + Tip to Tip. Greatest Width

uter Curve. Circumference. inside.
548 63 214 372
534 6g 265 44
534 6s 30 454
53 9 ? 40
52 8 P
492 72 38 455
48% 63 195 38%
478 58 20% 403
439 58 238 35

With the exception of three, all the specimens in this series have five
points on each side ; in the exceptions, one antler in each has five points,
while on the opposite side the number of points is respectively four, six,
and seven.

Distribution —Apparently the district lying to the north of Bhutan,
and probably the valley lying to the east of the Chumbi valley, and drain-
ing northwards into the valley of the Sangpo.

Information is urgently required as to the coloration and habits of this
magnificent deer, which has never been exhibited alive in any European
menagerie, if, indeed, it has been seen alive by European eyes. Brian
Hodgson had, however, either a skin or a specimen in the flesh, and it
is possible that a deer formerly living in the Viceregal menagerie at
Barrackpoor, near Calcutta, may have belonged to the species. With
regard to the scientific name adopted for this species, the words of Mr.
Blanford may be quoted, which are as follows :—“’The animal to the
figure of which, by Duvaucel, Cuvier gave the name of C. wallichu lived
in the Barrackpoor menagerie, and was said to have been brought from
Muktinath, near Mount Dwalagiri in Nepal. This place is as nearly as
possible half-way between the localities inhabited by C. cashmirianus and C.
affinis respectively. It is difficult to believe that any large deer living in

Tre DEER or Att Lanps Puarte V.

THOROLD’S DEER.

Published. try Rowland Ward [ta

Thorold’s Deer on

Northern Nepal could have escaped the knowledge of Hodgson’s collectors.
The shed horns of the type specimen of C. wa/lichi are preserved in
Calcutta, and have been figured ; they are probably, according to Mr. W. L.
Sclater, of the third year, but whether they agree better with those of
C. cashmirianus, or C. affinis, of the stag of Eastern Turkestan, or of any
other species, is undecided.”

There is, therefore, a great probability that what Cervus wallichii really
was will never be determined, although I have a suspicion that it was the
same as the present species. If this prove to be the case, the name affinis
will have to be superseded.

5. THoRoLp’s DrEER—CERVUS ALBIROSTRIS

Cervus albirostris, Prezewalski, Catalogue of Zoological Collections of H. M.
Prezewalski, p. 16, no. 36 (St. Petersburg, 1887).

Cervus dybowsku, W. L. Sclater, ‘fourn. As. Soc. Bengal, vol. lviu. p.
186 (1889), mec Taczanowski, 1876.

Cervus thoroldi, Blanford, Proc. Zool. Soc. 1893, p. 444.

(?) Cervus nariyanus, Hodgson, ‘fourn. As. Soc. Bengal, vol. xx. p. 292
(1851) ; see Blanford, of. cit. p. 447.

Plate V

Characters.—Size approximately the same as that of the hangul.
Antlers much flattened, with no bez (second) tine, and five or four points,
the brow-tine arising a considerable distance above the burr, the trez nearly
in the same plane with the tines above it, and the succeeding tine (third in
this species, but representing the fourth in those with the bez developed)
longer than either of the others; the beam suddenly bent back at the
origin of the trez-tine. Tail short, and the caudal disk of moderate size
and embracing the whole tail. Ear half as long as the head. Hair on
middle of back reversed, so as to be directed forwards towards the neck,
and the muzzle, chin, and lips white. General colour of pelage (season
unknown) uniformly dark brown, with the hairs minutely speckled, and
scarcely paler on the under-parts than above ; caudal disk pale ochry buff,
without any white below the tail ; inner surface of ears whitish, and, as
already said, pure white on the muzzle, lips, and chin. Metatarsal tuft
very coarse, and situated nearly in the middle of the cannon-bone.

Q2 Elaphine Group

The hairs on the body differ from those of other species of the genus in
their coarseness and pithy interior, thus somewhat resembling those of the
musk-deer. Possibly this structure is a provision against extreme cold.
Each hair is long, with the greater portion brown, but the base buff, and
the tip whitish, thus communicating the speckled appearance to the whole
pelage. The light tips are, however, wanting on the margin of the
caudal disk, thus forming, as in the hangul, the blackish border to this
area. The reversal of the hairs on the middle line of the back extends
from the hips to the withers, where it ceases abruptly in a kind of hump.

Compared with other members of the group, the skull is relatively
short ; and the muzzle is also exceptionally short and broad. In several
other structural details the skull also shows an approximation to the sikine
type; but its most remarkable peculiarity is the great expansion of the
nasal bones a short distance below their upper extremities, this being so
great as almost to obliterate the lachrymal vacuity, which is consequently
much narrower than in other members of the group. In this respect the
species approximates to some of the smaller members of the sambar group.

In regard to the affinity of this deer, Mr. Blanford remarks that it does
not come very close to any other species, although its relations are probably
on the whole nearer to C. cashmirianus and C. afinis than to any other.
He also remarks on certain wapiti-like characters in the antlers. But these
antlers—especially when they have only four points—appear to me in their
general flatness, and also in the absence of the bez-tine, to present a decided
resemblance to those of the sikas, towards which the species also approxi-
mates in the relative shortness of the skull. And it seems not improbable
that this form may indicate the line of descent of the wapiti from the
sikine group, while the hangul may mark the evolution of the red deer
from the same stock.

Distribution —The two specimens forming the types of C. thoro/d: were
shot by Dr. W. G. Thorold during his journey across Tibet at a spot about
200 miles to the north-east of Lhasa, at an approximate elevation of 13,500
feet above the sea-level, in the snow among brushwood just above the forest.
The head and antlers of a third specimen had been previously purchased by
Mr. W. L. Sclater in the bazaar at Darjiling, and provisionally assigned
by him to the sikine group, under the name of Cervus dybowsku. At a still
earlier date specimens had been obtained from some part of Central Asia

Thorold’s Deer O3

by Col. H. M. Prezewalski, and named C. a/birostris, and since this name
is accompanied by a brief description, in Russian, and also by photographs
clearly showing the characteristic white muzzle, it is entitled to supersede
the one given by Mr. Blanford.

With regard to the antler described by Hodgson under the name of C.
nariyanus, which Mr. Blanford considers may have belonged to a young
individual of the present species, that writer’s own words may be quoted.
“‘This horn,” he writes, “was said to have been brought from Ladak, it
was 34 inches in length, and had four points, the two lower being more
than 4 inches apart, so that there was no bez-tine. Judging by the figure,
the horn was more massive than would be expected in a young specimen
of Thorold’s stag. Mr. Hodgson remarked that the Bhotiahs who brought
this horn say it belonged to a very young animal, and that the species,
which is proper to Gnari, in Western Tibet, is larger than the shou. The
stag obtained by Dr. Thorold is considerably smaller than the shou ; there
is, so far as is known, no stag in Western Tibet, C. cashmirianus being
limited to the Kashmir valley, at all events to the north and east of its
range, and, as is well known, young examples of C. e/aphus, and I believe
of the wapiti also, frequently want the bez-tine ; so that it is by no means
impossible the Ladak horn may have belonged to a young C. cashmirianus
from Kashmir, to C. yarkandensis from Eastern Turkestan, or even to C.
eustephanus (C. canadensis, var.) from the Thian Shan, Ladak being connected
with all those regions by trade routes. ‘The Bhotiah story was probably
pure fiction.

“There is, moreover, one very strong reason for not using the name
nartyanus for the present animal, even if, as is highly probable, it was the
species that furnished the horn described and figured by Hodgson. ‘The
name was taken from Nari, the western (or rather perhaps the south-
western) province of Tibet, often called Gnari or Nari-Khorsum, a tract, as
represented on maps, of no great breadth from north to south, but extending
along the north of the Himalayas from the western extremity of Tibet
proper, near Rudok, and between long. 80° and 85° E. This region, part
of which is known as Hundes, is on the frontier of our own territory, and
has been visited at several points by British sportsmen. If any stag
inhabited the region, it is incredible that nothing should have been heard
of it ; moreover, the whole of the upper valleys of the Sutlej and Yarot-

04 Elaphine Group

sanpo, or Brahmaputra, of which the area consists, is a barren, treeless,
almost bushless waste, differing essentially from the country inhabited, so
far as is known, by any species of Cervus. I think it extremely improbable
that any stag inhabits Nari ; and under these circumstances it is not desir-
able to apply the name of zariyanus to a species which does not occur there.”

With this conclusion I fully agree, although I believe it would not be
accepted by all zoological writers. So far as English literature is concerned,
nothing appears to have been recorded regarding the habits of this very
distinct species of deer.

6. THe Wapiti—Cervus CANADENSIS

Cervus elaphus canadensis, Erxleben, Syst. Regn. Animal. p. 305 (1777).

Cervus wapiti, Barton, Philadelphia Med. and Phys. “fourn. vol. iii. part
i. suppl. p. 36 (1808) ; Leach, ‘fourn. Physique, vol. Ixxxv. p. 67 (1817).

Cervus major, Ord, in Guthrie’s Geography, p. 292 (1815).

Cervus canadensis, Desmarest, Mammalogie, vol. i. p. 433 (1822) ;
Baird, N. Amer. Mamm. p. 638 (1857); Sclater, Trans. Zool. Soc. vol. vii.
p- 342 (1871); Gray, Cat. Rumiants Brit. Mus. p. 68 (1872); Caton,
Antelope and Deer of America, p. 77 (1877); Brooke, Proc. Zool. Soc. 1878,
p- 913; Merriam, Mammals of Adirondacks, p. 143 (1884); Merrick,
Mammals of Minnesota, p. 278 (1892) ; Ward, Records of Big Game, p. 33
(1896) ; Rhoads, Proc. Ac. Philadelphia, 1897, p. 207.

Cervus (Elaphus) canadensis, H. Smith, in Grifhith’s Animal Kingdom,
vol. iv. p. 96, v. p. 308 (1827).

Cervus strongyloceros, Richardson, Fauna Bor. Amer, p. 251 (1829) ;
Gray, Knowsley Menagerie, plate xxxvi (1850).

Elaphus canadensis, De Kay, Zool. N. York, vol. i. p. 118 (1842).

Cervus (Strongyloceros) canadensis, Gray, Knowsley Menagerie, p. 58 (1850),
Cat. Ungulata Brit. Mus. p. 193 (1852).

Strongyloceros canadensis, Fitzinger, SB. Ak. Wien, vol. \xvii. part 1.
p. 350 (1873), lxix. part 1. p. 556 (1974);

Characters—Height reaching to 5 feet 4 inches at the shoulder.
Antlers very large, with more than five tines, curving backwards, and much
flattened in the upper half; the bez-tine present, the fourth tine longer
than all the others, and with the fifth, which is also long, forming a nearly

Wapiti 95

symmetrical fork, the fourth, fifth, and sixth being situated nearly in the
same plane as the portion of the beam immediately below them, so as more

\
: \
\

yy \
\

a

Fic. 24.—Head of East American Wapiti. From a specimen in the possession of Mr. W. Moncreiffe.
Length of antlers on outside curve 58, basal circumference 72, greatest width inside 49 inches,

or less nearly to occlude one another when viewed from the front ; the

brow-tine rising close to the brow, and approximately equal in length to

96 Elaphine Group

the bez ; the crown normally not cupped. Caudal disk larger than in any
other species; throat-fringe greatly developed ; neck and under-parts dark
coloured ; no white on the muzzle, and no reversal of the hair of the back.
Tail very short ; ear about half the length of the head ; face rather short.
General colour of summer coat on upper-parts yellowish brown, sometimes
with a reddish tinge ; neck and under-parts varying from dark brown to
blackish ; limbs generally chestnut-brown. Cry of male in the pairing-
season typically a loud squeal, ending in a more guttural tone.
Distribution —North America and Central and North-Eastern Asia.

a. East AMERICAN RacE—CERVUS CANADENSIS TYPICUS

Cervus canadensis verus, Blyth, Proc. Zool. Soc. 1865, p. 618.

Characters—The typical, or East American, race of this fine and distinct
species is a rather short-legged and heavily-built animal, standing about 5
feet 4 inches (16 hands) at the shoulder, and weighing from 700 to 1000 lbs.
The antlers are not excessively large in proportion to the general size, and
there is usually a distinct reddish tinge in the early summer pelage, although
later on in the season this fades to a dirty yellow tawny. The general
coloration has been admirably described by Mr. Caton, who observes that
the summer coat is of a dirty yellowish white colour on the body, and
chestnut-brown on the legs, neck, and head ; the males and females being
almost alike. When this coat first appears in June, upon the peeling off of
the old winter dress, it is of a deeper shade and more glossy than at any
other season of the year, and is the most beautiful dress the animal ever
wears. It is exceedingly short, fine, and soft, and fairly glistens in the
bright spring sunshine. In September this coat gives place to the winter
dress, but the change is so gradual as to require the closest scrutiny to detect
it, although the new coat differs very materially from the old in some parts.
When the winter dress has been fully assumed, a marked difference is
noticeable in the coloration of the males and females when the darker shades
prevail. In the former the neck, legs, and under-parts are brownish black,
and the dark border on the lower part of the light caudal patch deep black.
On the other hand, in the female, the head, neck, legs, and under-parts are
chestnut-brown, although the middle line of the latter is nearly black. The

caudal disk is straw-coloured.

Wapiti Q7

With regard to this light caudal disk, the same writer observes that it
“commences at the top of the hip and extends back so as to embrace the
tail ; its outer border descends laterally in a circular form, so that when even
with the root of the tail, above that member, it nearly describes a semi-
circle ; thence the outer border descends down the ham, gradually drawing
inwardly, and contracting the white section, which, however, descends to
unite with the lighter shades of the inguinal region. This white portion
is bordered by an intensely black stripe, which commences on either side

Fic. 25.—East American Wapiti. From a male at Woburn Abbey.
Photographed by the Duchess of Bedford.

above the region of the tail and continues down to the posterior sides of the
thighs, where it fades out and is lost. The black mark appears on all
animals of all ages and both sexes, but is the most brilliant on the male in
the prime of life and in the fall of the year.” When the summer coat is
first assumed, the light caudal disk is straw-coloured, but it gradually fades
till in the latter part of the summer it becomes nearly white. Fading also
takes place in the winter pelage, which gradually bleaches from the tawny
yellow of autumn to a pale dirty white in winter, although there is much
individual variation in this respect, scarcely any two animals in a herd being
of exactly the same tint.

98 Elaphine Group

Although from six to eight tines is the normal number in American
Wapiti antlers, not unfrequently additional tines are developed in the region
of the crown, which may then form a cup with more or less extensive
palmation. Such a palmated head is figured on page 414 of the first volume
on Big Game Shooting in the “ Badminton Library.”

The following dimensions of antlers of both races of American wapiti
are taken from Mr. Rowland Ward’s Records of Big Game, an abnormal
specimen heading the list being omitted :—

olangthalong — Greumerence rip to'Tip. Wile inside, Numi of oat.
65 73 : 48 7-7 Wyoming
644 8 : 48 7-7 ‘5
634 84 494 483 7-7 ;
62 78 334 503 7-7 Colorado
614 1% ? 45 6-7 Wyoming
604 1 ? Lo 6-6 Rocky Mountains
603 74 43 464 6-6 Wyoming
602 8h ? 55 7-6 "
598 ? ? ? ae e
593 83 374 47 8-7 Nebraska
593 7s i 45 6-6 Wyoming
584 1% 364 47 9-8 f
583 9 ; 465 = 10-7 i
583 84 ? 44% 6-6 3
57 6% 244 352 6-6 Montana
573% 7 47 482 6-6 Wyoming
574 98 32 424 Te/ 2
57 7§ 61 49 8-8

Distribution.—North America, east of the Rocky Mountains, from latitude
57 N. in the interior of the continent through Labrador, the Alleghany
region of Pennsylvania and Virginia, Northern Wisconsin, Minnesota,
Dakota, Nebraska, Wyoming, and Montana. From the Adirondack region
of New York the typical wapiti has long since been exterminated, the only
remaining evidence of its former presence there being buried antlers.
Regarding its range in the South-Eastern United States, Mr. Rhoads writes
as follows! :—‘* The former range of this animal in Pennsylvania was closely

1 To avoid confusion, the names wapiti and bison are substituted in the extract for elk and buffalo.

Wapiti 99

coextensive with that of the bison, both species using the same trails,
feeding-grounds, and licks among the Western Alleghanies, and passing
thence eastwards by the same routes to the Delaware valley. The wapiti
was most numerous among the elevated mountain glades and eastern
tributaries of the Alleghany and Monongahela rivers. It was also fairly
abundant in the early part of the century in Clinton, Potter, Tioga, and
Lycoming counties. The range of the wapiti and bison into the south
central counties of Pennsylvania, east of Fulton county, is very improbable,
if, indeed, they ever wandered so far. The main line of their eastern range
was probably along the valley of Castleman’s River in Somerset county and
the main ridge of the Alleghany Mountains near that place, which formed
a continuous trail of safety between their haunts in West Virginia and the
Keystone State. North of this region their range probably spread north-
eastward as far south as the Juniata valley, but by far the greatest number
did not come south of the east and west branches of the Susquehana. The
presence of an Elk Mills and Elk Creek in Chester county, and of an Elkton
in Cecil county, Maryland, would indicate their former presence in that
vicinity, probably only as stragglers along the Susquehana valley.”
Incessant persecution is telling on the numbers of both the American
races of the wapiti very severely, although it may be hoped the day is far
distant when they will meet the fate which has already befallen the bison.
East of the Mississippi there is said to be only a single locality where they
exist in the wild state; and although some are reported to remain in the
forests of Lower Michigan, their days are probably numbered. At present
to the westward of the Missouri, in the more remote districts of Montana,
Wyoming, and Colorado, as well as in the Rocky Mountains, there are
localities where the wapiti is still fairly abundant, yet its numbers are now
to be reckoned by hundreds, instead of by thousands, as in the olden days.
Habits.—By an unfortunate misapplication of terms, the wapiti is very
generally known in America as the elk; and the reader of American
sporting works must consequently be on his guard against confusing the
animal under consideration with the one to which the latter title pertains.
The accounts of the habits of the wapiti, both from the point of view
of the sportsman and of the naturalist, are numerous, although the majority
are based on the excellent description given by Mr. Caton, from which the
following observations are in great part condensed. As regards food, these

100 Elaphine Group

deer are very promiscuous, consuming alike grass, rank herbage of various
kinds, and the leaves of deciduous trees ; while in winter scarcely any kind of
vegetable matter comes amiss. Leaves and twigs seem, however, to afford
decidedly the most favourite nutriment ; and among the Woburn herd the
hinds, at least, can always be attracted to the fence of their paddock by
offering them leafy boughs. Gregarious at all times, the stags keep more
or less separate from the hinds at the season when the fawns are born,
which is during May and June. At this time even the hinds scatter, the
stags collecting in small parties by themselves ; the latter, when in mountain
districts, ascending to within a short distance of the snow-line. With the
commencement of August the scattered groups begin to collect, and as
September draws nigh, when the pairing-season is at its height, the old
stags gradually collect around them a party of hinds, from which the smaller
males are, so far as possible, driven away. ‘The latter, however, hover
about in the vicinity of the party, and seize every opportunity of drawing a
straggling hind away from her allegiance. During this season the old
stags are continually roaring; and the roar must be heard to be fully
realised, as no description will give any adequate idea of its peculiar sound.
Still it may be described with considerable approach to accuracy as a kind
of squeal, ending up with several hoarse, guttural grunts. During the
time that the old stag has his harem with him, he treats its various members
badly, always endeavouring to wrest from them the choicest food, and
enforcing his claim with his antlers. Although in a wild state wapiti are
stated to be prevented by their natural timidity from attacking any but the
members of their own kind, in confinement old stags are vicious during
the pairing-season, and rush at all persons approaching their enclosure.
After the pairing-season wapiti collect in large herds, which used
formerly to number several hundred individuals, and wander about for a
time till they finally select their winter feeding-grounds. These are usually
open hills where the ground is kept more or less free of snow by the wind,
so that such food as there is at this season may be obtained with the least
difficulty. During the hot weather, when they are much persecuted by
flies and mosquitoes, wapiti resort to water, in which they will stand for
hours ; and, in the pairing-season at least, the old stags are fond of wallowing
in mud-holes, from which they emerge coated with dirt, and presenting

anything but a prepossessing appearance. The antlers are shed in March,

West American Wapiti LOI

and the new pair free from velvet by the end of August or beginning of
September. Saplings of aspen or pine appear to afford the favourite
rubbing-posts for freeing the antlers from the last remnants of the velvet.

In the wild state the hind breeds when two or three years old ; the
number of fawns at a birth being sometimes two, or rarely three, although
one is the most common. The beautifully spotted fawns are very active,
but it is some little time after their birth that the hinds rejoin the herd.
In defence of her offspring the female wapiti displays great boldness and
energy ; her great aversion, when in captivity, being a dog, in the pursuit
of which the whole herd will join. Like that of the elk, the favourite
pace of the wapiti is a long slinging trot, which can be maintained for a
great length of time without apparent fatigue ; when forced into a gallop
the animal is soon tired.

Although the ferocity of the old stags during the pairing-season makes
them undesirable occupants of an open park, wapiti are easily domesticated,
and thrive well in the English climate. They cross readily with the Asiatic
wapitis, and also with the red deer ; and there is no doubt that a cross of
wapiti blood improves the stamina and size of the latter when there has
been a long course of in-and-in breeding. For magnificence of appearance,
no deer can compare with the various races of wapiti as inhabitants of a
park. In America Mr. Herrick states that he has seen wapiti employed
with success for draught purposes.

6. West AMERICAN RacE—CERVUS CANADENSIS OCCIDENTALIS

Cervus (Elaphus) occidentalis, H. Smith,’ in Grifhith’s Animal Kingdom,
VOlay..P..1Ol, Vv. p. 208),(1827).
Elaphus occidentalis, Swainson, Classif. Quadrupeds, p. 292 (1835).
Cervus occidentalis, Jardine, Naturalists Library—Mamm. vol. iii. p. 139
(1835).
Cervus canadensis occidentahs, Blyth, Proc. Zool. Soc. 1865, p. 618.
Strongyloceros occidentalis, Fitzinger, SB. Ak. Wien, vol. 1xix. part 1.
p- 563 (1874).
Cervus roosevelt, Merriam, Proc. Soc. Washington, vol. xi. p. 272 (1897).
' In the original description the tail is stated to be long, and there appears to be some confusion

with the mule-deer. But the antlers, which are clearly those of a wapiti, are first mentioned, and stand
as the type. Blyth’s acceptance in 1865 of the name confirms the-identification of the sub-species.

Io2 Elaphine Group

Characters.—Very close to the preceding, but longer-legged and more
lightly built, with the portion of the antlers above the fourth tine aborted,
and the colour very dark.

Specimens of this race are needed in England for comparison with the
typical one. Hamilton Smith, who is confirmed by Blyth, stated that
there were important differences in the antlers; and the former writer
refers to two pairs of antlers in the British Museum which are said to agree
with the specimens he saw in America, and to differ from those of the
eastern race by the bez-tine exceeding the brow in length. A male from
Oregon was living in the London Zoological Gardens in 1863, and, from
his recollection of this specimen, Mr. Thompson, the head-keeper, informs
me that it was much longer in the leg, slighter in the body, and altogether
a more elegant-looking animal than the eastern race. If this be so, it
approximates to the Manchurian race, which is just what would be expected
on distributional grounds. The antlers, according to Dr. Merriam’s
description, also approximate to those of the latter.

Distribution America, west of the Rocky Mountains, extending from
British Columbia and Vancouver Island through Washington, Oregon, and
Northern California to North-Western Mexico. From British Columbia it
is now apparently exterminated, although some remain in Vancouver Island.
Its numbers in Oregon are alluded to under the head of the preceding race.

c. MANCHURIAN RAcE—CERVUS CANADENSIS LUEHDORFI

Cervus lihdorf, Bolau, Abh. Ver. Hamburg, vol. vii. p. 33 (1880) ;
Fitzinger, SB. Ak. Wien, vol. xxxiii. p. 373 (1881).

Cervus luehdorft, Sclater, List Anim. Zool. Gardens, p. 263 (1883).

Cervus tsubra, Noack, Humboldt, vol. viii. p. 6, fig. 5 (1889).

Characters.—Apparently very closely allied to the West American wapiti,
although perhaps not attaining quite such large dimensions. Two hinds from
Manchuria living in the menagerie at Woburn Abbey in 1896-97 were
decidedly larger than those of the Altai wapiti, and also relatively more
long-legged animals, and this difference affords one of the reasons for
regarding the present form as sub-specifically distinct from the latter,
although in the annexed description the Manchurian wapiti is said to be
smaller than the (? typical) American race. ‘The summer pelage (as shown

Manchurian Wapiti 103
by the hinds at Woburn Abbey) is distinctly red, brighter even than in the

red deer, while the winter coat is tawny yellow, of a different tint from
that of the East American race. No trace of red is observable in the Altai
wapitl.

The type specimens of Cervus /uehdorfi, which comprised two pairs, were
obtained from Transbaikalia, and were probably brought from the Bureatish
Steppe of Northern Manchuria by nomads. The original description runs
as follows :—‘‘ The Isubra deer,” as it is called, “is intermediate in height

Fic. 26.—Manchurian Wapiti Hind at Woburn Abbey. Photographed by the Duchess of Bedford.

between the European red deer (C. e/aphus) and the North American wapiti
(C. canadensis). In size it is closer to the former, in the shape of the antlers
to the latter. Its hair is in winter brownish gray, in summer light brown ;
the throat has a small whitish median streak ; the under lip is whitish, with
three black spots, one small one in the middle, and two larger ones on each
side. The strong mane is like that of the wapiti—in colour dark chestnut-
brown, in places almost black ; in summer it disappears almost completely.
The eye is smaller than in the red deer. The tail is much shorter than in
either the red deer or the wapiti ; in the male it is only two-thirds of the

104 Elaphine Group

absolute length of that of the red deer ; relatively it is much shorter, as the
Isubra deer is larger. The caudal disk is very strongly marked, extending,
as in the wapiti, largely over the base of the tail. In the male it is foxy,
but in the other specimens bright straw-yellow. A dark band, extending
inferiorly on to the thigh, borders the disk. In both sexes the head is
elongated, and the nose somewhat convex. A pair of antlers sent from
East Siberia have two forwardly projecting brow-tines and a middle tine.
While the beam goes in a regular curve to end in a terminal tine, there is
given off a strong hind-tine, which is likewise single. Wherever tines are
given off, the antlers are flattened, and resemble those of C. eustephanus, as
figured by Blanford.” It is added that the young are spotted. Antlers
figured in the original paper show two well-developed tines above the
fourth tine, but in a pair belonging to the Duke of Bedford this portion is
aborted, as in the West American race.

Distribution —Manchuria and Amurland, that is to say, the forest districts
lying to the eastward of the Mongolian desert. If, as seems evident from
the two hinds at Woburn, this form comes nearer in build and stature to
the West American wapiti than does the Altai wapiti, it accords exactly
with what would have been expected from its geographical distribution.

The name Cervus isubra seems to have been given in error by Herr
Noack, as it is stated to be the title assigned to the species by Herr Bolau,
who is the founder of the name C. /uehdorfi.

Habits—The little that is known regarding the habits of this wapiti has
been reported by Herr Dorries, who is one of the few Europeans who have
explored its habitat. Its favourite resorts are the dense pine forests extending
from the Amur to the Lower. Ussuri. From these it never issues forth in
order to graze till about ten o’clock in the morning, and when abroad, it
is one of the most wary ofall the deer tribe. The pairing-season commences
about the end of September or the beginning of October, and lasts for about
a fortnight, during which time the old stags are roaring incessantly from
morning till night.

d. AtTAt RacE—CERVUS CANADENSIS ASIATICUS

Cervus maral (canadensis var.), Severtzoft, Turkestan “fevotme, pp. 62, 103

(1873), Ann. Mag. Nat. Hist. ser. 4, vol. xviii. p. 377 (1873).

Tae Derror Att Lanps. Plate Vir

ALTAI WAPITI.

ub

nd Ward

Altar Wapiti 1O5

Cervus maral, var. astatica, Severtzoft, Turkestan “fevotnie, p. 109 (1873),
Ann. Mag. Nat. Hist. t.c. p. 386.

Cervus eustephanus, Blanford, Proc. Zool. Soc. 1875, p. 637, Scient. Results
Second Yarkand Misston—Mamm. p. 90 (1879) ; Brooke, Proc. Zool. Soc. 1878,
Peouzen WWewleesclater,. Caz. Mamm. Ind. Mus. patt ii. p. 184 (1891) ;
Ward, Records of Big Game, p. 42 (1896).

Cervus canadensts eustephanus, Blanford, Proc. Zool. Soc. 1893, p. 447.

Plate VI

Characters.—Although generally like the East American wapiti, this
sub-species may readily be distinguished by its inferior stature, relatively
longer body and still shorter legs, and absolutely larger antlers ; being, in
respect to the development of the latter, decidedly the finest member of the
group. Indeed, in this point it may be compared to the barren-ground
reindeer, which, although a smaller animal than the woodland race, has
larger antlers. The colour of the pelage of the upper-parts is yellowish
tawny at all seasons, with no tinge of red.

Distribution —The Altai and Thian-Shan Mountains, which form the
forest districts to the west of the Mongolian desert.

History. —The name C. eustephanus was applied by Mr. Blanford to certain
shed antlers purchased by the members of the Second Yarkand Mission in
Kashgar, and stated to come from the forests of the Thian-Shan. A pair
of these antlers (the types) are preserved in the Indian Museum, Calcutta,
and there are three single specimens in the British Museum, two of which
belonged to adults, and the third to an immature animal. Of the Calcutta
specimens, Mr. Blanford writes that “they have apparently been shed,
and they probably belonged to different animals. They are of large size,
each measuring 51 inches along the curve ; one is 10.9, the other 10.5 inches
in circumference at the base, just above the burr. Each shows seven well-
formed tines, so that the animal must have had fourteen points. The beam
is very much curved, and, so far as it is possible to judge from the form of
the burr, the horns must bend somewhat towards each other at the tips and
branch less apart than in most stags. The brow-antler and bez are close
together, the former slightly exceeds the latter in length, and the bez
(second) is rather larger than the royal (third). The greatest peculiarity
of the horns, however, is in the form of the crown. Above the royal the

P

106 Elaphine Group

beam curves inwards and gives out an anterior (fourth) tine which is much
the largest of all, and slightly compressed, being only a little shorter, and
scarcely smaller, than the beam itself. Above this the beam gives out two
other tines, each successively diminishing in length, and all these four
branches, that is, the beam itself and the three upper tines, are in nearly

Fic. 27.—Altai Wapiti at Woburn Abbey. Photographed by the Duchess of Bedford.

the same plane, so that by looking at the horn with either the beam or the
great fourth tine in front, the remainder of the crown can be concealed
behind either one or the other. . . . It appears to me that, as regards the
horns, the Thian-Shan stag approaches the wapiti more than any Asiatic
deer. The resemblance between the Asiatic stags and Cervus canadensis
has been discussed by many naturalists, and by none more fully than by Mr.
Blyth, who has pointed out that the most important characters in which

Altai Wapiti 107

the horns of the American stag differ from those of the animals found in
Eastern Tibet, Kashmir, and Persia are the smoothness of the former, their
tendency to flattening or palmation of the crown, their great subdivision in
the coronal region, and the marked backward curvature and want of
convergence in the upper portion of the beam. Now in all these characters
the horns brought from Turkestan appear to be intermediate between those
of the other Asiatic stags and the wapiti. The horns of the Turkestan stag
differ from those of the wapiti in being less smooth, more curved inwards
towards the ends, and in having the brow- and bez-antler much nearer, but
they are much nearer to the wapiti horns than they are to those of C.
cashmurianus or C. affinis.

‘There can, I think, be very little doubt that Cervus eustephanus is the
animal described by Severtzoff and Prezewalski as inhabiting the forests of
the Thian-Shan and neighbouring ranges. It is a very large animal—as
indeed is evident from the dimensions of the horns—adults being, according
to Severtzoff, as much as 6 feet high at the shoulder. It is probably
known as maral by the Arian tribes of Central Asia, the word being Persian
for deer. The true C. mara/, however, inhabiting the forests on the
southern coasts of the Caspian, and in the Caucasus, etc., is a much smaller
animal, with differently shaped horns.”

The antlers in the British Museum were subsequently examined by Sir
Victor Brooke, who pronounced them indistinguishable from those of the
true wapiti, of which species Mr, Blanford finally came to the conclusion
that the present form is only a variety. All the characters he gives are
essentially those of wapiti antlers, of which they are an excellent descrip-
tion. That the deer described by Severtzoff are identical with C. eustephanus
may be considered beyond doubt, certain slight points of difference from the
type of the latter being of no importance ; and as the name asvaticus is the
earlier of the two, it is entitled to stand for the sub-species.

The first living examples of the Altai wapiti received in England were
a small herd from the Altai purchased in the autumn of 1896 by the Duke
of Bedford from Mr. Hagenbeck. These comprised three stags and several
hinds. On their arrival at Woburn Abbey the antlers of the stags were
so injured as to afford no distinctive characters, although they seemed to
indicate that the animals were immature. The new antlers developed in
the spring of 1897 were, however, precisely similar to the shed specimens

108 Elaphine Group

trom the Thian-Shan described as C. eustephanus, and served not only to
show the identity of the Altai and Thian-Shan deer, but also that the
Woburn specimens had by this time become fully mature, their antlers
carrying seven tines each, with the great development of the fourth tine
characteristic of the wapiti. The stags are, however, considerably smaller
than some full-grown American wapitis in an adjoining paddock at Woburn
Abbey, although their antlers appeared decidedly larger. And as a full-
grown American wapiti stands not more than 5 feet 4 inches (16 hands) at
the withers, the height of the Altai specimens may be roughly estimated at
5 feet, or 5 feet 2 inches. ‘This leads to the conclusion that the height
(6 feet) given by Severtzoff is excessive ; and it may be suggested that it is
either due to a misprint in his text, or that the height of the animal was
estimated from the antlers, which, as being larger than ordinary examples
of the true wapiti, would naturally lead to the conclusion that the animal
itself was superior in stature to the latter.

Habits.—Mr. St. George Littledale, who is one of the few Englishmen
that have seen it in its native haunts, tells me that this wapiti inhabits
thickly wooded districts in the Altai, and is very difficult of approach.
During the winter months large numbers are captured by the natives, who
drive them into nets. Extensive herds of such captive animals are kept in
domestication for the sake of their antlers, which are exported to China for
medicine, fine specimens realising as much as {10 each. During the
pairing-season the cry of the two old stags at Woburn Abbey was somewhat
intermediate between that of the red deer and that of the American wapiti,
commencing with a roar exactly like that of the former, but sometimes
concluding in a squeal resembling that of the latter.

7. THE BokHAaRA DEER—CERVUS sp. nov. (?)

Characters.—This deer, which appears to be an undescribed species, is
represented by an adult male now living in the Zoological Gardens at
Moscow, to which institution it was presented by the Governor of Russian
Turkestan. ‘The Duke of Bedford has a pair of the shed antlers. For the
following notes, as well as the photograph from which the accompanying
figure was taken, I am indebted to Mr. Carl Hagenbeck.

Height at shoulder, 4 feet. Antlers of the general type of those of the

Bokhara Deer 109

shou, showing a nearly similar forward curvature near the trez-tine, but
normally with only four points, owing to the absence of the bez-tine. Head
short and broad. ‘Tail about 4 inches in length. General colour of pelage
at all seasons ashy gray with a light yellowish sheen ; a darker line down
the middle of the back ; a small light-coloured caudal disk, including the
tail, and bordered inferiorly by a blackish streak ; legs brownish.

The antlers, which have been of the same general type for the last four

Fic. 28.—The Bokhara Deer. From the Moscow specimen.

years, are simpler than in any other member of the group, but come nearest
to those of the shou. The absence of the bez-tine appears to be constant ;
but in the shed pair in the possession of the Duke of Bedford there are
five tines on the left side, apparently due to abnormal development, as the
terminal fork is much smaller on this than on the opposite side. The
brow-tine, as in the shou, rises immediately above the burr. Probably
this deer indicates a small western representative of the last-named species.
Distribution —Russian Turkestan.

110 Sikine Group

ii. THE SIKINE Group—Sus-GeEnus PsEUDAXIS

Pseudaxts, Gray, Cat. Ruminants Brit. Mus. p. 70 (1872) ; Brooke, Proc.
Zool. Soc. 1878, p. 907, as a sub-genus.

Elaphoceros, Fitzinger, SB. Ak. Wien, vol. Ixviii. part 1. p. 602 (1873),
Ixix. part i. p. 596 (1874).

Fic. 29.—Antlers of Japanese Sika. From specimens presented to the British Museum
by Viscount Powerscourt.

Sika, Heude, Mém. hist. nat. emp. Chinots, vol. ii. p. 17 (1888).

Stkelaphus, Heude, op. cit. vol. 1. p. 146 (1894).

Characters.—Antlers smaller and simpler than is usually the case in the
preceding group, flattened, generally four- (occasionally five-) tined, with

:
«

i

.

fe
]
.

Tur DEER or Aut Lanps. Puate VII.

MANCHURIAN SIKA IN WINTER PELAGE.

Publiahed. yy Rowland. Ward) ltd

Common Sika REE

a trez (third) but no bez (second). Pelage of adult spotted with white,
at least in summer, and a pure white area bordered with black in the
region of the tail, which is also white and black ; young more or less
distinctly white-spotted ; metatarsal tuft generally whitish. Throat maned ;
head shorter than in the preceding group ; ears and tail moderate. Naked
portion of muzzle larger than in the preceding group, extending well on to
the upper surface of the face, and being very wide between the nostrils and
the upper lip. Rudimentary upper canines present ; upper molars of the
general type of those of the elaphine group. The face-gland, or tear-pit,
moderately developed. Size medium. In the growing antlers the velvet
is of a deep red colour, passing into black at the tips of the tines, and thus
matching the summer pelage. Sir Victor Brooke stated that the winter
coat is always without spots, and this error has been frequently repeated.

Distribution —Now confined to the south-eastern portion of the Eastern
Holarctic and some adjacent parts of the Oriental region, but apparently
represented in Europe during the Pliocene epoch.

In their spotted coloration and comparatively simple antlers, the sikas
are evidently a less specialised type than the elaphine group, and if, as
appears probable, they date from the Pliocene, they might well be the
ancestors both of the latter and of the fallow deer group. They have
frequently been considered as near relatives of the sambar group,' although
I cannot satisfy myself that this is well founded.

1. THE Common SIKA—CERVUS SICA

Cervus (Hippelaphus) japonicus, Sundevall, K. Svenska Vet. Ak. Hand. for
1844, p. 177 (1846), as commonly quoted ; no specific name was, however,
really given, the word japonicus (printed in ordinary type) merely signifying
that the new deer came from Japan, and not being its scientific title.

Cervus stka, Temminck and Schlegel, Fauna ‘fapon—Mamm. p. 54
(1847) ; Sclater, Proc. Zool. Soc. 1860, p. 377, Trans. Zool. Soc. vol. vii.
p- 346 (1871) ; Brooke, Proc. Zool. Soc. 1878, p. 908 ; Powerscourt, zbid.
1884, p. 208; W. L. Sclater, Cat. Mamm. Ind. Mus. part il. p. 39
(1891) ; Ward, Records of Big Game, p. 21 (1896).

Cervus syka, Pucheran, Arch. Mus. Paris, vol. vi. p. 398 (1852).

1 See Fitzinger, SB. Ak. Wien, vol. xix. part i. p. 603.

112 Sikine Group

Rusa japonica, Gray, Ann. Mag. Nat. Hist. ser. 3, vol. vi. p. 218 (1860),
Proc. Zool. Soc. 1861, p. 236; Sclater, Proc. Zool. Soc. 1860, p. 365, 1862,
p- 150—yavanica.

Pseudaxis stka, Gray, Cat. Ruminants Brit. Mus. p. 72 (1872), Hand-list
Ruminants Brit. Mus. p. 142 (1873).

Elaphoceros stka, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 352
(1873), lxix. part 1. p. 602 (1874).

Cervus euépis, Swinhoe, Proc. Zool. Soc. 1874, p. 151; Brooke, sid.
1878, p. go8.

Axis stka, Ritimeyer, Adbh. schweiz. pal. Ges. vol. viii. p. 93 (1881).

Characters.—Size very variable. Pelage bright rufous chestnut, spotted
on the body with white in summer ; uniformly coloured, or nearly so, in
winter, when it is dark umber-brown, with the hairs annulated ; a light
chestnut patch on the shoulder ; white on the side of the upper, and the
whole of the lower lip ; tail mainly white, frequently with a narrow black
line on the upper surface and sometimes a dark terminal tuft; caudal
patch large. Inside of ears and part of base of outer surface of same
covered with pure white hairs.

Distribution.—Northern China, Manchuria, and Japan.

a. JAPANESE RAcCE—CERVUS SICA TYPICUS

Characters.—Size small, the height of the typical form varying from
2 feet 8 inches to 2 feet 10 inches ; white area of caudal region very large,
extending on to the sides of the buttocks, and completely bordered with
black above and at the sides.

The typical Japanese sika is a beautifully coloured little deer, with a
bright chestnut red coat profusely spotted on the body with longitudinal
rows of white spots in summer, but turning to a uniform blackish brown
with the assumption of the winter dress. Generally all traces of the white
spots completely disappear at the latter season, although occasionally more
or less distinct vestiges of these may persist, more especially in hinds.
When in the velvet, the antlers are of a bright chestnut red, with black tips,
and at this season the bucks look their handsomest. The head and neck are
uniformly coloured at all seasons, the back of the neck in the summer pelage
being of the same tint as the ground-colour of the body, while the head and
throat are more fawn-coloured. The black-bordered white area in the

Japanese Sika ra

region of the tail stands out in bold contrast to the general colour at all
seasons. ‘The metatarsal tuft is very distinct.

Originally this deer was supposed to be confined to Japan, but Mr.
Swinhoe described, under the name of C. ewdpis,a specimen from China, now
in the British Museum, which, as Sir Victor Brooke states, can in no wise
be distinguished from the typical island form. The first examples of this
deer brought to England were received at the Zoological Society’s Gardens in
1860, but later on other Japanese examples arrived which were of larger
size, and thus served to connect the typical form with the larger Manchurian
race referred to below.- This led Sir Victor Brooke’ to express the opinion
that when a larger series of these deer were acquired, it would be found
impossible to separate them into definite species, but that they would
constitute local races of a single very variable specific type ; and it is very
difficult to believe that this is not the correct view, although in certain
points the typical manchuricus differs very markedly.

A large herd of sikas maintained in the park at Woburn Abbey exhibit
considerable variation in size, and tend to show that eventually a series may
be obtained graduating in height from the 2 feet 8 inches of the typical
Japanese race to the 3 feet 3 or 5 inches of the Manchurian race.

From the following list, which is taken from Mr. Rowland Ward’s
book, it would appear that from eight to ten points are occasionally
developed in the antlers of the species, although I have never seen specimens
with more than five on each side.

ee tear, wan, Ne Lat
314 St 278 278 4-4 :
258 + 20} : i ?
224 34 164 lox 8-9 Japan
29 4 163 132 5-4 Bred in Ireland
212 3 165 204 10-6 Yezzo
184 33 P 12 9 Bred in Ireland
183 L 11 12+ 4-4 Japan
164 38 : 13 4-3 3
162 g 92 10 4-3 Bred in Ireland
154 2$ oF 104 4-4 Bred in England
1st 32 124 lod 4-4 Bred in Ireland

1 Proc. Zool. Soc. 1878, p. 909.
Q

114 Sikine Group

Distribution.—Japan and Northern China.

Habits.—N othing of importance has been recorded of the habits of these
pretty little deer in their native country. In English and Irish parks they
thrive exceedingly well, and have much the same general habits as red deer.
The cry of the bucks in the pairing-season is a kind of whistle, sometimes
passing into a scream. In Lord Powerscourt’s park male sikas have crossed
with red deer hinds. In view of Sir Victor Brooke’s statement as to the true
red deer refusing to interbreed with the Caspian race of the same species,
this is a most remarkable circumstance. ‘Two pairs of antlers belonging
to the hybrid race are preserved in the British Museum, both of which
possess bez-tines. In the park at Woburn Abbey, where these deer run
with the fallow and red deer, they feed chiefly by grazing, although they
eagerly take boughs when offered. When startled and running off,
these and the Manchurian sikas expand and dilate the long white
hairs of the caudal disk, thereby increasing the distinctness of this con-

spicuous recognition mark.

4. MANCHURIAN RAcE—CERVUS SICA MANCHURICUS

Cervus mantchuricus, Swinhoe, Proc. Zool. Soc. 1864, p. 169, 1865, p. 1 ;
Sclater, Trans. Zool. Soc. vol. vii. p. 344 (1871) ; Brooke, Proc. Zool. Soc.
1878, p. 908; Moellendorff, Zool. Fahrb. 1887, p. 588 ; Ward, Records of
Big Game, p. 21 (1896).

Pseudaxis mantchurica, Gray, Cat. Ruminants Brit. Mus. p. 72 (1872),
Hand-list Ruminants Brit. Mus. p. 141 (1873).

Cervus kopschi, Swinhoe, Proc. Zool. Soc. 1873, p. 5743; Brooke, zbzd.
1878, p. gog.

Elaphoceros mantchuricus, Fitzinger, SB. Ak. Wien, vol. xix. part i.
p- 597 (1874).

Axis mantschuricus, Riitimeyer, Abh. schweiz. pal. Ges. vol. vill. p. 93
(1881).

Cervus sica manchuricus, Lydekker, Proc. Zool. Soc. 1897, p. 39.

Plate VII
Characters.—Size larger than the last, the height at the shoulder ranging
up to 3 feet 3 inches. The white area in the region of the tail much
smaller, so as scarcely to be apparent in a side view, but completely bordered

Manchurian Sika ris

with black. Spots and a tinge of red frequently retained on the hind-quarters
of does in winter pelage.

The type specimen of this form was a living buck procured by Mr.
Swinhoe at Ying-tzu-kou (Nuchwang), the treaty port of Manchuria, and
forwarded to the London Zoological Gardens, where it arrived in July 1864.
It is figured by Mr. Sclater, in plate xxxi of vol. vii. of the Society’s
Transactions, in summer pelage, and in plate xxxii in its winter dress.
The summer coat is well spotted, whereas the winter dress is a rich uniform

Fic. 30.—Buck and Doe of Manchunan Sika in winter pelage. From a photograph
by the Duchess of Bedford.

umber-brown, with light chestnut patches on the shoulders, faint traces of
the spots showing on the fore part of the back, and the under-parts being
dark. In the summer coat there is a black patch above the white area on
the tail, and a black line down the middle of the upper surface of the latter ;
but in the winter pelage very little black remains.

The height of the type specimen is given as 3 feet 8 inches, but I am
inclined to think this is a misprint for 3 feet 3 inches, or 5 inches, as a
stuffed buck in summer pelage at Woburn Abbey agrees in all respects
with the type, but only measures about 3 feet 2 inches at the shoulder ;

116 Sikine Group

and although evidently not fully mature, I cannot think it would ever have
grown six inches more. It shows a distinct light metatarsal tuft, which is
not apparent in the figure of the type, but may have been accidentally
omitted. Except for the reduced size of the white caudal patch, this
animal is in all respects essentially a large variety of the Japanese sika.
The type of C. kopschi, from China, is in the British Museum, and appears
rightly identified by Sir Victor Brooke with this race.

Distribution —South-Western Manchuria, or Shingking, typically from
the neighbourhood of Ying-tzu-kou (Nuchwang), but probably occurring
in other parts of Northern China. In habits this form is doubtless iden-
tical with the last.

2. THe FormMosAN SIKA—CERVUS TAEVANUS

Cervus tatouanus, Blyth, ‘fourn. As. Soc. Bengal, vol. xxix. p. go (1860) ;
Sclater, Proc. Zool. Soc. 1860, p. 376.

Cervus taévanus, Sclater, Proc. Zool. Soc. 1862, p. 152, Trans. Zool. Soc.
vol. vii. p. 345 (1871) ; Swinhoe, Proc. Zool. Soc. 1862, p. 362; Brooke,
ibid. 1878, p. gog; W. L. Sclater, Cat. Mamm. Ind. Mus. part i. p. 45
(1891) ; Ward, Records of Big Game, p. 22 (1896) ; Lydekker, Proc. Zoo/.
Soc. 1897, p. 45.

Pseudaxts tatvanus, Gray, Cat. Ruminants Brit. Mus. p.'70 (1872), Hand-
list Ruminants Brit. Mus. p. 141 (1873).

Elaphoceros taévanus, Fitzinger, SB. Ak. Wien, vol. \xix. part 1. p. 599
(1874).

Axis taivanus, Riitimeyer, Abh. schwerz. pal. Ges. vol. viii. p. 93 (1881).

Plate VIII

Characters.—Nearly allied to the common sika, but distinctly spotted
in winter. Height medium, apparently about 2 feet 11 inches at the
shoulder. The face shorter, the muzzle more pointed, the limbs shorter,
and the body longer than in the common sika. Ground-colour of summer
pelage light chestnut, with large white spots, and a deep red tinge on the
hinder part of the neck ; in winter the spots less numerous ; the black
border to the white caudal area forming a more distinct bar superiorly,
the median black line on the tail broader, and the dark line down the back

very strongly marked.

Tue Deer or Aut La

sf

%

FORMOSAN SIKA IN WINTER PELAGE.

Published. by Rowland Ward Ltd.

Nay ek

INiIJ OD .

pal ee ied vy" Oe

>

Tae DEER or Att Lanps.Puatre IX.

PEKIN SIKA IN WINTER PELAGE.

Pablithed. by Rowland, Ward Ld.

Pekin Sika kI7

Such are the characteristics to be drawn from the descriptions of
Messrs. Sclater and Swinhoe, supplemented by the specimens at Woburn
Abbey. The first example of this deer seen in England was sent by Mr.
Swinhoe, and arrived at the Zoological Gardens in December 1861.
Between that date and 1867 six other specimens were received, and in the
following year a fawn was born. Up to 1896 the only other specimen
was a male presented in the summer of 1889. I have had no opportunity of
studying living specimens which could be confidently referred to this form,
with the exception of two living at Woburn Abbey in 1896-97, one of
which is figured in plate viii. The shortness of the face is a common
feature in many island deer.

Distribution.—Formosa. There is nothing to lead to the belief that
this deer has been introduced into the island.

3. THe Pekin StkA—CERVUS HORTULORUM

Cervus pseudaxis, Gray, Proc. Zool. Soc. 1861, p. 236, nec Eydoux and
Souleyet.

Cervus hortulorum, Swinhoe, Proc. Zool. Soc. 1864, p. 169 ; Lydekker,
ibid. 1897, p. 42.

Cervus dybowskii, Taczanowski, Proc. Zool. Soc. 1876, p. 123; Brooke,
ibid. 1878, p. 909; Noack, Humboldt, vol. vii. p. 4, fig. 1 (1889); Lydekker,
Proc. Zool. Soc. 1897, p. 40; nec W. L. Sclater, fourn. As. Soc. Bengal,
vol. lviii. p. 186 (1889), Cat. Mamm. Ind. Mus. part ii. p. 182 (1891).

Cervus dybowski, Ward, Records of Big Game, p. 22 (1896).

Plate IX

Characters.—Size large, intermediate between that of a chital and a red
deer, the height at the shoulder being about 3 feet 7 inches (113.5 centi-
metres). Antlers very large and rugged, less flattened than in the common
sika, with the brow-tine given off from the beam some distance above the
burr. Summer pelage with large white spots ; winter coat of adult very
long and shaggy, more or less uniformly coloured, except on the flanks and
lower part of the thighs, where it is light fawn, with traces of spots on
the hind-quarters ; little or no pure white on the upper lip, the middle of
which is chestnut ; the white caudal area in adults small and inconspicuous,

118 Sikine Group

not extending on to the outer surface of the buttocks ; tail bushy, white
with a black tip.

The coloration of the winter pelage is described by Professor Noack as
follows :—General colour yellowish umber-brown, tending more to yellow
in front and to umber behind, and becoming darker on the back ; head as
far as the nose yellowish brown, forehead and neck reddish brown, nose
grayish red, upper lip yellowish red, the dark
spot on the grayish white lower lip moder-
ately large; ear thickly haired, dirty gray
internally, rusty red externally. Mane on
head and neck very long and shaggy, whitish
gray in colour; chest nearly black ; under-
parts whitish gray ; the white caudal patch
bordered in front with black; tail white
with a black tip. Metacarpus yellowish red,
metatarsus umber-brown, each with a dark
streak in front ; no light metatarsal tuft.

Two hinds in transition dress are described
as bright roe-colour; the individual hairs
being umber mingled with blackish brown ;
the forehead darker than in the winter coat,
and the whitish base of the ear very con-
spicuous.

In immature animals (plate ix) the

pelage is spotted with white at all seasons,
Fic, 31.—Head of Pekin Sika. From although more fully in summer than an
iinet Toate Wine, when tt 1 77 Jia eg
che Duchess of Bette ’ Bucks in winter with the ground-colour of
the body bright chestnut-brown ; neck with-
out spots, bluish gray at the base, then a blackish collar, followed by
chestnut ; face bluish gray ; thighs and fore-legs grayish brown; under-
parts grayish white; and often a dark band above the white on the
buttocks. In summer the general body-colour chocolate-brown.
The plate represents an immature stag and two hinds living in the
park at Woburn Abbey in early winter pelage. Later on in the winter
the coat becomes more shaggy, and the under-parts a deep leaden gray.

Pekin Sika 119

In their second winter season the spotting was much less conspicuous, less
white was apparent on the buttocks, and the whole colour was browner.

The history of this species is somewhat remarkable. In the spring of
1861 the Zoological Society received from the late Mr. R. Swinhoe the
skins of three sikine deer which had been shot after the taking of the
Summer Palace, Pekin (12th October 1860), when they would have
assumed the winter pelage. These specimens were shortly after transferred
to the British Museum, and one, a buck (No. 61. 6. 2. 1), was described
and figured by Dr. Gray in the Proceedings of the Zoological Society for 1861,
under the name of Cervus pseudaxis, with the express statement that it was
killed in autumn. As now mounted it stands 3 feet 4 inches at the shoulder.
The hair is now much faded. In Dr. Gray’s figuré! the hair of the body
is a chestnut-brown colour, with numerous distinct white spots of con-
siderable size ; and there is a white glandular patch on the outer side of
the hind leg just below the hock. The neck is unspotted, and its lower
portion is of a slaty-blue colour, above which there is a dark collar,
followed by chestnut-brown, the lower part of the face being also bluish
gray. The under-parts are whitish ; and the tail is white with a narrow
black median line and tip, but there is no distinct black cross on the
buttocks. This deer, which is about two years old, is therefore quite
unlike C. sica in its winter coat.

The female (No. 61. 6. 2. 2) appears to be similarly coloured, with
the exception that there is no slaty-blue on the neck, and the under-parts
are grayish, while there is a distinct black cross on the buttocks. Its
height at the shoulder is 2 feet g inches.

In 1864 Mr. Swinhoe, being satisfied that the buck figured by Dr.
Gray was not the Cervus pseudaxis of Eydoux and Souleyet, proposed for
it the name of C. hortulorum. In the same letter the name C. mantchuricus
was suggested for certain specimens sent home alive at the same time for the
Zoological Gardens ; and it is quite evident that Swinhoe was satisfied of
the distinctness of the two forms. The young stag represented in the
plate differs from the type specimen mainly by the larger amount of black
and white on the buttocks, but as the Woburn hinds are precisely similar

1 From a comparison with the original specimen, I find that the colouring of the figure is inexact in
several particulars, the under-parts being too white and the distribution of the colours on the neck very

badly shown.

120 Sikine Group

to the original hind sent home by Mr. Swinhoe, no doubt can be
entertained as to the specific identity of all the specimens.

Dybowski’s deer was originally described by Professor Taczanowski from
several examples obtained from the Ussuri district of North-Eastern Man-
churia, one of which (a male) was presented to the British Museum, where
it is preserved. Although not fully mature (the antlers being small), it
measures 3 feet 5 inches at the shoulder. It is in winter dress, and has
somewhat the general appearance of a very large Manchurian sika, although
with a much smaller white caudal patch, no light metatarsal tuft, and the
general colour somewhat lighter, although this last difference may be due
to fading. If Professor Taczanowski’s description be referred to, it will be
found that the Ussuri deer is compared with Cervus axis and C. dama,
without any reference to the sikas. One specimen in the original
description is stated to be the colour of a roe in winter pelage, with faint
traces of dappling on the hind-quarters, but another was lighter ; and the
Museum example shows little traces of spots. A more mature head (Fig. 31)
is in the possession of the Duke of Bedford. Sir Victor Brooke considered
that this deer might prove identical with C. manchuricus, and this view
was adopted by Herr Moellendorff, and subsequently by myself. I had,
however, at that time not seen the paper by Herr Noack, who regards
the North Manchurian sika as a perfectly distinct species. And at
that time the specimens at Woburn, figured in plate ix, were not
nearly so like the types of C. dybowsku as they have since become.
The present form seems to be a large, shaggy-haired deer, adapted for
living in a cold climate.

Distribution—The district of the Ussuri river, in North-Eastern
Manchuria, but recorded by Herr Noack from the south-west of
Vladivostock, near the borders of Corea. The distributional area is,
therefore, probably more northerly and easterly than that of the Man-
churian sika.

Habits—From information furnished by Herr Dorries, a few notes on
the habits of this deer have been published by Herr Noack in Der Weid-
mann (the German Fre/d) for 14th August 1891.

From these it appears that the Pekin sika is found in thin forest near
the coast, especially on the island of Ascold. The pairing-season com-
mences at the end of September, during which the old stags will call

Mandarin Sika I21

without ceasing an hour at a time. Most of the specimens captured are
taken after deep falls of snow ; and the antlers, like those of the Manchurian
wapiti, find a ready sale in China for medicinal purposes.

4. THE MaAnparin Sica—CERVUS MANDARINUS

Cervus mandarinus, Milne-Edwards, Rech. Mamm. p. 184 (1871) ;
Brooke, Proc. Zool. Soc. 1878, p. 968 ; Lydekker, sbid. 1897, p. 44.

(?) Cervus axis, Swinhoe, Proc. Zool. Soc. 1864, p. 169, nec Erxleben.

(?) Cervus mantschuricus major, Noack, Humboldt, vol. viii. p. 5, fig. 4
(1889). .

Characters.—This doubtful species is definitely known by the type
specimen preserved in the Museum at Paris, and from the figure of which
the following characters are taken. Size very large. Pelage spotted at all
seasons, and apparently very long and shaggy in winter. Colour redder
than in the preceding species, with the spots abundant in the winter pelage,
when the neck and limbs are similar in tint to the ground-colour of the
body ; under-parts dark ; metatarsal tuft apparently similar in colour to the
rest of the leg ; tail comparatively long, mainly reddish, with but little
white. In his original description Professor Milne-Edwards remarks that this
deer is distinguished from C. manchuricus by the more profuse spotting of
the summer coat, and the retention of a large, although somewhat smaller,
number of distinct spots in the winter dress on the body—the ground-
colour of the latter being dark chestnut-brown, and the neck and under-parts
also brown ; while there is a very thick frill of long hairs on the throat.
It is also mentioned that the colour of the summer coat is much brighter
than in C. manchuricus. These observations are fully borne out by the two
plates accompanying the memoir, which illustrate the type specimen at both
seasons. Sir Victor Brooke gives mandarinus as a synonym of manchuricus,
without a word as to the persistence of the spotting in winter. And the
influence of one so well known as an authority on the group has led to
C. mandarinus being ignored by subsequent writers.

In a letter published in the Proc. Zool. Soc. for 1868 Mr. Swinhoe wrote
as follows :—‘In the gardens of Messrs. Jardine, Matheson, and Co., in
Hong-Kong, I saw several bucks and does of C. sica and C. taévanus, as also
of C. axis in winter dress. The bucks of the two former had manes about

R

122 Sikine Group

the neck ; C. sca was spotless, C. ¢aévanus with indistinct spots, while C.
axis Was of a rich yellowish-brown colour, with distinct white spots. The
latter had long, thin, reddish tails, and, I think, are identical with the true
C. axis. They are from Hankow, interior China.” Now as Mr. Swinhoe
must be presumed to have been well acquainted with both C. manchuricus
and C. hortulorum, and as C. axis is unknown beyond India, it is possible
that these deer were really C. mandarinus. The “long, thin, reddish tails”
appear to accord well with Professor Milne-Edward’s figure of the latter
in winter dress.

Distribution. — Northern China and (?) Manchuria; whether the
specimens mentioned by Mr. Swinhoe really came from Hangchau may
perhaps be doubtful.

So far as the available material admits of forming an opinion, there
seems considerable probability that the stag from Manchuria described and
figured by Professor Noack under the name of C. mantschuricus major may
prove to be identical with the present form. It is stated to be as large as a
red deer, standing 4 feet at the shoulder, and is apparently very rare. It
was identified by its describer with C. hortulorum (C. pseudaxis, Gray).
In any case, the name C. major cannot be used for it on account of pre-
occupation by C. major, Ord, and also by Axis major, Hodgson. Possibly

it is the same as the Pekin sika.

5. THe PLiocene StkKA—CERVUS PERRIERI (Extinct)

Cervus perriert, Croizet and Jobert, Oss. Foss. Puy-de-Déme—Cervida,
plate iv (1828) ; Dawkins, Quart. “fourn. Geol. Soc. vol. xxxiv. p. 407 (1878) ;
Deperet, Bull. Soc. géol. France, ser. 3, vol. xii. p. 268 (1884) ; Lydekker,
Cat. Foss. Mamm. Brit. Mus. part ii. p. 107 (1885).

Cervus etuertarum, Croizet and Jobert, op. cit. plate vi (1828) ; Dawkins,
op. cit. p. 410 (1878) ; Depéeret, op. cit. p. 265 (1884) ; Lydekker, op. cit.
p. 107 (1885).

Cervus pardinensts, Croizet and Jobert, of. c7t. plate xi (1828) ; Dawkins,
op. cit. p. 409 (1878) ; Deperet, of. cit. p. 262 (1884).

Cervus tsstodorensis, Pomel, Cat. méthodique, p. 105 (1853) ; Dawkins,
op. cit. p. 407 (1878) ; Depéret, of. cit. p. 263 (1884) ; Lydekker, op. cit.
p. 107 (1885).

Pliocene Sika [23

Cervus rusoides, Pomel, op. cit. p. 106 (1853).

Characters—This species is known merely by detached antlers, which
are typically four-tined, and agree closely with those of the existing members
of the present group. Cervus etueriarum and C. pardinensis, which are
doubtless identical, were founded upon three-tined antlers from the same
deposits. These have very generally been regarded as indicating rusine
deer, although Professor Boyd Dawkins has already pointed out that the
typical three-tined antler of C. pardinensis might well have belonged to an
immature individual of C. perrieri—the four-tined form. In this view I
thoroughly concur, the three-tined antlers being decidedly more like those
of immature sikas than those of adult members of the sambar group.
Moreover, both types occur in the same deposits, which is of itself strong
prima facie evidence of their specific identity. Since, with the exception
of those of Anoglochis ardeus, antlers of a more complex type are unknown
from these beds, it would seem that four is the maximum number of tines
developed, and hence that the species belongs to the sikine group—a matter
of some importance from an evolutionary point of view. The brow-tine is
situated close to the burr, and branches off from the beam at an acute angle.

By Professor Depéret the antlers described as C. pardinensis are referred
to the rusine group, while those to which the other names belong are all
regarded as. indicating primitive members of the elaphine group, the
latter opinion harmonising to a great extent with that followed here.

Distribution —Europe (France and Italy) during the latter portion of the
Pliocene period.

INcERT# SEDIS
1. Cervus pseudaxis

Cervus pseudaxis, Eydoux and Souleyet, Voyage de‘ La Bonite, Zoology, vol. 1.
p- 64 (1841-52); Brooke, Proc. Zool. Soc. 1878, p. 909 ; Lydekker, zd. 1897, p. 38.

Axis pseudaxis, Gray, Cat. Ungulata Brit. Mus. p. 214 (1852); Fitzinger, SB.
An ten, vol. \xix. patti. p. 274 (1874).

Sikelaphus pseudaxis, Heude, Mém. hist. nat. emp. Chinois, vol. i. p. 146
(1894).

With regard to this nominal species, I can only follow Sir Victor Brooke, who ob-
serves that he has hesitated to identify it with “any species of the sub-genus. The type
specimen is still preserved in the Muséum d’Histoire Naturelle at Paris; but though

124 Sikine Group

I have often carefully examined it, the absence of the skull, and the great uncertainty
of the locality where it was procured, render it impossible to form a decided opinion.”
Mr. Sclater has suggested that it really belongs to C. ¢aévanus, in which case that
name would have to be superseded ; psewdaxis being the earliest of all.

‘The animal which has been figured under the name of Cervus pseudaxis,” writes
Dr. Gray, “was obtained by MM. Eydoux and Souleyet in Java, but they did not
believe that it was a native of that country. It lived several years in the Jardin des
Plantes at Paris, and hence a series of its horns was procured and figured ; and while
there it bred with the common axis, and the mule produce was fertile. Some
naturalists have given the Sooloo [Sulu] Islands, near the Philippines, as the habitat
of this specimen, but I do not know on what authority.” Mr. A. H. Everett
informs me that there are deer in the Sulu Islands, doubtless introduced. From his
description they appear to be sikas, and if so, they are not improbably C. saévanus.
Pere Heude has assigned a name to these Sulu deer.

2. Other Names

The following names have been applied by Pere Heude (Mem. hist. nat. emp.
Chinois) to specimens from China, Japan, and the Sulu Islands belonging to the present

group, Viz. :—
1. Stkelaphus soloensis, vol. i. p. 147 (1894).
2. Sitka poouvrelianus, 3 Ps Po. es.
3. 5, brachyrhinus, 5 Dea lant ‘
4. 4, a@ndreanus, 55). Pe LS =
5. 4, grilloanus, a ee ae bate
6. ,, dugenneanus, Sy ps UGE aa
Fo. aun JOLesanUs, 5 Pe Sy Ss
8. ,, oxycephalus, 3 a Ds cu ease
9. ,, frinianus, Spe TSO h ates
10. ,, cycloceros, oP lOGe
Li. 5 saraescens, Ay eemauleyi f
12. ,, /acrymans, je. Prwo2 i:
13. 4, @7terinus, ap woe A
14. 4, yuanus, Dh aon me
15. ,, sendaensis, vol. ii. p. 98 (1896).
16. ,, blakistonius, pe pes a
17. ,, dolichorhinus, io ip POOy aay
18. ,, aplodonticus, io Pe TeOaaas,
19: 4, schizodonticus,. ,5 p.. 108 Ws
20. ,, orthopodicus, 3 SDieloC, mas

Damine Group 126

21. Sika mitratus, vol. il. p. 102 (1896).
D2 ma MA ClUEpiECUs, Fe eyoraaileys, ss
DR Wy) CELANS, weep: 1O38 .
24. 4, Mminoensts, oP alOa .*5,
Dea Wye BCHUIES, Epos *
26. 5, yesoensts, Sie PeelOS . = ;;

iil. THE Damine Group—Sus-Genus DAMA

Alce, Blumenbach, Bevtrdge Naturgeschichte, 1st French ed. vol. ti. p. 407
(1803).

Dama, H. Smith, in Griffith’s Ammal Kingdom, vol. v. p. 306 (1827) ;
Gray, List Mamm. Brit. Mus. p. 181 (1843); Fitzinger, SB. 4k. Wren,
WOlebavitt. parti p. 249 (1073), Ixix. part 1. p. 546 (1874) ; Brooke, Proc.
Zool. Soc. 1878, p. 913; Rutimeyer, AdA. schwerz. pal. Ges. vol. vill. p. 43
(1881).

Megaceros, Owen, Rep. Brit. Assoc. for 1843, p. 237 (1844).

Platyceros, Wagner, Schreber’s Sdugethiere, vol. iv. p. 347 (1854).

Dactyloceros, Wagner, op. cit. vol. v. p. 352 (1855).

Palmatus, Giebel, Saugethiere, p. 351 (1859).

Characters.—Antlers normally without a bez (second), but with a trez
(third) tine, above which the beam is more or less palmated, and generally
furnished with numerous snags. Pelage of adult normally spotted with
white in summer, uniform in winter with a black-bordered white area in
the region of the tail, as in the sikine group; young spotted. Throat not
maned ; ears moderate ; tail rather long; head relatively short. Naked
portion of the muzzle much the same as in the red deer group. No upper
canines ; molars unusually broad transversely ; face-gland rather small ;
size medium or large.

Distribution.—Now restricted to the Mediterranean sub-region of the
Eastern Holarctic region, but during the Plistocene and Pliocene epochs
ranging over a large part of Europe.

Although the fallow deer were widely separated by Sir Victor Brooke
from the sikine group, there seems no doubt that the two are closely allied.
A year previously the late Professor Garrod! had written that ‘“‘ Dama

1 Proc. Zool. Soc. 1877, p. 18. —

126 Damine Group

vulgaris, as well as D. mesopotamica, from the shape of their antlers—neglect-
ing the palmation, an evidently insignificant character—are intimately
allied to the pseudaxine [equal sikine] group.” It may be added that, in
addition to the elaphine, these are the only groups of Cervus in which a
true trez-tine is developed, and that the two resemble one another in the
black and white markings of the caudal region. Moreover, occasionally a
bez- (second) tine is developed, thus adding to the elaphine resemblances.
Such a tine is exhibited on one side of a pair of antlers of the common
species in the collection at Woburn Abbey, where there are also two other
pairs showing a small knob on each side just above the brow-tine, which evi-

Fic. 32.—Fallow Deer at Woburn Abbey. From a photograph by the Duchess of Bedford.

dently represent a rudimentary bez. The occasional development of a bez-
tine in the common fallow deer has been pointed out by Mr. G. H. Fowler
in a paper published in the Proc. Zool. Soc. for 1894. Mr. Sclater, while
including the sikas in Cervus, separates the fallow deer as a distinct genus
(Dama), but there seems little justification for this course.

Typically the skull differs from that of the red deer group not only by
its shortness, but also by its greater breadth, and the larger size of the brain-
cavity and the eye-sockets. These characters are also found to a great
extent in the giant extinct fallow deer, which was shown by Ritimeyer to
be closely allied to the existing forms ; and since a connecting link, so far
as the antlers are concerned, has been subsequently discovered in Ruff’s

Common Fallow Deer 1a

fallow deer, there appears every reason for including all in a single group,

of which the proper name should be Ale.

1. THE Common FAttow DEER—CERVUS DAMA

Cervus dama, Linn. Syst. Nat. ed. 12, vol. i. p. 93 (1766) ; Jenyns, Brit.
Ve wAniuigss, p30 (0535) Bell, Brit. Quadrupeds, p. 402 (1837) ;
Keyserling and Blasius, Wirbe/th. Europ. p. 26 (1840) ; Brooke, Nature, vol.
Mieepe 2 ter(o74). iv0c. Zool. Soc. 1878, p. 913 ; Busk, Trans. Zool. Soc. vol.
x. p. 114 (1877) ; Ward, Records of Big Game, p. 43 (1896).

Cervus platyceros, Cuvier, Tableau élém. Hist. Nat. p. 160 (1798).

Cervus mauricus, de Blainville, fourn. Physique, vol. xciv. p. 261 (1822).

Cervus (Dama) dama, H. Smith, in Grithth’s Animal Kingdom, vol. iv.
Posy. p. 300 (1827).

Dama platyceros, Fitzinger, Bertr. Landesk. Osterreichs, vol. i. Pe 37
Gicne Speeder vol. ixvinl, part 1p. 350 (1873), Ixix. part 1. p. 547
(1874).

Dama vulgaris, Gray, List Mamm. Brit. Mus. p. 181 (1843), Cat.
Ungulata Brit. Mus. p. 200 (1852), Cat. Ruminants Brit. Mus. p. 74 (1872) ;
Loche, Expéd. Algérie, Mamm. p. 64 (1867) ; Sclater, Last Animals Zool.
Gardens, p. 171 (1883).

Cervus (Platyceros) dama, Wagner, Schreber’s Sadugethiere, vol. iv. p. 347
(1854).

Cervus (Dactyloceros) dama, Wagner, op. cit. vol. v. p. 352 (1855).

Cervus (Palmatus) dama, Giebel, Sdugethiere, p. 351 (1859).

Dama dama, \ataste, Act. Soc. Linn. Bordeaux, vol. xxxix. p. 288
(1885).

Characters—Height at shoulder about 3 feet. Antlers well palmated
and directed largely upwards, with the brow- and trez-tines simple, and the
front edge normally devoid of snags, which are, however, numerous on the
summit and hinder margin of the palmated portion. General colour of
summer pelage in the wild race a brilliant fawn, with large white spots
irregularly distributed over the back, upper part of the sides, and haunches ;
this spotted region being bounded inferiorly on the sides and posteriorly on
the haunches by indefinite white lines ; a blackish line running down the
middle of the back and tail; a white area, bordered above by black on the

128 Damine Group

buttocks below the tail, and the under side of the tail, under-parts, inner
surfaces of upper portions of limbs and inner sides of ears white or whitish.
In winter the whole pelage of the upper-parts uniformly grayish fawn.
Owing to long domestication, the fallow deer of the British parks
frequently display great variation from this original type of coloration, and
a uniformly dark brown breed has been long established, while white or
whitish varieties are far from uncommon. Indeed, specimens exhibiting any
shade from pure white to nearly jet black may be observed in large herds.

Fic. 33.—Head and Antlers of Common Fallow Deer. (Rowland Ward, Records of Big Game.)

As already mentioned, a bez-tine may occasionally be developed on the
antlers, and another abnormality is the presence of an additional tine on the
front edge immediately above the trez."

The antlers make their first appearance in the stags in the second year,
when they are in the form of simple snags, the animal being then known
as a pricket. In the third year the brow- and trez-tines show themselves,
while the extremity of the beam becomes palmated. In the succeeding
year the palmation increases, with strongly marked serrations on the hinder

1 See Brooke, Nature, vol. xi. p. 210 (1874).

Common Fallow Deer 129

border ; and in the fifth season the antlers have attained nearly their full
dimensions, although the palmation and its posterior snags do not reach their
complete development till the succeeding season. The antlers are shed
somewhat earlier in the spring than those of the red deer.

Mr. Rowland Ward gives the following dimensions of fallow deer
Antlers ——

Length on Basal Tip Spread Number of Width of

Outer Curve. _. Circumference. to Tip. inside. Points. Palmation.
30 43 235 265 10-9 43
295 5 17 285 14-13 Ts
29 45 305 24% 10-11 5S
284 4 144 26 IO-II 6
272 5 204 214 13-7 52
274 4 m8. 10-8 ?
26% 33 2 174 7-10 ?
265 45 235 205 7-9 56
26 5 202 232 10-10 +
254 ? 26 y si
258 4 26 8-8 sh
= ot 243 254 Pam St

The fallow deer of Epping Forest, which all belong to the uniformly
brown breed and have reverted to an almost wild state, are remarkable for
the narrowness of the palmation of the antlers, and since the same condition
obtains in those of the skeleton of a wild specimen from Asia Minor in the
British Museum, it might seem that this is the original form, and that the
great width characteristic of the antlers of most British park herds is due to
high feeding. But their dark pelage indicates that the Epping herd is a
specialised breed ; and a pair of antlers in the paleontological gallery of the
British Museum (No. M. 4104) obtained from Clacton, Essex, and said to
be from a deposit of the polished stone age, are as broad as those of any
modern park herd.

Distribution.—Southern Europe, namely Greece, Spain, Portugal, Ana-
tolia, Rhodes, and Sardinia ; parts of Asia Minor, Northern Palestine, and
North-Western Africa. In a semi-domesticated state in Great Britain, the
south of Sweden, Italy, and Tasmania. Fossil in the Plistocene deposits
of the Gibraltar caverns. |

With regard to the occurrence of the species in North Africa, the

s

130 Damine Group

evidence is not as full as might be desired, although that region doubtless
forms part of the habitat. Cuvier stated that wild fallow deer occur near
Tunis, and this was confirmed by Gervais, who said that they are found in
Barbary, Tunis, and Algiers, especially in the forest of Calle. Loche, in
1867, stated that he had seen a few specimens from the latter locality ; but
M. Lataste, writing in 1885, was unable to obtain any further information
with regard to them.

Since remains of fallow deer appear unknown in the peat of the English
fens, it has been generally believed that the existing form has been intro-
duced into the country from abroad ; the introduction being attributed to
the Romans. The above-mentioned antlers from Clacton, if really of the
age assigned to them, point, however, either to a much earlier introduction,
or to the species being indigenous.

In Jenyns’s British Vertebrate Animals it is stated that the dark brown
breed was introduced into Britain by King James the First from Sweden
(misquoted Norway in Bell’s British Quadrupeds), but it has been shown’
that this breed was living in Windsor Park soon after the middle of the
fifteenth century, so that its introduction (if introduced it was) must have
been at a much earlier date than the one given by Jenyns.

Habits. —Of the life of the wild fallow deer of the Mediterranean
countries there seems to be but little recorded, nearly all the voluminous
literature relating to park herds. The fallow deer is everywhere a
thoroughly gregarious species, going about in large herds; and in
most British parks these herds while feeding graze in the open. In
Epping Forest, however, Mr. J. E. Harting states that “they do not
associate in one herd, but roam about in ‘small parties, keeping to the
thickest underwood and most unfrequented part of the forest. Consequently
they are but seldom seen, and many people doubtless are unaware of their
existence.” For the greater portion of the year the bucks keep apart from
the does, grazing in separate herds, but in early winter, as well as during
the pairing-season, the two sexes are found in company. ‘The pairing-
season commences in September, by which time the antlers of the bucks
are free from the velvet, and the young are born in the following June or
July. Asa rule, only a single fawn is produced, although twins occasion-
ally occur. Mr. Harting was informed that the fawns of the brown breed

1 See Harting, Essex Naturalist, vol. i. p. 52 (1887).

Brown’s Fallow Deer Ean

in Epping Forest are not spotted, but those of the same breed are dappled
at Woburn, and I am told that faint spotting may be seen in Epping fawns.
The old bucks lose their antlers in May, and only a short interval elapses
before the new ones begin to bud. The date of changing the coat varies
somewhat in different parks. ‘The brown breed has the pelage darkest in
summer, whereas in the spotted variety the winter coat is the darker.

Although subsisting to a great extent by grazing, fallow deer are fond
of the leaves of most deciduous trees, and in autumn consume a considerable
quantity of horse-chestnuts. To reach overhanging boughs they may
frequently be seen raising themselves on their hind legs, and Bell states that
the bucks will use their antlers to knock off horse-chestnuts. Their times
of feeding vary according to the season. In winter, with the exception of
a short mid-day rest, they graze most of the daylight hours ; but in summer,
after feeding during the early morning, their period of repose lasts from
between nine and ten till two, after which they are again usually on the
move till about four. About this time they take another interval of rest,
which lasts till near six, when they once more take a three hours’ feed, after
which they lie down for the night. During winter they require to be
supplied with hay and corn. The only cry uttered by this species is a
grunting bark. According to Jenyns, the terms buck, doe, and fawn are
properly restricted to the two sexes and young of the fallow deer, the
names stag, hind, and calf being those originally applied in the same
senses to the red deer.

2. Brown’s Fattow DeER—CERVuS BROWNI (Extinct)

Cervus brownt, Dawkins, Quart. “fourn. Geol. Soc. vol. xxiv. p. 511
(1868), Brit. Pleist. Mamm. part vi., Cervide (Mon. Pal. Soc.), p. 17 (1887) ;
Lydekker, Cat. Foss. Mamm. Brit. Mus. part ii. p. 89 (1885).

Cervus clactomanus, Falconer, Pal. Memoirs, vol. ii. p. 478 (1868).

Cervus dama brown, Dawkins, Early Man in Britain, p. 98 (1880).

(?) Cervus savini, Dawkins, Brit. Pleist. Mamm. op. cit. p. 11 (1887).

Characters.—Closely allied to the last, of which its describer states it
may be only a variety. Antlers simpler, with the palmation smaller, the
posterior series of snags wanting, and an additional tine on the front edge
above the trez, which is only very rarely developed in the existing form.

132 Damine Group

The basal half of the antlers so closely resembles those of the latter that it
would be almost impossible to differentiate specimens with the coronal half
broken away.

Distribution —Typically from the Plistocene deposits at Clacton, Essex,
overlying the forest-bed. It does not appear, however, that the antlers
from the latter deposit described as C. savini can be satisfactorily dis-
tinguished as representing a distinct species, on account of the still smaller
development of the palmation. Not improbably this species, or race,
indicates a transition towards the next.

3. THe Mesoporamian Fattow DrER—CERVUS MESOPOTAMICUS

Cervus (Dama) mesopotamicus, Brooke, Proc. Zool. Soc. 1875, p. 265, 1876,
p. 298.

Cervus mesopotamicus, Brooke, op. cit. 1878, p. 914; Fitzinger, SB. Aa&.
Wren, vol. \xxix. part i. p. 62 (1879).

Dama mesopotamica, Sclater, List Anim. Zool. Gardens, p. 171 (1883).

Characters —Larger than C. dama ; the colour much brighter (as bright
as in the Indian spotted deer), the row of elongated white spots running on
each side of the dark median line of the back in the former coalescing into
a continuous band, and the black on the upper surface of the tail narrower
and confined to its root. Antlers of a totally different type, being some-
what expanded at the origin of the trez-tine, which is large and situated
some distance above the very short brow-tine, but at the summit are only
somewhat flattened, and break up at the crown and summit of the posterior
border into some four, five, or more snags.

The typical specimen was described in 1875, and in the following year
Sir Victor Brooke figured several additional shed antlers, which, allowing for
age, are all of the same general type as those of the former. In 1877 the
Zoological Society of London received a living male (which subsequently
bred with a hind of the common species), and in 1878 a female, while a
second specimen of the latter sex was presented by Lord Lilford in 1881.
The original pair bred together, although their progeny was unfortunately
a female. Beyond this little appears to be known of this very interesting
species. Fitzinger observes that “‘in my opinion this form is only a variety
of the common fallow deer, and the figured antlers are apparently of an

Mesopotamian Fallow Deer Lae

abnormal type, such as frequently occurs in other species of deer.” The
general constancy of type of the antlers—peculiar as they are—figured by
Sir Victor Brooke is sufficient to disprove this view, and to indicate that the
Mesopotamian fallow deer is a perfectly distinct species. Additional
specimens would, however, undoubtedly be most interesting. One of the

Fic. 34.—Head of Mesopotamian Fallow Deer. From a specimen in the British Museum.

figured antlers shows a duplication of the trez-tine. In addition to the type
specimen, the British Museum possesses a flattened skin belonging to a buck
with the antlers in a fairly advanced stage of development. It is from this
specimen that the accompanying figure has been drawn, the head being now
mounted in the Museum.

Distribution —The mountains of Luristan, in Mesopotamian Persia.

134 Damine Group

4. THe GriaAnr Farrow DrER—CERVvus GIGANTEUS (Extinct)

Alce gigantea, Blumenbach, Beitrdge Naturgeschichte, 1st French ed. vol.
ll. p. 407 (1803).

Cervus tslandicus, de Blainville, ‘fourn. Physique, vol. xciv. p. 261 (1822).

Cervus hibernus, Desmarest, Mammalogie, vol. ii. p. 446 (1822).

Cervus euryceros, Hibbert, Edinburgh ‘fourn. Sct. vol. vill. p. 129 (1825) ;
Pohlig, Paleontographica, vol. xxxix. p. 217 (1892).

Cervus megaceros, Hart, Descript. Foss. Deer Ireland (1826) ; Riitimeyer,
Abh, schweiz. pal. Ges. vol. x. p. 108 (1883).

Megaceros hibermcus, Owen, Rep. Brit. Assoc. for 1843, p. 237 (1844),
Brit. Foss. Mamm. and Birds, p. 444 (1846).

Cervus eurycerus, Giebel, Fauna Vorwelt, p. 145 (1847).

Cervus (Megaceros) hibernicus, Johnson, Trans. Norfolk Soc. vol. i. p. 288
(1877) ; Ball, Trans. Dublin Soc. ser. 2; vols it.cp. 337 4(109)5)

Cervus giganteus, Lydekker, Cat. Foss. Mamm. Brit. Mus. part ii. p. 82
(1885) ; Scharff, Mém. Soc. Zool. France, vol. viii. p. 453 (1895) ; Ward,
Records of Big Game, p. 45 (1896).

Characters—When fully developed, probably the largest of the brow-
antlered deer, males standing fully 6 feet in height at the shoulder when
in the flesh. Antlers very variable, but always with at least three tines on
the front border, of which the one above the trez is the longest, the beam
much expanded, and the brow-tine often flattened and bifurcated.

As represented by the Irish race—popularly but incorrectly known as
the Irish elk—this splendid deer was first described just two centuries ago,
an account of a skeleton being given by Dr. Molyneux in the PAz/osophical
Transactions of the Royal Society for the year 1697. It was then regarded
as indicating the former existence of the American elk in Ireland, the
author prefacing his description with the following observation :—‘ That
no real species of living creatures is so utterly extinct as to be entirely out
of the world since it was first created, is the opinion of many naturalists ;
and it is grounded on so good a principle of Providence taking care in
general of all its animal productions, that it deserves our assent.”

Since that date skeletons and skulls have been dug up by hundreds in

Irish bogs, while more imperfect remains have been discovered in many

Giant Fallow Deer mays

other European countries. In Ireland remains of hinds are much less
common than those of stags, and it was accordingly at first supposed that
both sexes were antlered. Probably the cause of the much greater abund-
ance of male skeletons is due to the antlers becoming entangled in roots
or on snags, and thus preventing the carcases from being washed away. To
support the enormous weight of the antlers, the neck-vertebre of the stags
are of very large proportionate size. Although the beam of the antlers is
very frequently directed at first outwards, the pedicles arise from a ridge on
the skull in a nearly vertical direction after the manner of other species of

Fic. 35.—Skull and Antlers of Irish Race of Giant Fallow Deer. From a specimen in the British Museum.

the genus Cervus, so that there is no resemblance in this respect to d/es ;
from which the species also differs in regard to the lateral metacarpal bones
of the fore-limb. In the presence of a brow, trez, supra-trez, and back-
tine to the antlers, the species resembles Brown’s fallow deer, to which
animal the closest approximation is made by one variety of the German
race, in which the antlers are directed more upwardly than usual, and the
supra-trez is situated very high up (Fig. 36). Whether the giant fallow
deer had a dappled summer coat like its living relatives, can, unfortu-
nately, never be determined. The maximum development of the antlers
occurring only in Western Europe, is a fact paralleled by the case of the
red deer. |

136 Damine Group

Distribution —Ireland, Isle of Man, England, Scotland, Denmark, France,
Belgium, Germany, Austria, Hungary, North of Italy, Russia, the Altai,
and Siberia,' during the Prehistoric and Plistocene periods.

a. GERMAN RacE—CERVUS GIGANTEUS RUFFI

Cervus megaceros rufi, Nehring, SB. Ges. Naturf. Berlin, 1891, p. 151
(1892), p. 3.

Cervus euryceros germania, Pohlig, Paleontographica, vol. xxxix. p. 220
(1892).

Cervus ruff, Lydekker, Royal Natural History, vol. ii. p. 365 (1894).

Characters-—Distinguished from the typical race by the smaller, nar-
rower, and more upwardly directed antlers, in which the whole palmation
is much curved inwardly, and the tines on the front border are much less
developed, the one immediately above the trez not being markedly larger
than the others. The specimens to which the name ruff was originally
applied (Fig. 36) have the antlers directed still more upwardly, with the
plane of the palmation placed nearly in a longitudinal direction, and the
tines above the trez situated near or at the summit. Although in all these
respects this type makes a most decided approximation to the existing
fallow deer, Dr. Pohlig has shown that it is merely an ultra-development of
the form he calls germani@,—a name which must give place to the earlier ruff.
Although I formerly regarded the type of the latter as indicating a distinct
species, more mature consideration leads to the conclusion that it would be
most improbable to have two species inhabiting the same area (the typical
germania being undoubtedly a mere race of giganteus), and I accordingly
adopt Dr. Pohlig’s view.

Distribution.—Europe during the upper portion of the Plistocene and
Prehistoric periods, remains of this form having been discovered in Ger-
many, France, Belgium, England, Russia, and probably Siberia and
Denmark. In geological age this race appears somewhat older than the
following, its remains occurring in cavern and river deposits in company
with those of the extinct cave-bear, whereas the Irish race is commonly
found in association with remains of a variety of the brown bear and of the
beaver.

1 See Tcherski, Das Janaland und die Neusibirischen, Inseln Mém, Ac. St. Pétersbourg, vol. x1. (1892).

Giant Fallow Deer L377

5. Irish RACE—CERVUS GIGANTEUS TYPICUS

Cervus euryceros hibernia, Pohlig, Paleontographica, vol. xxxix. p. 217
(1892).

Characters —The antlers very large, with the beam above the origin of
the brow-tine directed almost immediately outwards, the palmation enor-
mously developed and situated almost in a horizontal plane ;’ the brow-

Fic. 36.—Skull and Antlers of variety of the German Race of the Giant Fallow Deer. a, burr;
b, brow-tine ; ¢, trez-tine ; 4, back-tine ; ¢, supra-trez tine. (From Nehring.)

tine large, flattened, and generally forked ; and either four or five tines on
the front edge above the trez, of which the one immediately above the
latter (corresponding to the fourth tine of the wapiti) is the longest and
stoutest of all.

The back-tine is situated almost immediately opposite the trez, and Mr.

1 This is when the face is in the normal inclined position ; in the specimen here figured the skull is
placed horizontally and the front edge of the antlers consequently thrown upwards.

1h

138 Damine Group

Gordon Cameron has published a figure showing the almost exact corre-
spondence in the position of the tines of this race and those of a Scotch red
deer in which the bez-tine is wanting but the back-tine developed.
Whether the fact of the tine immediately above the trez being larger in this
form and the wapiti than either of the others is anything more than a
curious coincidence, is very difficult to determine. The large tine in ques-
tion corresponds to the one normally developed in the same situation in
Brown’s fallow deer, and very occasionally in the common fallow deer.
Rarely the brow-tine is undivided and much bent down over the face.

A large series of antlers of this race are preserved in the British and the
Dublin Museum ; the following dimensions being taken from Mr. Row-
land Ward’s book and Professor V. Ball’s memoir :-—

Toole Cares Sicatieene: sey idiom oe eile
P 10 Ii ft. 46 174 12-11
7 ft. 54 124 i | 194 |
5 » 83 98 10 4, 2 19 19
? 5 98 P ?
P 14+ Gigs) 20 BR
eis 134 Ones ait 13-15
? ? Gj, ee 154
6 ft. 13 83 8 ,, 112 173 10-13
? oh Pah te! 134
5 ft. 33 82 Fea 10 P 10-11

Distribution.—Ireland, and probably parts of Britain and Western Contin-
ental Europe in deposits belonging mainly to the Prehistoric period. This
race is the culminating form of the species, both as regards the development
of the antlers and its relative age. It certainly existed during the human
period, and in Ireland commonly occurs in the shell-marl beneath the peat,
although stated to be sometimes found in the peat itself.! Remains not
improbably referable to this form have been discovered in the peaty mud
near Newbury, Berkshire, and also in the marl below the peat at Maybole,
Ayrshire.

With much apparent truth, Dr. Pohlig points out that the enormous
dimensions and wide span of the antlers of this magnificent deer would seem

1 See Boyd Dawkins, Ear/y Man in Britain, p. 258.

Giant Fallow Deer I 39

to preclude its being an inhabitant of forests, and that it must apparently
have dwelt in more or less open country, although the other races of the
species may well have been woodland animals. The cause of its extinction
presents a difficult problem.

It may be mentioned that as the name C. Aibernicus is a synonym of
giganteus, it cannot be employed in the sense proposed by Dr. Pohlig as the
title of the typical race.

Fic. 37.—Antlers of Irish Race of Giant Fallow Deer at different ages. a, burr; 4, brow-tine ; ¢, trez-
tine ; 4, back-tine. After Owen.

c. IraLtIAN RaAacE—CERVUS GIGANTEUS ITALLE

Cervus euryceros italia, Pohlig, Paleontographica, vol. xxxix. p. 228
(1892).

Characters.—Size relatively small; the antlers comparatively simple,
with the palmation narrow, much inclined upwards, and the front border
curved inwardly so that much of the outer surface is seen in a front view ;
all the tines being small, and those above the trez few in number and
placed near the summit of the palmation.

Distribution.—Italy, Hungary, and probably other Mediterranean
countries, in deposits of approximately equivalent age to those yielding
remains of the German or northern race.

d. FRENcH RacE—CERVUS GIGANTEUS BELGRANDI

Cervus belgrandi, Lartet, in Belgrand’s La Seine, vol. i. Pa/. p. 13, plate
xvill (1869).
Cervus euryceros belgrandi, Pohlig, Paleontographica, vol. xxxix. p. 232

(1892).

140 Damine Group

Characters.—Impertectly known, but characterised by the rudimentary
condition of the brow-tine, and the almost horizontal direction of the base
of the beam of the antlers, of which the extremity is unknown.

Dr. Pohlig suggests that this race shows close affinity with the true
elks, but this view I am not prepared to accept. He further points out
that in the abortion of the brow-tine the antlers approximate very closely
to those of the so-called Cervus dawkinsi, mentioned under the head of the
next race, although he adds that the former come very close to those
described as C. verticornis, with which they are contemporary in age.

Distribution —Germany and France during the middle portion of the
Plistocene period.

e. FoREST-BED RAcCE—CERVUS GIGANTEUS CARNUTORUM

Cervus megaceros carnutorum, Langel, Bull. Soc. géol. France, ser. 2, vol.
Mi. p. 711 (sez).

Cervus carnutorum, Dawkins, Quart. “fourn. Geol. Soc. vol. xxvili. p. 409
(1872).

Cervus verticornis, Dawkins, op. cit. p. 406 (1872), Brit. Pleist. Mamm.
part vi. Cervide (Mon. Pal. Soc.), p. 22 (1887) ; Lydekker, Cat. Foss. Mamm.
Brit. Mus. part 1. p. 92 (1885).

Cervus dawkinsit, Newton, Vertebrata of Forest-bed, p. 54 (1882),
Vertebrata of Phocene Deposits, p. 26 (1891); Dawkins, Brit. Pkest.
Mamm. op. cit. p. 7.

Cervus gunn, Newton, Vertebrata of Forest-bed, p. 57 (1882), Vertebrata
of Pliocene Deposits, p. 28 (1891).

Cervus fitchii, Newton, Vertebrata of Forest-bed, p. 56 (1882), Vertebrata
of Pliocene Deposits, p. 28 (1891).

Cervus euryceros carnutorum, Pohlig, Paleontographica, vol. Xxxix. p. 233
(1892).

Cervus euryceros dawkinst, Pohlig, /oc. cit.

Characters.—Antlers stout, directed upwards and outwards, with the
palmation generally slight, and the brow-tine large, simple, and inclined
mainly downwards ; higher up come the trez and the tine above it, while
the crown apparently forms two or three points, with a back-tine on the

posterior border.

Rusine Group 141
Such is the typical C. verticornis of the Norfolk forest-bed, which I

agree with Dr. Pohlig in regarding as inseparable from the earlier-named
C. carnutorum from equivalent beds in France. The single antler from the
forest-bed described as C. dawskinsi differs by the rudimentary condition of
the brow-tine, and the greater palmation of the crown, which terminates
in more points. As already mentioned, this type approximates in some
respects to the French race, while in its greater expansion it indicates a
transition towards the Irish race. As both types occur in the same
geological horizon, Dr. Pohlig appears undoubtedly right in regarding
them as mere individual variations of one and the same form. There
can likewise be no question as to their near relationship with the more
typical forms of C. giganteus, of which they indicate the ancestral stock.
Whether they should be regarded merely as a race of that species, or as
a separate species, may be a doubtful point, although I have provisionally
adopted the view of Dr. Pohlig. In the case of extinct forms constituting
a completely connected series, it seems advisable to regard sub-species in a
somewhat wider sense than when dealing with recent animals.

Distribution —England and France during the early portion of the
Plistocene period.

iv. THE RusInE oR SAMBAR GRoup—SuB-GENus Rusa

Rusa, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 309 (1827), as
a sub-genus ; Gray, List Mamm. Brit. Mus. p. 179 (1843), Cat. Ungulata
Brit. Mus. p. 205 (1852), Cat. Ruminanis Brit. Mus. p. 76 (1872) ;
Fitzinger, SB. 4k. Wien, vol. \xviil. part i. p. 354 (1873), Ixx. part 1.
p- 279 (1874); Brooke, Proc. Zool. Soc. 1878, p. goo; Riitimeyer, AdA.
schwetz. pal. Ges. vol. vill. p. 45 (1881).

Axis, H. Smith, in Griffth’s Animal Kingdom, vol. v. p. 312 (1827), as
a sub-genus ; Gray, List Mamm. Brit. Mus. p. 178 (1843), Cat. Ungulata
Brit. Mus. p. 212 (1852), Cat. Ruminants Brit. Mus. p. 79 (1872) ;
Paginect. oi. 42. Wen, vole Ixvau: patt 1. p. 354 (1873), xx. part 1.
p. 264 (1874) ; Brooke, Proc. Zool. Soc. 1878, p. 907; Riitimeyer, 4d/.
schweiz. pal. Ges. vol. viii. p. 44 (1881).

Hippelaphus, Sundevall, K. Svenska Vet. Ak. Hand/. for 1844, p. 178
(1846), as a sub-genus.

142 Rusine Group

Hyelaphus, Sundevall, op. cit. p. 181 (1846) ; Gray, Cat. Ungulata Brit.
Mus. p. 215 (1852), Cat. Ruminants Brit. Mus. p. 79 (1872).

Ussa (Oussa), Heude, Meém. hist. nat. emp. Chinois, vol. 11. p. 20 (1888).

Sambur, Heude, op. cit. pp. 20 and 41 (1888).

Melanaxis, Heude, of. cit. p. 47 (1888).

Characters —Antlers rounded, three-tined, with both the bez (second)
and trez (third) tines wanting, and the beam simply forked at the ex-
tremity. Pelage either uniform, or spotted, at all seasons and all ages, or
intermediate between the two, without a light-coloured caudal disk, or
black-bordered white area on the buttocks. Throat and neck often heavily
maned ; ears usually large; tail relatively long. Naked portion of the
muzzle large, extending on to the upper surface of the face, and not
constricted above its junction with the upper lip. The face - gland,
or tear-pit, frequently very large and capable of complete eversion.
Upper canines small or wanting; upper molars tall-crowned, with a
small additional column on the inner side. Size large, medium, or
small.

By previous writers the Indian spotted deer has been invariably
separated as a distinct sub-genus or genus (Axis), but its antlers are of
essentially the same type as those of some forms of Rusa proper, and the
latter are intimately connected with it through the Philippine spotted
deer and the hog-deer. Indeed, by those writers who do not separate
it sub-generically or generically as Hye/aphus, the latter species is
referred alternately to Rusa and to Axis. Accordingly, the latter group
is merged in the former, of which it doubtless represents a less specialised
type.

It has been customary to regard the anterior or inner tine of the
terminal fork of the antlers in this group as corresponding to the trez
(third) tine of the red deer group,’ but there seems no reasonable doubt
that Mr. A. Gordon Cameron is correct in his view that the trez-tine is
unrepresented in this group, as it also is in the rucervine group.

The simple form of the antlers indicates that the group is a generalised
one. The original type would appear to be represented by the Indian
spotted deer, or chital, in which the white spots are persistent at all ages
and all seasons. A more advanced modification is displayed by the hog-

' Compare the figures in Sir Victor Brooke’s monograph of the Cervide.

*

Characters 143

deer, in which the spots disappear in the adult during the winter. But
the most specialised forms of all are the various kinds of sambar, in which
the adult is uniformly coloured at all seasons, while even the fawn may
have lost its dappling. It is probable that the group reached the Oriental
region at a comparatively early epoch (remains of fossil species occurring

Fic. 38.—Frontlet and Antlers of Indian Sambar. From a specimen from Simrol, in the
British Museum. (Rowland Ward, Recoras of Big Game.)

in the Pliocene strata of the Siwalik Hills), and that it has had no con-
nection with the ancestors of any of the preceding groups since the early
part of the Pliocene epoch, when it may have branched off from the
sikine stock. Numerous remains from the Pliocene deposits of Europe
have been assigned to the present group, but at least a considerable propor-
tion of these seem rather to belong to sikas.

Distribution —The Oriental region.

144 Rusine Group

1. THE SAMBAR—CERVUS UNICOLOR

Cervus unicolor, Bechstein, A//gem. Uebers. vierfiis. There, vol. i. p. 112
(1799); Blanford, Fauna Brit. India—Mamm. p. 543 (1891); Ward, Records
of Big Game, p. 3 (1896).

Cervus albicornis, Bechstein, /oc. cit. (1799).

Cervus niger, de Blainville, Bul/. Soc. philom. Paris, 1816, p. 76.

Cervus aristotelis, Cuvier, Ossemens Fossiles, ed. 3, vol. iv. p. 503
(1825) ; Sclater, Trans. Zool. Soc. vol. vii. p. 350 (1871) ; Brooke, Proc.
Zool. Soc. 1878, p. gor ; Lydekker, Cat. Foss. Mamm. Brit. Mus. part ii.
p. 103 (1885) ; W.L. Sclater, Cat. Mamm. Ind. Mus. part ii. p. 176 (1891).

Cervus leschenaulti, Cuvier, op. cit. p. 506 (1825).

Cervus (Rusa) unicolor, H. Smith, in Grithth’s Animal Kingdom, vol. iv.
p. 108; v. ps gio (1827).

Cervus (Rusa) aristotelis, H. Smith, of. cit. pp. 110, 310 (1827).

Cervus jarat, Hodgson, Gleanings in Science, vol. ill. p. 321 (1831).

Cervus heterocervus, Hodgson, ‘fourn. As. Soc. Bengal, vol. x. p. 721
(1841).

Rusa jarya, Hodgson, of. cit. p. 914 (1841).

Rusa nepalensis, Hodgson, /oc. cit. (1841).

Rusa heterocervus, Hodgson, /oc. cit. (1841).

Rusa aristotelis, Gray, List Mamm. Brit. Mus. p. 179 (1843), Cat.
Ungulata Brit. Mus. p. 205 (1852), Cat. Ruminants Brit. Mus. p. 76
(1872) ; Jerdon, Mamm. India, p. 256 (1867) ; Fitzinger, SB. Ak. Wien,
vol. Ixx. part i. p. 279 (1874) ; Sterndale, Mamm. India, p. 503 (1884).

Cervus Hippelaphus aristotelis, Sundevall, K. Svenska Vet. Ak. Hand. for
1844, p. 178 (1846).

Cervus (Hippelaphus) niger, Sundevall, op. cit. p. 183 (1846).

Cervus (Hippelaphus) leschenaulti, Sundevall, /oc. cit. (1846).

Cervus (Hippelaphus) unicolor, Sundevall, /oc. cit. (1846).

Rusa aristotelis migra, Fitzinger, SB. Ak. Wien, vol. \xx. part 1. p. 284
(1874).

Rusa aristotelis leschenaulti, Fitzinger, op. cit. p. 286 (1874).

Rusa aristotelis unicolor, Fitzinger, op. cit. p. 287 (1874).

Rusa aristotelis heteroceros, Fitzinger, op. cit. p. 289 (1874).

Sambar ies

Characters.—Size typically large. Hair coarse and shaggy, the hairs
on the back not distinctly banded with differently coloured rings ; colour
of adult uniformly some shade of dark umber-brown, with chestnut or
whitish on the inner side of the buttocks, and often on the under-parts and
limbs ; young more or less uniformly coloured, or faintly spotted on the
hind-quarters. Antlers large, stout, and very rugose, with the brow-tine
generally long and making an acute angle with the beam, and the front or
outer tine of the terminal fork forming the continuation of the line of the
beam when there is any inequality in the length of the two tines; the
space enclosed by the antlers of opposite sides generally V- or U-shaped,
although the tips of the antlers frequently are inclined inwards ; pedicles
of antlers short. Ears typically large, but relatively shorter in the smaller
races ; face-gland greatly developed, and capable of complete eversion ;
neck and throat thickly maned ; tail comparatively long, and more or less
bushy.

Few groups of deer are more difficult to understand than the various
kinds of sambar; and unfortunately the series of specimens in our
museums is far too incomplete to render possible a decisive solution of the
difficulty. This can only be done by the acquisition of an extensive series
of examples with the localities fully authenticated. From time to time
numerous examples of the group are exhibited in the Zoological Society’s
Menagerie, and many are now living in the park at Woburn Abbey ; but
in too many instances such specimens are purchased from dealers without
any definite information as to their place of origin, and they consequently
rather increase than diminish the confusion.

Very different views have been entertained as to whether the various
modifications of the sambar type indicate distinct species, or merely races
of one very variable form ; the latter view being adopted by the late Mr.
Blyth and subsequently by Mr. Blanford. My own observations lead to
the belief that all the kinds in which the front or outer tine of the terminal
fork of the antlers forms the continuation of the line of the beam, where
there is any inequality in the length of the two, are but varieties of a single
species. But, on the other hand, I am inclined to regard those forms in
which the back or inner tine is situated in the direct line of the beam as
representing a second specific type. Whether all the local modifications
of the first type are truly indigenous forms, and therefore entitled to rank

U

146 Rusine Group

as valid sub-species, is still an open question; and I confess to a great
ditiiculty in determining specimens of which the place of origin is unknown.
In the case of hinds and stags without antlers, I believe it to be frequently
a matter of impossibility even to determine to which of the two specific
types they belong.

In the skull of the present species the nasal bones, when the animal is
fully adult, develop a plate at the posterior expansion, which tends to grow
over the lachrymal vacuity ; and in some of the smaller races—notably the
Philippine—the extension is so great as to reduce the vacuity to a mere
fissure.

Distribution.—The undulating or hilly wooded districts of a large
portion of the Oriental region, namely India, Ceylon, Assam, Burma, the
Malay Peninsula, Borneo, Siam, W. China, Hainan, Formosa, and the
Philippines.

a. INDIAN RacE—CERVUS UNICOLOR TYPICUS
Plate X

Characters.—Size very large, the height at the shoulder reaching 5 feet
4 inches, or perhaps more. Antlers long, with the two tines of the terminal
fork generally of approximately equal length, but if unequal, the front one
usually the shorter (Fig. 38), and the hinder one arising from the posterior
surface of the beam and not forming the continuation of the axis of the
latter. Colour almost uniformly dark umber-brown, but tending to gray or
yellowish in some individuals ; under-parts very little paler than the back,
but the chin, inner portion of the buttocks, the lower surface of the tail,
and the inner side of the upper-part of the limbs more or less distinctly
chestnut ; hinds paler than the stags; young reddish, probably with a
black tail and dorsal band, but spots wanting even when newly born. Ears
very broad, about equal to half the length of the head ; the latter fairly
long, with a comparatively straight profile ; tail moderately bushy, and
longer than the ear.

The following dimensions of antlers are taken from Mr. Rowland
Ward’s Records of Big Game, but Mr. Blanford mentions a larger pair, of
which the length is 48 inches. Antlers vary enormously in girth, length,
and span, the longest specimens being by no means the stoutest. Any

S.

7
We

A

Ee AE AN

THe DEER <

INDIAN SAMBAR, STAG AND HIND.

Pablished by Rowland Ward. Ltd

Indian Sambar 147

examples over 35 inches in length may be considered good ; even such
dimensions being, I believe, never reached by the Malayan sambar.

oaeatnces above Brow-tine, TIP to Tip. Tae Lae
463 6% 245 30% ;
45} E 173 324
45 72 224 332 Central Provinces.
45 7t 445 Western Ghats.
444 1 442 45% Central Provinces.
44 z 34 ? >
44 6 Ox 24% >
43 714 354 38 Gwalior.
422 52 26 324 Central Provinces.
424 (Fig. 38) 64 15 272 Ghats of Simrol.
41g 64 335 354 Western Ghats.
41f 284 34 ;
412 M 312 362 Nepal.
4it 64 304 363 Central Provinces.
41 63 183 29 »
403 58 33 374 :

Out of a total of sixty-eight specimens catalogued by Mr. Rowland Ward
in only seven does the number of points reach four, the extra tine in most
cases being only developed on one side; so that all such variations can only
be regarded as abnormal sports.

Distribution.—India and Ceylon, probably extending eastwards as far as
the Assam valley, where the range may be separated by the Bramaputra
from that of the next sub-species, with which, however, it may intergrade
in this district. The Ceylon form is a smaller animal than the one from
the mainland. Sir Victor Brooke included Burma, Siam, and Hainan in
the range of this race ; but, as suggested by Mr. W. L. Sclater, there can
be no doubt that those countries should be included in the habitat of the
Malayan sambar. Regarding its distribution in India, Mr. Blanford remarks
that “the sambar ascends the Himalayas in places to gooo or 10,000 feet,
and is common on the summits of the ranges in Southern India and Ceylon.
It is not common on alluvial flats, though it is occasionally found on them,
at considerable distances from the hills. It is, of course, wanting in the
treeless plains of the Punjab, Sind, and Western Rajputana.”

148 Rusine Group

Till the name waicolor was revived by Mr. Blanford, the sambar was
very generally known as C. arvstote/is. Concerning the reasons for the
rejection of the latter name that author writes as follows :—‘‘ This fine deer
appears to have been first mentioned by Pennant, who described it as the
middle-sized and greater axis (Cervus axis unicolor and C. axis major of Kerr).
To these forms the names of Cervus unicofor and C. albicornis were applied
by Bechstein. Cuvier, in the second edition of his Ossemens Fossiles, named
different varieties C. Aippelaphus and C. eguinus, and two years afterwards
added the names of C. artstotehis and C. /eschenaultn, given to horns only.
Why the name C. arvstote/is, given to an abnormal horn, has been preferred
for the Indian sambar it is difficult to say. The name C. wnicofor, employed
by Hamilton Smith, is preferable on account of both priority and suitability,
being an appropriate term for the only Indian deer with unspotted young.”

Imperfect skulls and antlers of a rusine deer from the Plistocene
deposits of the valley of the Narbada, India, have been provisionally
assigned by myself to the sambar, of which they may indicate an extinct
race. Other remains from the Pliocene formations of the Siwalik Hills
not improbably belong to more or less closely allied extinct species.

Although sambar is the Hindustani name, and the one generally adopted
by sportsmen, in Nepal the stag is known as jarao, and the female as
jarai, and these latter terms are sometimes employed in sporting literature.
Plate x represents a stag and hind living in the menagerie at Woburn
Abbey in 1897.

Habits.—The life-history of the Indian sambar has been described by
several authors, among whom may be mentioned the late Sir Samuel Baker!
and Captain Forsyth.2- Although both stags and hinds are not unfrequently
found alone, sambar generally go about in small parties, varying in number
from four or five to about a dozen. Their favourite haunts are rocky hills
and ranges, which are well wooded, and thus afford abundance of shady
retreats. From such covert sambar issue forth at evening to graze in any
open grassy glades there may be in the neighbourhood, or to feed upon
plantations of sugar-cane, cinchona, etc. The young shoots and leaves of
trees, as well as various wild fruits, also form an important part of their diet.
Whether feeding on the upland glades or the cultivated tracts lower down,

1 Wild Beasts and their Ways, p. 408.
2 The Highlands of Central India.

Indian Sambar 149

these deer always retire to the higher woods to pass the day ; and there
select a lair well sheltered from the sun where grass is abundant. To obtain
water they travel long distances, but there is some difference of opinion as
to whether a daily drink is necessary to their well-being.

According to Sir Samuel Baker, a sambar does not attain its full develop-
ment of body and antlers until ten years old. A remarkable peculiarity of
this, and probably several of the other races of sambar, is that the antlers of
the stags are not replaced annually. On this point Sir Samuel Baker says :-—
“They are not shed annually, but with great irregularity every third or
fourth year. This has been established as a fact by those which have been
for some years kept in confinement, and it is generally accepted by all
natives who are experienced shikaris. During eight years’ hunting in
Ceylon, I killed a vast number of sambar throughout all seasons, and there
was no particular month when the antlers were shed ; the deer were found
with horns in every stage of growth, irrespective of periods or localities.”
Similar testimony is afforded by Messrs. Forsyth and Blanford, the former
of whom states that he knew several individual stags which retained their
antlers for successive years. In the plains of India the antlers are usually
dropped in March, but not till about a month later on the spurs of the
Himalaya. Among the stags at Woburn Abbey the antlers are shed
annually, but the time of shedding in all the members of the group, and
also in the chital, is very variable ; and the fawns may be dropped at any
time of year.

In the plains of India the new antlers of those individuals in which the
old ones have been shed are generally free from the velvet about September,
and the pairing-season takes place during the months of October and
November, although in the Himalaya it is stated by Brian Hodgson to be
deferred till the spring. During the season in question sambar collect in
larger parties than at other times of the year ; the old stags calling in the
mornings and evenings and sometimes far on into the night. The call of
the stags has been described as a loud and somewhat metallic-sounding
bellow, and that of the hinds as a faint grunting low. In the pairing-season
the stags stalk about with the tail erected, the muzzle stretched out, and
the eye-pits so completely everted as to expose the pink mucous membrane
with which they are lined. In the wild state the stags at this time seem
to confine their attacks to members of their own species, on whom they

150 Rusine Group

inflict frightful gashes with the long brow-tines of their antlers; but in
captivity they are dangerous to approach. In addition to the pairing-call,
sambar utter a hissing kind of snort, frequently accompanied by a stamp of
the foot, when alarmed by the presence of a leopard or tiger, or the approach
of a human being.

The fawns in the plains of India are born during June and July, and it
is but seldom that more than one is produced at a birth. As already said,
they are entirely without spots.

Although extremely active, and capable of rushing down the steep side
of a rocky cliff with great rapidity, sambar in the open display only a
moderate degree of speed, and can be ridden down with comparative ease
by a well-mounted man. They are, however, difficult to kill, and will
carry wounds that would be fatal to many others of their kindred. Although
coarse-grained, the flesh is well flavoured.

A large herd of sambar of various races is kept at Woburn ABER
where they flourish well, although apt to get out of condition during the
winter. The secret of success seems to be that they are kept in large open
paddocks, devoid of covert. In Ireland, where they were tried some years
ago by Lord Powerscourt, they were allowed to wander in the park, but
failed to thrive. ‘It was a curious thing with the sambar deer,” writes his
lordship,' ‘‘and it was no doubt the cause of their death, that they would
never come out of the thickets in the daytime. They unfortunately could
not be taught that Ireland has not a tropical climate, and they used to skulk
in the thickest covert they could find, out of the sun, all day, and only
came out to feed at night, as they would in the jungles of Mysore, where I
have seen them pursuing the same tactics. Of course by this unnecessary
precaution on their part against the meridian rays they got chilled through,
and eventually died.” At Woburn sambar may be seen feeding at all hours
of the day in the open.

5b. MALAYAN RaceE—CERVUS UNICOLOR EQUINUS

Cervus equinus, Cuvier, Ossemens Fossiles, ed. 2, vol. iv. p. 45 (1823) ;
Brooke, Proc. Zool. Soc. 1878, p. go1 ; Giinther, sid. 1880, p. 452 ; Ward,
Records of Big Game, p. 7 (1896).

! Proc. Zool. Soc. 1884, p. 208.

Tum DEER or ALL LANDS. Puate Xl,

MALAYAN SAMBAR.

Published by Rowland Ward Lid

Malayan Sambar FS

Cervus mataccensis, F. Cuvier, Hist. Nat. Mamm. vol. i. plate x (1824).

Cervus (Rusa) equinus, H. Smith, in Griffith’s Animal Kingdom, vol. iv.
Pe LS, ive pr 210 |(1 8277).

Rusa equina, Gray, List Mamm. Brit. Mus. p. 179 (1843); Jerdon,
Mamm. India, p. 260 (1867) ; Fitzinger, SB. Ak. Wien, vol. |xx. part i.
p- 290 (1874).

Axis pennanti, Gray, List Mamm. Brit. Mus. p. 179 (1843).

Cervus (Hippelaphus) equinus, Sundevall, K. Svenska Vet. Ak. Hand/. for
1844, p. 178 (1846).

Rusa equinus, Gray, Cat. Ungulata Brit. Mus. p. 210 (1852), Cat.
Ruminants Brit. Mus. p. 77 (1872).

Rusa equina mataccensis, Fitzinger, SB. Ak. Wien, vol. |xx. part i. p. 294
(1874).

Rusa equina pennantu, Fitzinger, /.c. p. 296 (1874).

Cervus brooket, Hose, Ann. Mag. Nat. Hist. ser. 6, vol. xu. p. 206

(1893).
Plate XI

Characters—Nearly as large as the typical Indian sambar ; but the
antlers generally shorter and thicker, with the hinder or inner tine of the
terminal fork much shorter than the front or outer tine, and arising as a
spur from the inner hind margin of the beam, of which the hinder tine
forms the direct continuation ; the brow-tine also in most cases relatively
longer. General colour darker, approaching to black or slaty gray in old
stags ; generally a light ring round the eye; the ears rather smaller, often
with a distinct white margin ; the legs frequently light-coloured ; and the
tail much more bushy. Face comparatively long and straight. Very
young fawns, at least frequently, distinctly or faintly spotted on the hind-
quarters, the general colour being foxy red, with the tail and a line down
the back blackish er black.

That this deer is not entitled to rank as a distinct species may, I think,
be regarded as certain, although it appears to have good claims to sub-specific
distinction. After remarking on its difference from the typical sambar, Sir
Victor Brooke states that he has seen skulls and antlers from Siam which are
intermediate between the Indian and the Bornean races. None have, how-
ever, ever come under my own notice from the countries to the east of the

Bay of Bengal which resemble those of the Indian race. Although very

152 Rusine Group

general constant, the form of antler described above is not absolutely so,
Mr. Blanford possessing a Burmese skull in which the hind tine of the
terminal fork is the longer on one side, and the front tine on the other ;
but even in the former case the antler is not of the type of that of C. Aipp-
elaphus.

More precise information is required with regard to the coloration of
the young. In the writings of Messrs. Jerdon and Brooke all the sambar are
credited with having unspotted young;
and this view was followed by Mr. Blan-
ford. A correspondent of the Aszan
newspaper, writing about the year 1892
or 1893, mentioned, however, that in
Cachar sambar always produce spotted
young, and I have some reason to believe
that this is also the case in Burma.” Tn
1893 Mr. C. Hose described a very young
sambar fawn from Mount Dulit, in North
Borneo, which he regarded as distinguish-
able from the present form on account of
being spotted, and to which he applied
the name C. drookei. In this specimen,
which is now in the British Museum,
the general colour is foxy red, with faint

yellowish spots on the hind-quarters ; the
Fic. 39.—Frontlet and Antlers of Malayan tail and.a streak down the back being
Sambar. From a Burmese specimen in black. Another fawn in the Museum,
iva eee aa a a also sent by Mr. Hose from Borneo, and
labelled C. equinus, is of approximately the
same age, and shows a few very faint traces of spots on the hind-quarters.
In the skin of a third and somewhat older fawn in the Museum, likewise
sent by the same gentleman from Borneo, the spots are wanting, and the
head and neck have begun to assume the adult grayish colour. Probably
the spots have disappeared. A spotted fawn was born at Woburn Abbey
in January 1898.
Mr. Hose’s original description of his presumed species is as follows,

viz. :—

Malayan Sambar 15.3

“General colour of the fawn rufous, browner on head, neck, and
shoulders, richer on the posterior back, fore-limbs from elbows downwards,
thighs, and outer sides of hind-limbs. Muzzle and sides of face brown ;
forehead and crown rufous. Ears externally dark blackish brown, edged
with pale rufous, internally whitish. Back with an indistinct mesial black-
ish line. Shoulders and rump spotted, the spots small, yellowish, few in
number on the shoulders, more numerous on the upper side of the rump.
Inner sides of limbs, both fore and hind, and belly whitish, with a strong
suffusion of rufous, especially towards the feet. Chest between the fore
limbs deep shining black, this colour extending backwards along the lower
part of the sides, where it is divided in the centre by the whitish or
yellowish belly-colour. Middle of metatarsals with an elongated tuft of rich
rufous hairs, tipped with black. Feet rufous, without black markings.”

He adds that the young differs from the typical C. unicolor equinus by
being fully spotted, instead of being almost or quite unspotted ; and also
states that it does not show the same striking contrast between the deep
black of the chest and tail and the brilliant rufous of the sides and rump.
The latter differences, if constant, I should not regard as of even sub-
specific value. And the cases cited prove that very young fawns of the
Malayan sambar are frequently, if not invariably, faintly spotted.

The following dimensions of antlers are taken from Mr. Rowland
Ward’s book ; the first five specimens in his list are, however, omitted, as
being from Java, and thus apparently referable to C. hippelaphus.

Length along Basal Widest

Outer Curve. Circumference. Tip to Tip. inside, LENA
1 3 S) 1
308 4¢ 207 2 Aa Burma
292 6 162 202 e
1 ab al il
281 44 114 124 P
264 63 a4k 2 Perak
234 q] 152 Borneo
5. aL 4b WE ?
192 44 164 16g f
164 4d 52 ? Perak

Distribution—Apparently ranging from Assam and Cachar through
Burma and the Malay Peninsula to Siam, Hainan, Borneo, and, it is said,
Sumatra. The reasons for regarding the Burmese sambar as referable to
this and not the preceding race have already been given. There are antlers

x

154 Rusine Group

trom Assam in the British Museum showing all the characteristics of the
present race ; and the aforesaid occurrence of sambar with spotted young in
Cachar shows that the form from that district does not belong to the
typical Indian race.

In respect to habits this race probably shows no important difference
from the preceding.

c. Formosan Race—CEeErRvus UNICOLOR SWINHOEI

Rusa swinhon, Sclater, Proc. Zool. Soc. 1862, p. 152, Trans. Zool. Soc. vol.
vil. p. 333 (1871) ; Gray, Cat. Ruminants Brit. Mus. p. 77 (1872).

Cervus swinhou, Brooke, Proc. Zool. Soc. 1878, p. go1; W. L. Sclater,
Cat. Mamm. Ind. Mus. part ii. p. 178 (1891).

Rusa swinhoet, Fitzinger, SB. Ak. Wien, vol. xx. part i. p. 298 (1874).

Cervus swinhoet, Ward, Records of Big Game, p. 8 (1896).

Characters.—Very closely allied to the preceding, from which it may be
distinguished by its shorter head and concave profile, the relatively longer
legs, and the undermentioned differences in colour ; the stature being some-
what less. In the winter pelage the general colour of the upper-parts is
uniform reddish black-brown, the head and ears being reddish yellow-
brown, and the upper surface of the nose having a V-shaped blackish brown
mark reaching to the eyes ; the under surface of the body is nearly as dark
as the back, but the inner side of the thighs and the whole lower legs are
brownish or whitish yellow, and the bushy tail is black all round. In the
summer coat the general colour is light yellowish red-brown, darker in
front than behind, and lightest on the under surface. The antlers are of
the type of those of the Malayan race, but smaller, the longest recorded
length being just under 20 inches; and the skull is very similar to that
of the Luzon race.

In his description Mr. Sclater remarked that this deer was very similar
to the Malayan race, from which he was unable to point out distinctive
differences, having never seen an adult of the latter. A short face and
concave profile are stated by Sir Victor Brooke to be characteristics of red
and fallow deer living under unfavourable conditions ; and in the case of
the present form the same features may be due to the comparatively small
area of its habitat. Indeed, it seems highly probable that the Malayan,

Formosan Sambar rcs

Formosan, and Szechuan races of the sambar will eventually have to be
included under a single sub-specific title.

Mr. Swinhoe states that the young, when about half-grown, “is reddish
brown, with the tail bushy and black, but reddish at its root ; sides of the
body paler, and the belly blackish brown ; legs pale towards the hoofs, the
latter black ; under surface of tail, abdomen, and inner sides of hind-legs
down to middle of shank yellowish white, the breast and belly being
blackish brown ; under surface of head and neck mottled whitey-brown ;

Fic. 40.—Formosan Sambar, from a male at Woburn Abbey. Photographed by the Duchess of Bedford.

crown of the head, with many of the hairs, tipped with black ; from the
occiput a dark line runs down to the base of the tail ; ears blackish brown,
tipped and margined with ochreous white, and whitish on their insides.”
It is not known whether the very young are spotted ; if they are not, there
would be a distinction from the preceding race.

Distribution —The island of Formosa. The type specimen was pre-
sented to the London Zoological Society in 1862, and is now in the British
Museum. ‘The society subsequently received three other examples, the
last in 1868. Recently two examples apparently referable to this form
were imported by a London dealer, one of which was sold to the Paris
Zoological Gardens, while the second is living in the park at Woburn

156 Rusine Group

Abbey, and shows all the characteristics of the type. It is the latter that
forms the subject of Fig. 40.

Habits.—It is to the present race that the Chinese apply the name of
four-eyed deer, on account of its power of everting the large face-glands.
Mr. Swinhoe states that “‘in the central ranges of Formosa, near Mount
Morrison, this brown deer is very common; and on a visit I paid to the
wild tribes of these parts in February 1866, I found them hunting the deer
with dogs. A place is cleared in the forest, where a party of men hide
armed with matchlocks ; the dogs yelp after the deer and drive them into
the open, where the hidden sportsmen get easy shots at them.” There
appears to be nothing recorded as to the breeding habits.

d. SZECHUAN RAcE—CERVUS UNICOLOR DEJEANI

Rusa dejeani, Pousargues, Bull. Mus. Paris, 1896, no. 1, p. 2.

Cervus dejeant, Ward, Records of Big Game, p. 22 (1896).

Characters.—Described as being very similar to the Formosan race, but
as large as the Indian sambar ; the colour being a more sombre brown than
in the latter, and the tail longer and more bushy. It is stated to be in all
probability only a larger continental form of the Formosan sambar, and
would accordingly appear indistinguishable from the Malayan sambar,
with which it is not compared by its describer. In the type and only
known specimen, which is preserved in the Paris Museum, the length of
the antlers is 304 inches, and the basai circumference 534 inches. As
already said, it is highly probable that this and the two preceding forms
should be classed as one sub-species.

Distribution —The province of Szechuan, North-Western China, about
30° north latitude. The distribution is interesting, as being the most
northerly recorded for the sambar, or indeed for any member of the
rusine group. The habitat serves in some degree to connect the dis-
tributional area of the Malayan with that of the Formosan race ; and it is
probable that this race will be found to intergrade with the former in the
Yunnan district.

Luzon Sambar PS 7

e. Luzon RAcE—CERVUS UNICOLOR PHILIPPINUS

Cervus mariannus, Desmarest, Mammalogie, vol. 11. p. 436 (1822);
Brooke: Prac. Zool. Sec.'1877, p. 53, 1878, p. 901.

Cervus (Stylocerus) philippinus, FH. Smith, in Griffith’s Animal Kingdom,
VOlPiIVs (Ds h47)¥e"P. -319) (1827).

Cervus (Rusa) martanus, H. Smith, of. cit. pp. 304, 311 (1827).

Cervus philippinus, Fischer, Synopsis Mamm. p. 622 (1829) ; Brooke,
Proc. Zool. Soc. 1877, p. 51, 1878, p. gol.

Cervus (Hippelaphus) philippinus, Sundevall, K. Svenska Vet. Ak. Hand.
for 1844, p. 179 (1846).

Cervus (Hippelaphus) marianus, Sundevall, op. cit. p. 180 (1846).

Rusa philippinus, Gray, Knowsley Menagerie, p. 63 (1850), Cat. Ungulata
Bra. Mus, p. 211 (1852).

Rusa paradoxa, Brehm, Zool. Garten Hamburg, 1864, p. 11.

Rusa mariannus, Gray, Cat. Ruminants Brit. Mus. p. 78 (1872).

Rusa marianna, Fitzinger, SB. Ak. Wien, vol. Ixviill. part 1. p. 355
(78373); lex: part 1. p. 304 (1874).

Rusa philippina, Fitzinger, op. cit. xviii. p. 355, Ixx. p. 306 (1874).

Ussa marianus, Heude, Mém. hist. nat. emp. Chinois, vol. 11. p. 40 (1888).

Cervus marianus, Ward, Records of Big Game, p. 9 (1896).

Characters.—Nearly allied to the two last. Height at shoulder about
28 inches; general build stout and massive, with the hind-quarters not
specially elevated, and the form that of a small Malayan sambar. General
colour of pelage rich ruddy brown, darkest on the back and lightest on the
neck ; forehead and cheek rufous fawn; a blackish streak starting from
over each eye to form a median line down the face, which is separated by
a pale band of fawn from a narrow moustache-like mark on the muzzle ;
under-parts uniformly brown ; metatarsal gland forming a rufous spot much
lighter than the rest of the leg ; ears moderate, covered externally with
short, close-set hairs. Antlers nearly similar to those of the Malayan race,
being very massive, nearly straight, with a very long brow-tine, and the
inner tine of the terminal fork markedly shorter than the outer one. Skull
rather elevated in the nasal region, and the lachrymal vacuity large and
triangular.

In a skull from the Marianne Islands in the British Museum, one of

158 Rusine Group

the antlers of which is figured by Sir Victor Brooke on page 56 of the
Proc. Zool. Soc. for 1877, the length from the occiput to the tip of the
nasals is 10.5 inches, and the length of the antlers along the -outer curve
18 inches. A Luzon skull in the Museum is of approximately the same
size, with similar antlers.

The Luzon sambar is a very rare animal in collections, the British
Museum possessing only a very few skulls, with antlers. The type of
C. philippinus is preserved in the Paris Museum. Judging from the figure
given by Sir Victor Brooke in his above-quoted paper of 1877 and from the
skulls in the British Museum, in the shortness of the face and the shape of
the antlers it appears to be very similar to the Formosan sambar, but it is
a shorter, less “leggy” animal, with the distinctive moustache-like mark-
ings on the upper lip. That it is only a race of the sambar, I am fully
convinced.

The so-called Marianne deer (C. mariannus) was considered by Sir Victor
Brooke to be in all probability identical with this form ; and the specimens
in the British Museum leave no reasonable doubt in my mind that this view
of the case is the true one. Now the Marianne or Ladrone Islands are of
very small size, and situated more than twenty degrees east of Luzon, very
nearly in the longitude of New Britain ; and it seems inconceivable that
any species of deer could naturally be found in islands of such a type situated
almost in the heart of Polynesia. Sir Victor Brooke has indeed suggested
that the Marianne sambar was introduced from Luzon by the Malays, and
this explanation must, I think, receive definite acceptation. Unfortunately,
the name C. mariannus antedates C. philippinus, but as the animal seems not
to be native to the Mariannes, I think the latter name must be adopted.
There is of course the possibility that even the Philippine race may be an
introduced variety of the Malayan sambar.

Distribution.—The island of Luzon, at the northern extremity of the

Philippine group ; introduced into the Marianne Islands.

ff. BasitAn RAcE—CERVUS UNICOLOR NIGRICANS

Cervus nigricans, Brooke, Proc. Zool. Soc. 1877, p. 57, 1878, p. 902.
Ussa nigricans, Heude, Mém. hist. nat. emp. Chinois, vol. 1. p. 31

(1888).

Basilan Sambar 159

Cervus steertt, Elliot, Publ. Field Columbian Mus.—Zool. vol. i. p. 72
(1896), wid. p. 157 (1897).

Characters —Smaller than the last, the height at the shoulder being
from about 24 to 26 inches; the general build much more slender, with
the hind-quarters much higher than the withers. General colour of
pelage blackish brown, slightly tinged with rufous, the face, neck, and
shoulders being almost black ; no moustache-like markings on the muzzle ;
chin and under-parts and the inner surfaces of the thighs varying from
dirty white to whitish brown, the white showing most on the front of the
thighs ; metatarsal gland generally not distinctly seen ; ears oval, relatively
small, and almost naked externally. Antlers moderately stout, somewhat
curved forwards, with a very short brow-tine, and the hinder tine of the
terminal fork not much shorter than the front one. Skull much depressed
in the nasal region, and the nasal bones in fully adult individuals so much
expanded as almost or completely to obliterate the lachrymal vacuity.

In a skull in the British Museum collected in Basilan by Mr. A. H.
Everett, the length from the occiput to the extremity of the nasals is 9.5
inches, and that of the antlers measured along the outer curve 13.5 inches.

The species was founded upon a female from an unknown island in the
Philippines, in which the lower portions of the legs are perhaps rather
lighter than in other examples, and the metatarsal gland indistinctly visible
as a light patch.’ This specimen is in the British Museum. In the year
18go the Museum received from Mr. E. L. Moseley a small male sambar-
like deer which had been collected by Mr. J. B. Steere in Basilan, and
has been for some years mounted in the mammal gallery, and labelled
C. nigricans. From the comparatively small size of the antlers it is probably
immature ; and the only difference, apart from sex, between this and
the type specimen is that in the former the front and outer side of the
lower portion of the legs are slightly darker, and show no distinct light
gland-patch. As mounted, it measures 254 inches at the shoulder, and 27
inches at the rump. In its relatively high rump it accords with Sir
Victor Brooke’s “crouching aguti-like carriage” of the type specimen,
and from actual comparison with the latter there seems no doubt that both
belong to the same kind of animal, the nearly naked ears being a very con-
spicuous common feature. The antlers are strongly curved. Two skulls

1 Both the legs and the gland-patch are made too light in Sir Victor Brooke’s plate.

160 Rusine Group

in the Museum collected by Mr. A. H. Everett in Basilan evidently belong
to older animals, having the antlers larger, although of the same type.

In 1896 Mr. D. G. Elliot described a small deer from Basilan collected
by Mr. Steere, by whom the British Museum mounted specimen was also
obtained. The author states that “the specimen serving as the type, and
which is the only one I have seen, is a fully adult male, and the horns, for
their size, are heavy and rough. In shape they are very different from
those of C. philippinus, the brow-antler being quite short, diverging rapidly
to a point, and joining the beam at a sharp angle. The inner tine is of the
same shape and about the same size as the brow-antler, and bends backwards
and very slightly inwards. The burr is rather small, and but slightly larger
than the beam in circumference. The latter is thick and straight, with
little or no taper, the outer tine, which curves slightly inward, being simply
the natural prolongation of the beam to a point. It is widest at the forking
of the inner and outer tines.

“From C. mgricans, the present animal differs in the colour of the coat,
which is much brighter than that of Sir Victor Brooke’s species, and also
by the form of the skull.”

The author then gives details of the points of difference between the
skull of his specimen and that of the typical female of the present form
figured by Sir Victor Brooke, remarking that these are apparently too great
to be due to sex alone.

In a subsequent communication Mr. Elliot adds the following :—“ Last
autumn, having had an opportunity to examine the type of C. phi/ippinus in
the Natural History Museum in the Jardin des Plantes, Paris, I ascertained
that my new species was hardly comparable with it in any way save a
similarity in the shape of the antlers; C. steer is not much over half the
size of C. philippinus, and differs also in the colour of the coat. In fact, it
would seem to belong to quite another section of the group than that
represented by C. philippinus. Besides the type, there were several other
examples of the species in the Museum, but all of them, even the young
animals, were much larger than the type of C. steer. It would seem,
therefore, that we must look for its near ally in some other species of
Philippine deer, as yet unknown to me.” It is also stated that in the
Basilan sambar the metatarsal gland is not different in colour from the hair
of the rest of the leg, that there are no dark moustache-like markings on

Basilan Sambar 161

the upper lip, no dark streak down the back, and that the tail is dark all
round.

Although the antlers appear somewhat straighter, I have no doubt,
from the figure of the head, that the type of C. steerii is identical with the
British Museum mounted specimen referred to C. nigricans, and since I
cannot distinguish the latter from the type of that species, I have no hesita-
tion in pronouncing C. steer to be a synonym of C. nigricans. Indeed, all
the specimens from Basilan agree in their small size and short brow-tine.
Mr. Elliot states that his specimen measures 1 foot 104 inches (224 inches)
at the shoulder, and 2 feet 84 inches at the withers, but the latter must
obviously be a misprint for 2 feet 34 inches (274 inches) ; and this relatively
high elevation of the hind-quarters is, as already said, a special feature in
the description of the type of C. mgricans. The alleged differences in the
skull I consider due to sex and age.

It is with some little hesitation that I include this tiny little deer as a
sub-species of sambar, but since it has the same type of antlers and similarly
coloured hair, it seems preferable to regard it as a very diminutive and
short-eared race of that very variable species.

Distribution.—The island of Basilan, situated near the southern extremity
of the Philippine group.

It is interesting to note that as Basilan is one of the smallest of the
Philippine Islands, so the Basilan sambar is the most diminutive of all the
deer to which the name sambar can properly be applied, it being a well-
known fact that the larger species of mammals gradually diminish in size
with the shrinkage in the size of their habitat. Although the creature is
not larger than a roe, and has relatively smaller ears than its relatives of the
larger Oriental islands and continent, yet, especially as regards its antlers,
pelage, colour, and tail, it is essentially a sambar, and as such seems better
placed as a sub-species than as a species.

Habits.—Dr. Steere writes that the Basilan sambar ‘“ was found occupy-
ing the higher parts of the island, in a country of steep, rocky ridges,
covered thickly with timber and thick undergrowth. In the valleys were
shallow streams. I took it rather for a mountaineer than a swamp-inhabiter,
though the whole country was dripping with moisture at the time of my
visit, and the higher lands in the Philippines are always the swampiest. . .

I noticed that all the deer procured in Basilan were closely alike in colour,
Y

162 Rusine Group

size, and character of horns. . . . I have an impression that the same
differentiation of species of Cervus exists in the Philippines as I found so
prominent in many genera of birds.” All these traits are essentially those
of a sambar. Its crouching, skulking gait and elevated hind-quarters have
been already mentioned.

2. Tue Rusa, oR JAVAN SAMBAR—CERVUS HIPPELAPHUS

Cervus hippelaphus, Cuvier, Ossemens Fossiles, ed. 3, vol. iv. p. 40 (1825) ;
Brooke, Proc. Zool. Soc. 1878, p. 903; W. L. Sclater, Cat. Mamm. Ind.
Mus. part ii. p. 179 (1891).

Cervus (Rusa) hippelaphus, H. Smith, in Griffith’s Animal Kingdom, vol.
LV. Pps 105, W.up. 300. {1927).

Cervus russa, Muller and Schlegel, Verh. Ges. Neder/. Zool. p. 217, plate
xliv (1839-44).

Rusa hippelaphus, Gray, List Mamm. Brit. Mus. p. 179 (1843), Cat.
Ungulata Brit. Mus. p. 209 (1852), Cat. Ruminants Brit. Mus. p. 77 (1872) ;
Fitzinger, SB. Ak. Wien, vol._\xviii. patt i. p. 355 (1873),-lxx part a
p- 312 (1874).

Cervus (Hippelaphus) hippelaphus, Sundevall, K. Svenska Vet. Ak. Hand.
for 1844, p. 178 (1846).

Cervus tunjac, Blyth, Cat. Ind. Mus. p. 151 (1863).

Characters.—General form, pelage, and coloration sambar-like, but the
ears never very large, the tail thin, the hairs on the back banded with
differently-coloured rings, and the under-parts, chin, and inner sides of
thighs and buttocks more or less distinctly whitish. Size medium. Antlers
comparatively slender and only moderately rugose, with the brow-tine of
medium or short length, and making a large acute angle with the beam ;
the hinder or inner tine of the terminal fork much longer than the front
or outer one, and forming the continuation of the beam, from the front or
front-outer surface of which the front tine arises as an offshoot ; the two
antlers enclosing a more or less distinctly lyrate space. Young uniformly
coloured.

Although included by Mr. Blanford among the synonyms of the sambar,
the rusa, or Javan sambar, is undoubtedly entitled to rank as a separate
species. Not only is there the marked difference in the form of the antlers,

Tur DEER or Att Lanps. PuateE Xi].

MOLUCCAN RUSA.

Published by Rowland Ward, Ltd.

Javan Rusa 163

as also the ringed hairs of the back and certain peculiarities of coloration,
but the fauna of Java differs so remarkably from that of the other Malayan
Islands and Peninsula, that on this ground alone there would be very strong
reason for separating the rusa as a species. Although used by Horsfield
and others for Indian varieties of the sambar,' the name /ippelaphus has by

Fic. 41.—Skull and Antlers of Javan Rusa. From a specimen in the British Museum.

most other writers been restricted to the Javan animal. Whether it was
applied by Cuvier to Javan specimens, I have no means of knowing, but
have retained it in the usually accepted sense. In the case of specimens of
which the locality is unknown, I find some difficulty in assigning them to
their proper race, and a good series of examples with authenticated localities

1 See Blanford, Fauna Brit. India—Mamm. p. 545.

164 Rusine Group

is essential before finality can be reached in this respect. Sir Victor Brooke
speaks of the various races as differing mainly in point of size ; but Fitzinger
alludes to colour differences and the varying length of the hair on the neck
and tail ; and examples living at Woburn display considerable differences in
the former respect.

Distribution.

Mauritius.

Java, Timor, Celebes, and the Moluccas, introduced into

a. JAVAN Racr—CERVUS HIPPELAPHUS TYPICUS

Characters.—Smaller than the Indian sambar, and of the approximate
size of a red deer. Head of moderate length, with the facial profile nearly
straight ; throat and neck of males with a well-developed mane ; ear broad,
and less than half the length of the head ; tail only slightly longer than the
ear, thinner than in the Malayan sambar, and ending in a tuft of thick
coarse hairs. Antlers as described above, a pair in the British Museum
having the front tine of the terminal fork nearly in the same plane as the
hinder one. General colour of the upper-parts in summer pelage dark,
grizzled, ochry-brown with a tinge of red, darker on the hind-quarters and
thighs than elsewhere ; front of neck, chest, and under-parts varying from
dirty white to brownish gray, and a dark reddish brown longitudinal streak
on the front of the chest ; flanks shot with rusty brown, and frequently
with a patch of that colour; inner side and lower part of legs, as well as
inner side of buttocks, dirty whitish ; chin, lips, and under surface of lower
jaw whitish, with a brownish spot on the lower lip at the angle of the
mouth ; inner surface of ear also whitish; base of tail yellowish brown
above and dirty white beneath, at or near the tip uniformly blackish brown.
In winter the general colour of the upper-parts is more grayish brown ; the
under-parts and inner surfaces of the upper portion of the fore-legs, thighs,
and buttocks dirty yellowish white ; the tip of the lower jaw, the border of
the upper lip, and the neighbourhood of the nose white ; a blackish spot
beneath the angle of the mouth, and often a brownish band round the
muzzle. In the female the streaks on the chest and the tail are somewhat
lighter. The antlers are relatively long and stout.

Such is Fitzinger’s description of this species, which accords fairly well
with specimens mentioned under the head of the next form. The British
Museum has no skins, two from Hainan labelled C. Aippelaphus being

Javan Rusa 165

apparently referable to the Malayan sambar. The species has not been
exhibited in the London Zoological Gardens since the year 1877.

Two stags apparently belonging to the present race now living in the
menagerie at Woburn Abbey differ from the description given above in
that the colour of the winter pelage is reddish tawny, with no pure white
on the under-parts. As they have the large antlers characteristic of the race,
this variation in colour may be analogous to that occurring in the larger
races of sambar. As shown in the accompanying photogravure, they have

SPs ff

Fic. 42.—Javan Rusa, from a Male at Woburn Abbey. Photographed by the Duchess of Bedford.

a well-developed mane on the neck and throat, and a dark terminal tail-tuft.
In the stag figured by Miller the general colour is similar to that of the
specimen of the Moluccan race represented in plate xii. ‘“ Rusa,” it may
be observed, is the Malay term for all deer, and is consequently applied by
the natives alike to the Malayan sambar and the present form, to which
latter it may conveniently be restricted in natural history.
Distribution.—Java, introduced into Mauritius, and, it is said, Borneo.
As indicating the extent to which the Malays have introduced deer into the
various islands they frequent, it is interesting to note that on Horsburgh

166 Rusine Group

Island, forming one of the Cocos-Keeling group, lying east of Northern
Sumatra, Mr. W. H. Forbes! mentions the existence of a herd which are
“a cross between the Javan rusa (Cervus hippelaphus) and the darker

”
.

Sumatra species (Cervus equinus)

6. Motuccan RaceE—CERVUS HIPPELAPHUS MOLUCCENSIS

Cervus moluccensis, Quoy and Gaimard, Voyage Astrolabe—Zool. vol. i.
p- 133, plate xxiv (1830) ; Eydoux and Gervais, Mag. Zoo/. vol. vi. p. 26
(1836) ; Brooke, Proc. Zool. Soc. 1878, p. go4.; Ward, Records of Big Game,
p. 12 (1896).

Cervus russa moluccensis, Muller and Schlegel, Verh. Neder/. Zool. p. 212
(1839-44).

Cervus (Hippelaphus) moluccensis, Sundevall, K. Svenska Vet. Ak. Hand.
for 1844, p. 179 (1846).

Rusa moluccensis, Gray, Knowsley Menagerie, p. 62 (1850), Cat. Ungulata
Brit. Mus. p. 209 (1852), Cat. Ruminants Brit. Mus. p. 77 (1872) ;
Fitzinger, SB.*_4k. Wien, vol. Vxwui. part 1. “p. 935401673) ec ance
p- 320 (1874).

Hippelaphus moluccensis, Heude, Mém. hist. nat. emp. Chinois, vol. iii.
p- 94 (1896).

Plate XII

Characters.—According to Fitzinger, the Moluccan rusa may be dis-
tinguished from the Javan race by its inferior size, and especially the want
of a distinct mane on the neck of the male, the tail having no distinct
terminal tuft, and the hair on the neck being short. The antlers are
relatively small, the build is low and stout, and the head large and thick.

The British Museum possesses the mounted skin of a male rusa formerly
living at Woburn Abbey, and presented by the Duke of Bedford, which
may be provisionally assigned to this race, although, as its place of origin is
uncertain, the determination may be open to doubt. As mounted, it stands
3 feet at the shoulder ; and is in the winter pelage. On the back the hairs
are distinctly marked with alternate dark and light rings, giving the speckled
appearance to this part of the pelage characteristic of the species. The

1 A Naturalis?s Wanderings in the Eastern Archipelago, p. 31.

Moluccan Rusa 167

general colour of the upper-parts is speckled rufous yellowish brown,
passing into uniform sambar-like brown on the flanks, where the hairs are
not ringed, and speckling is consequently absent. The face is short ; the
ears are of moderate size, and sparsely haired externally, where they are
white on the outer side of the base. The inner surface of the ears, a line
on each side of the nostrils, the lower lip, chin, under-parts, and the inside
of the thighs and buttocks are white or whitish ; the lower portion of the
legs being whitish brown. The antlers differ from the pair belonging to

Fic. 43.—Moluccan Rusa. From a photograph by the Duchess of Bedford.

the Javan race in the Museum, figured on page 163, by the circumstance that
the front tine of the terminal fork arises markedly from the outer side of the
beam, instead of from the front ; but I have no means of knowing how far
this difference is constant.

Distribution.—Celebes and the islands of Boru, Batchian, and Amboina
in the Molucca group ; all these islands lying between Borneo and New
Guinea, and having a fauna with a certain number of Australian types.

Sir Victor Brooke writes as follows concerning the occurrence of the
rusa in these islands and Timor :—“ Deer resembling Cervus hippelaphus in

168 Rusine Group

every particular excepting size are found in many of the islands in the
Indo-Malayan and Austro-Malayan regions. How far these are the result
of natural distribution, or how far their occurrence is attributable to
artificial transportation, is now most difficult to decide. The best
authorities on the fauna of the East Indies, Wallace, Schlegel, and Miller,
are inclined to lean towards the latter alternative ; and it is, I think, certain
that the introduction of many of the larger mammals (e.g. monkeys, pigs,
and deer) into the islands of the Austro-Malayan region has been affected
by the Malays, who, according to Wallace, are much given to taming
animals and conveying them from island to island. At the same time, the
existence of a species of Fe//s peculiar to Timor, and of the singular Anca
depressicornis in Celebes, seems to me to necessitate caution in accepting
this solution as of universal application. I shall, therefore, for the present
keep the references to Cervus timoriensis of Timor and C. moluccensis
distinct.”

Mr. Wallace! himself, who is much more confident about the intro-
duction by human agency, writes as follows :—‘‘The only Moluccan
ruminant is a deer, which was once supposed to be a distinct species, but
is now generally considered to be a slight variety of the Rusa /hippelaphus
of Java. Deer are often tamed and petted, and their flesh is so much
esteemed by all Malays, that it is very natural they should endeavour to
introduce them into the remote islands in which they settled, and where
luxuriant forests seem so well adapted for their subsistence.”

The question is one of extreme difficulty, to which there seems no
possibility of giving a definite answer. So far as Celebes is concerned, the
occurrence of the anoa and the babirusa renders it quite probable that a
deer might also have obtained an entrance into the island. ‘The Moluccas
are, however, more essentially Australian ; but if the pigs found in Ceram
and New Guinea are really indigenous, it is difficult to say that the deer
might not be so likewise. On the other hand, recent researches tend to
show that both Celebes and Timor belong to the Oriental, instead of the
Australian, region ; the deep channel running to the east, instead of to the
west of these islands. And this indicates a considerable probability that
these deer may be indigenous.

Habits—In Batchian Dr. Guillemard? writes that ‘‘the common

The Malay Archipelago, p. 300. 2 Cruise of the * Marchesa,’ p. 357.

Moluccan Rusa 169

Malayan deer was numerous in the forest and plantations of this part of the
island. It furnishes a permanent livelihood to a tribe of Gilolo Alfuros,
who have been settled in Batchian for many generations. Living for the
most part in the hills, they kill and smoke the deer, and bring the meat
into the villages for sale. We were fortunate enough to assist at one of
their hunts, in which no other weapon but the spear is used. The side of
_a large ravine which had been partially cleared, and presented a confused

Fic. 44.—Antlers of a Moluccan Rusa in four successive years. From a photograph by the
Duchess of Bedford.

jumble of fallen trees and low brushwood, was assigned to us as our post,
and from the extensive view it commanded we were able later in the day
to watch one run almost from start to finish, although at first the sport
appeared to be successful in every direction but our own. At length a
stag broke covert about 500 yards above us, and descended the slopes of
the ravine, but shortly afterwards turned and made for the forest again.
He was met by some of the hunters and driven back, but the dogs were
now in full cry, and pressed him hard, the hunters meanwhile racing at

Z

170 Rusine Group

their utmost speed above, in order to prevent his regaining the jungle.
He now altered his direction, and turned down once more towards us, but
the fallen trees were so thick that the dogs gained rapidly on him. He
made one more effort for his life by doubling, but it was too late, and in
another moment the dogs and hunters had fairly run him down.”

From this account it would appear that the ground frequented by the
Moluccan rusa is of the same general nature as that favoured by the Indian
sambar.

c. TIMORESE RACE—CERVUS HIPPELAPHUS TIMORIENSIS

Cervus timorensis, de Blainville, fourn. Physique, vol. cxiv. p. 267 (1822).

Cervus peronu, Cuvier, Ossemens Fossiles, ed. 3, vol. iv. p. 46 (1825).

Cervus (Rusa) peroni, H. Smith, in Griffith’s Amal Kingdom, vol. iv.
p. T14e Vops Bla a27,)-

Cervus russa timoriensis, Miller and Schlegel, Verh. Nederl. Zool. p. 212
(1839-44).

Cervus (Hippelaphus) peroni, Sundevall, K. Svenska Vet. Ak. Hand!. for
1844, p. 179 (1846).

Rusa peronu, Gray, Knowsley Menagerie, p. 63 (1850), Cat. Ungulata
Brit. Mus. p.. 211 (1852), Cat. Ruminants Bri. Mis, p. 75-0372
Fitzinger, SB. 4k. Wien, vol. \xvni. parti. p. 355 (1873), xx pane
p. 317 (1874).

Cervus timortensis, Brooke, Proc. Zool. Soc. 1878, p. 903.

Hippelaphus timortensis, Heude, Mém. hist. nat. emp. Chinois, vol. i.
p. 51 (1896),

Characters. —In size about the same as the preceding race, but
distinguished by the thicker hair on the neck, the more distinctly tufted
tail, certain differences in coloration, especially on the face and rump, and
the wider antlers. Fitzinger describes the coloration as follows :—The fore-
head is gray, the face and the eyebrows are dark blackish brown, the neck,
the upper part of the flanks and the greater portion of the chest being of
the same tint, the dark brown of the chest forming a long streak between
the legs ; the middle of the back is almost black ; the under-parts and inner
surface of the thighs are brownish or ochry yellow, passing into dirty
white on the hinder portion of the abdomen; a band above the hoofs

Timorese Rusa GAs

being also of the same colour ; the lips and inner surface of the ear are
white, as is also the inner surface of the buttocks ; the tail-tuft being dark
blackish-brown. In the males a whitish streak runs from above the eye
across the cheek to the side of the neck.

This deer has only once been exhibited in the London Zoological
Gardens, a pair having been presented by Captain L. Brayley in 1864.

Fic. 45.—Undetermined Rusine Stag at Woburn Abbey. From a photograph by the
Duchess of Bedford.

Distribution.—The islands of Timor, Semao, and Kambing, forming the
eastern extremity of the Sumatra-Java line. The observations with regard
to a possible introduction by man given under the heading of the last race,

apply equally in this case.

Undetermined Specimens

Here may be mentioned a pair of small dark-coloured rusine deer
living in the menagerie at Woburn Abbey, whose antlers, as shown in the
accompanying photograph, differ markedly from normal specimens of any

£72 Rusine Group

of the races of rusa or sambar. They have four points ; and in the figured
example recall to a certain extent those of the rucervine group. The
general similarity of these appendages in the two stags is decidedly against
abnormality, although they may possibly be hybrids. They are certainly
members of the present group; but nothing is known as to their place of
origin. The pedicles of the antlers were longer when young than at the
time when the figure was taken, and it is possible these deer may prove to
be the adult of C. culionensts.

3. THe PuivippineE SporTrep DEER—CERVUS ALFREDI

Cervus alfredt, Sclater, Proc. Zool. Soc. 1870, p. 381 ; Brooke, sid. 1877,
p. 59, 1878, p. go2; Meyer, bid. 1879, p. 666.

Axis alfredi, Gray, Cat. Ruminants Brit. Mus. p. 80 (1872).

Melanaxis alfredi, Heude, Mém. hist. nat. emp. Chinois, vol. ii. p. 47
(1888).

Characters.—-Size relatively small, the height at the shoulder being
about 28 inches; build long and low; face rather long. Hair less coarse
and shaggy than in any of the preceding forms. Ground-colour of pelage
of upper-parts of adult dark blackish brown, frequently with a deep black
line along the middle of the back ; the whole body marked with a number
of distinct whitish spots, forming a regular row on each side of the back,
but less regularly distributed elsewhere ; under-parts, chin, lower lip, inner
surface of buttocks and of upper portion of legs, as well as the front of the
thighs, white ; young also spotted. Antlers supported on short pedicles ;
apparently of the general type of those of the Malayan sambar, but re-
latively smaller, and with a shorter brow-tine ; good specimens are,
however, not known. Ears short and rounded, about one-third the length
of the head, almost naked behind ; face-gland fairly large ; no mane on
neck ; tail rather short and thinly haired, with some white on the lower
surface.

This very handsomely coloured little deer was first made known from a
living male specimen presented to the Zoological Society in 1870, of which
the skin is now preserved in the British Museum. A female was received
by the Society in 1871, and its skin is now mounted in the same collection.
The male first bred with the type female of the Basilan sambar, and subse-

Calamianes Deer 173

quently with its own species. Beyond the progeny of those three animals,
no other specimens have been exhibited in the Society’s gardens. In the
female skin in the British Museum the black line down the middle of the
back is very conspicuous, but it is not noticeable in that of the male. Sir
Victor Brooke regarded this species as closely allied to the Basilan sambar,
which it resembles in its small, bare ears and its flattened skull. The tail
and pelage are, however, quite unlike the sambar type, and the species
should apparently be regarded as indicating a distinct modification of the
group, not improbably related to the Indian spotted deer.

Distribution —The islands of Samar and Leyte, on the eastern side of the
centre of the Philippine group.’

4. THE CALAMIANES DEER—CERVUS CULIONENSIS

Cervus sp., Elliot, Puél. Field Columbian Mus.—Zool. vol. 1. p. 68
(1896).

Cervus culionenis, Elliot, op. cit. p. 157 (1897).

Characters—A small uniformly brown deer, differing from all the pre-
ceding forms by the great length of the pedicles supporting the antlers,
and the large relative size of the auditory bulle at the base of the skull ;
these two characters affiliating the species to the Bavian deer and the
hog-deer.

The original description states that the build of this deer is rather stout,
with the hind-quarters elevated, the head being slender, with a rather long and
pointed nose. The hair is somewhat coarse and stiff, especially on the
flanks, where it is longest. The ears are relatively long, slightly rounded,
and thickly haired externally ; and the tail is rather bushy. As the antlers
of the type were in velvet and not fully developed, their characters could
not be given. The general colour of the upper-parts is uniform cigar-brown,
with a tinge of ochry, but a black stripe of about an inch in width runs
from the shoulders to the root of the tail; the latter being brown above
and white beneath. The under-lip and chin are yellowish white, the chest,
shoulder, and much of the under-parts purplish brown, with long white
hairs intermingled, but the abdomen and inside of the thighs white, and

1 See Meyer, op. cit.

174 Rusine Group

the legs blackish brown, with a long narrowish white stripe on the front of
the hinder pair. In the female the general colour is paler.

I only knew this deer from the description, but in addition to the
peculiarities of the skull, the small size (height not given), thickly haired
ears, and dark dorsal stripe seem to differentiate it from all the foregoing
forms. And, from the analogy of the other island types, it seems primd
facie probable that the deer of the Calamianes would be a distinct form,
especially as the fauna of that island is very distinct from that of the
Philippines proper, and more nearly related to that of Palawan and
Borneo.

Distribution. —The island of Calamianes, or Culion, on the western side
of the centre of the Philippine group to the north of Palawan.

5. Tue Bavian DeER—CERVUS KUHLI

Cervus kuhlu, Miller and Schlegel, Verh. Ges. Nederl. Zool. p. 223
(1839-44).

Cervus (Hippelaphus) kuhli, Sundevall, K. Svenska Vet. Ak. Hand. for
1844, p. 179 (1846).

Rusa kuhli, Gray, List Osteol. Brit. Mus. p. 68 (1847), Cat. Ruminants
Brit. Mus. p. 79 (1872); Fitzinger, SB. 4k. Wren, vol. \xviil. part 1.
p. 355. (1973), [xx. patta. p; 200) (2874):

Cervus kuhli, Brooke, Proc. Zool. Soc. 1878, p. go2; Ward, Records of
Big Game, p. 11 (1896).

Characters—A small uniformly coloured brown species, apparently
nearly allied to the last, but without a dark line down the back.

Height at shoulder about 27 inches; build light and tall; face com-
paratively short. Hair moderately coarse and long ; the hairs on the back
ringed. General colour of adult pelage uniformly brown, apparently about
the same tint as that on the back of the hog-deer; no dark line on the
back, and the under-parts of a rather darker tint than in the latter ; young
also uniformly coloured. Antlers not much longer than the head,
supported on relatively long pedicles ; their general form similar to those
of the Malayan sambar, but much thinner and less rugose, and the brow-
tine very short. Skull differing from that of all the preceding forms, with

Hog- Deer B75

the exception of the Calamianes deer, by the great development of the
auditory bulle, which form marked projections on the inferior surface.
Ears small and pointed, thickly haired externally ; tail moderately long and
bushy ; face-gland small; metatarsal tuft only slightly lighter than the
rest of the leg; no mane on the neck.

The type specimen of this little deer is preserved in the Museum at
Leyden, and the British Museum possesses a mounted specimen, now much
faded and in bad condition, which was received in exchange from the
former establishment. Living examples have been exhibited years ago in
the Zoological Gardens at Amsterdam. The species appears to be in many
respects intermediate between the sambar and rusa on the one hand, and
the hog-deer on the other.

Distribution.—The small Bavian, or Bawean Islands, lying between
Borneo and Java.

6. THe Para, or Hoc-DEER—CERVUS PORCINUS

Cervus porcinus, Zimmermann, Species Zool. Geogr. Quadr. p. 552 (1777) ;
Brooke, Proc. Zool. Soc. 1878, p. go2; Lydekker, Cat. Foss. Mamm. Brit.
Mus. part i. p. 104 (1885) ; W. L. Sclater, Cat. Mamm. Ind. Mus. part ii.
p. 178 (1891); Blanford, Fawna Brit. India—Mamm. p. 549 (1891) ; Ward,
Records of Big Game, p. 10 (1896).

Cervus (Axis) porcinus, H. Smith, in Griffith’s Animal Kingdom, vol. iv.
Pang, V2 p.<gi2 (1827).

Cervus (Axis) pumilio, H. Smith, op. cit. pp. 120 and 313 (1827).

Cervus pumilio, Fischer, Synop. Mamm. p. 621 (1827).

Axis porcinus, Jardine, Naturalist’s Library — Mamm. vol. iii. p- 196
(1835); Jerdon, Mamm. India, p. 262 (1867) ; Sterndale, Mamm. India,
p- 508 (1884).

Cervus (Hyelaphus) porcinus, Sundevall, K. Svenska Vet. Ak. Hand!. for
1844, p. 181 (1846).

Hyelaphus porcinus, Gray, List Osteol. Brit. Mus. p. 67 (1847), Cat.
Ungulata Brit. Mus. p. 215 (1852), Cat. Ruminants Brit. Mus. p.79 (1 $72);
Fitzinger, SB. 4k. Wien, vol. Ixviii. part i. p. 354 (1873), Ixx. part 1.
P- 253 (1874).

176 Rusine Group

Axis oryzus, Kelaart, Prodr. Faun. Zeylan. p. 83 (1852).

oe porcinus pumilio, Fitzinger, SB. Ak. Wien, vol. \xx. part A,
p. 258 (1874).

Cervus minor, Sclater, List Anim. Zool. Gardens, p. 169 (1883), nec Axis
minor, Hodgson (1841).

Characters.—Allied to the preceding, but the antlers larger, the build
longer and lower, and the summer pelage of the adult, as well as that of
the young, spotted.

Height at shoulder ranging from about 25 to 29 inches ; build heavy
and low, with the legs short ; face comparatively short ; the hairs on the
back not ringed, but with pale tips. General
colour of adult pelage in winter rufous or yellow-
ish brown, with a somewhat speckled appearance
owing to the pale tips of the hairs ; under-parts
much darker ; in summer the upper-parts paler
and more or less spotted in the early part of the
season with pale brown or white, the spots some-
times limited to one or two rows on each side
of a dark stripe down the middle of the back ;
young fully spotted for about the first six
months. Antlers supported on long pedicles,
and of the same general type as those of the
preceding species—that is to say, with the
Big, 46, Head of Hoe a ainiden or inner, tine of the terminal fork the

(Rowland Ward, Records of
Big Game.) shorter, and the brow-tine relatively short—but
of greater proportional length, being consider-
ably longer than the head in fully adult individuals. Skull structurally
similar to that of the last, but much less compressed in front of the orbits,
and the latter less prominent. Ears rather large, well haired externally,
white internally ; tail rather long, whitish beneath ; face-gland small ;
metatarsal tuft slightly lighter than the rest of the leg ; no mane on the
neck or throat.

A pair of antlers of an immature hog-deer in the British Museum
are almost indistinguishable from those of an adult Bavian deer. The
following dimensions of the antlers are recorded by Mr. Rowland

Ward:

Hog-Deer t77

Length along Basal

Outer Curve. Circumference. ue: AGREE
21 34 rst Pegu
204 3 9 N.-W. Provinces
202 34 9 Nepal
19s 34 132 N.-W. Provinces
19+ 34 gs India
19t 34 163 Burma
19t 32 183 India
174 34 9h Nepal
174 i gt India
16e g 103 5
164 qe 144 Nepal

Doubts have been expressed as to whether the hog-deer is spotted in
the adult summer pelage, and it is probable that many individuals are never
so marked, and that in most or all the dappling disappears before the
assumption of the autumn coat. Mr. Blanford states that he has watched
the appearance of the spotted summer garb in several individuals for two
or three successive seasons in the Calcutta Zoological Gardens, and the
spotting has been very noticeable in specimens living in the park at Woburn
Abbey, one of which is shown in the accompanying figure. Mr. Sclater
regards the spotted hog-deer as a distinct species, under the name of
Cervus minor. According, however, to Mr. Blanford, Hodgson’s Axzs
minor is a synonym of the Indian spotted deer, and therefore if the spotted
hog-deer were a distinct species or sub-species, it would require a new name.
In Mr. Blanford’s description the under-parts of the hog-deer are stated to
be lighter in colour than the back, whereas, in the summer coat at least,
they are much darker ; this species and the preceding being almost the only
deer besides the wapiti in which this peculiar type of coloration obtains.

Distribution.
the Punjab to Assam, thence through Sylhet to Burma and Tenasserim.
Although it has been reported from the highlands of Central India and
Bombay, it is very doubtful if the species ranges to any extent into Penin-

India, throughout the Indo-Gangetic plain from Sind and

sular India, except that it may be found some distance up the larger

tributaries of the Ganges. It has been reported from Madras, but apparently

owing to the name hog-deer having been improperly applied to the Indian

chevrotain and muntjac. In Ceylon it has been introduced into certain
2A

178 Rusine Group

districts. An antler from the Plistocene deposits of the Narbada valley has
been tentatively assigned by myself to the para ; if the reference be correct,
the species formerly had a larger range.

Habits.—The para, as this little deer is called in Hindustani, is essentially
a dweller on open alluvial flats, where it generally prefers grass-clad plains,
although occasionally found among tall trees. Shunning the dense and lofty
grass-jungles which form the haunt of the Indian rhinoceros and buffalo,
the hog-deer selects localities where the grass and bushes attain but a

moderate height. In such spots it is to be met with abundantly, although

Fic. 47.—Male Hog-Deer in Summer Pelage. From a photograph by the Duchess of Bedford.
The light tint of the head and neck is due to the strong sunlight on these parts.

it is seldom that more than two or three are seen in company, as this
animal is not of a sociable nature, and never collects in herds after the
manner of its relative the spotted deer. Indeed, except during the pairing-
season, even individual males and females keep apart from one another and
lead a more or less completely solitary life. The pairing-season is in
September and October, and the fawns are born eight months after. April
is the usual month for shedding the antlers. In its movements the hog-deer
is not a particularly graceful animal, running in an awkward-looking manner,
with the head carried low. In captivity the bucks show great tameness,
and walk with the head well elevated somewhat after the manner of a

* aye my
“—s

=

Tue DeER or Att Lanps. Puate XIU

CHITAL, or, INDIAN SPOTTED DEER.

}ublished by Rowland. Ward. Lid

Chital 179

sambar, only with less aggressiveness, and without such constant dilata-
tion of the face-glands, which are much smaller than in the latter.
The species has bred both in the London Zoological Gardens and at
Woburn Abbey.

7. THE CHITAL, oR INDIAN SpoTTED DEER—CERVUS AXIS

Cervus axis, Erxleben, Syst. Regn. Anmmal. p. 312 (1777) ; Brooke, Proc.
Zool. Soc. 1878, p. 906; W. L. Sclater, Cat. Mamm. Ind. Mus. part ii.
p. 181 (1891); Blanford, Fauna Brit. India—Mamm. p. 546 (1891); Ward,
Records of Big Game, p. 18 (1896).

Cervus (Axis) axis, AH. Smith, in Grifhth’s Ammal Kingdom, vol. iv.
Paebh 7. %. po Ar2(1827).

Cervus axis ceylonensis, H. Smith, of. cit. vol. v. p. 786 (1827).

Cervus nudipalpebra, Ogilby; Proc. Zool. Soc. 1831, p. 136.

Axis major and minor, Hodgson, fourn. As. Soc. Bengal, vol. x. p. 941
(1841).

Axis maculata, Gray, List Mamm. Brit. Mus. p. 178 (1843), Cat. Ungulata
Brit. Mus. p. 212 (1852), Cat. Ruminants Brit. Mus. p. 80 (1872) ; Fitzinger,
SB. Ak. Wren, vol. xx. part 1. p. 264 (1874).

Cervus (Hippelaphus) axis, Sundevall, K. Svenska Vet. Ak. Handl. for 1844,
p- 180 (1846).

Axis maculatus, Jerdon, Mamm. India, p. 260 (1867) ; Sterndale, Mamm.
India, p. 506 (1884).

Axis maculata ceylonensis, Fitzinger, SB. Ak. Wien, vol. 1xx. part i. p. 269
(1874).

Axis nudipalpebra, Fitzinger, op. cit. p. 270 (1874).

Hyelaphus maculatus, Fitzinger, op. cit. 259 (1874).

Plate XIII

Characters —Distinguished from all the other members of the group,
except the Philippine spotted deer, by the pelage being profusely spotted
with white at all seasons and at all ages. Size medium, the height at the
shoulder varying from 36 to 38 inches; build light and slender, with the
legs long. Colour of pelage bright rufous fawn, spotted all over the body
with white ; a dark stripe running from the nape of the neck to the

180 Rusine Group

extremity of the tail, bordered along the back by one or two rows of white
spots ; the spots low down on the flanks sometimes blending into a line ;
the chin, upper portion of throat, inside of ears, the under-parts, inner
surface of limbs, and under side of the tail white ; head uniformly brownish,
darker on the face, with a blackish ring on the muzzle. Antlers supported
on short pedicles, long, slender, and moderately rugose ; the brow-tine long
and making nearly a right angle with the beam; the front, or outer, tine
of the terminal fork much longer than the hinder, or inner one, and forming
the continuation of the beam, from the inner side of which the hinder tine

Fic. 48.—Antlers of a Chital in six successive years. From a photograph by the Duchess of Bedford.

arises ; space enclosed by the two antlers more or less distinctly lyrate ;
sports, or irregular snags near the base of the brow-tine very frequently de-
veloped. Ears moderate ; tail long, pointed, and evenly haired throughout ;
face-gland not excessively large ; no mane on either the neck or the throat.

The skull differs from that of the hog-deer by the normal size of the
auditory bulla on its lower surface. A dark variety, to which the name of
Cervus nudipalpebra was given, is occasionally met with, and shows scarcely
any indications of spotting; but it has no apparent claim to rank as a
distinct sub-species. The largest individuals occur in Northern and Central
India, where the height of the stags commonly reaches to between 36 and

Chital 181

38 inches ; in Southern India the height is less, usually varying from 30
to 34 inches, although 36 has in one instance been recorded. It was to
the small South Indian form that Hodgson gave the name of Axis minor.
The shape of the antlers is exactly the reverse of that obtaining in the
rusa. Those of the present species may be described as consisting of a
brow-antler and a beam, the latter of which gives off an inner tine some-
where near the middle of its length ; whereas in the rusa the beam gives
off an outer tine. On the other hand, in all the varieties of the sambar
(C. unicolor) the beam may be described as more distinctly forked, although
there is great variation in the relative length of its constituent tines.
Ordinary specimens of chital antlers from North and Central India measure
about 30 inches, and any examples over this length may be considered
good ; in Southern India they are smaller. The following are some of the
largest specimens recorded by Mr. Rowland Ward :—

Cree Cieeeh reams! pee: Eeealy:
384 42 194 Central Provinces
37 44 243 :

372 4 24% P

ua 44 165 ?

364 43 19% 7

364 44 25 Berar

364 4 152 Nepal

36 “4 25h

353 44 184 P

353 48 23 ?

355 Ae 220 Western Ghats
35 44 North Kanara

A still finer head is shown in Fig. 5, p. 31.

The reasons for including this species in the rusine group, instead of
referring it to a separate group by itself, have already been mentioned.
From the retention of the spots at all ages and seasons, the Indian spotted
deer may be regarded as the most primitive and least specialised member
of the whole group; the long tail being probably also a primitive
character. Many writers have considered the Indian spotted deer as nearly
allied to the sikas ; but almost the only features common to the two are

182 Rusine Group

the spotted coat and the length of the tail, both these having doubtless
been inherited from the common early Pliocene ancestor. In the develop-
ment of a trez-tine to the antlers, as well as in the tendency to lose the
spots, and also in the black-bordered white caudal area, the sikas have
departed from the ancestral type apparently retained by the chital.

Fic. 49.—Skull and Antlers of Chital. From a specimen in the British Museum.
(Rowland Ward, Records of Big Game.)

Distribution—Almost the whole of India and Ceylon. Mr. Blanford
writes that this species ‘occurs at the base of the Himalayas, not, how-
ever, ascending the mountains beyond the lower spurs, from the neighbour-
hood of the Sutlej; to Nepal, but not in Sikhim. It is not found in the
Punjab plains, nor in Sind, and only to the eastward of Rajputana ; it is
wanting also in Assam and to the east of the Bay of Bengal, but is
common in the Sandarbans, apparently as far east as Mymensing, through-

Chital 183

out Bengal and Orissa, the North-West Provinces, Central India, Mysore,
Malabar, and Ceylon, in all suitable localities. It ascends the hills of
Southern India, in places to about 3500 or 4000 feet.”

Habits —This species being the only distinctly spotted deer inhabiting
India, it is natural that the natives of that country should have taken note
of that feature in naming the animal; the Hindustani name chital, or
chitra, meaning spotted, and being met with under the same or a nearly
similar form in the vernacular title of the spotted soft river-tortoises
(chitra), and the leopard (chita). This beautifully spotted coat renders
the chital one of the most lovely members of the whole deer tribe ; and it
is not a little remarkable that almost all the countries of the Old World
where deer are found have one spotted species or group, as if to remind us
of the beautiful coloration which was doubtless formerly common to the
entire family. In the Mediterranean countries we have the two species of
fallow deer, in China and Japan the sikas, and in India the chital. Not
only is the spotted deer one of the handsomest of all Indian mammals, but
it is likewise one of the most characteristic, being, as already mentioned,
unknown in the countries lying to the eastward of the Bay of Bengal, and
having certainly been an inhabitant of the peninsula since the time when
the latter was connected with Ceylon.

In marked contrast to the para, the chital is an essentially social species,
congregating in herds, which sometimes include hundreds of individuals,
and always at least one fully-antlered buck. Even in their native haunts
they are much less exclusively nocturnal than the sambar, and may fre-
quently be seen on the move for several hours after sunrise, and also some
time before sunset ; while when transported to English parks they will
feed at any hour. The neighbourhood of water is essential to the well-
being of this species, and if this be at hand, as well as covert for shelter,
chital may be found either on the open plains or in hilly ground. Their
partiality for water leads them to frequent spots where the foliage is of the
most luxuriant ; and no more beautiful sight can be witnessed in any part
of the world than a herd of Indian spotted deer feeding on an open grassy
glade dotted over with clumps of feathery bamboo by the side of a stream.
Bamboo-jungle, and covert of other bushes and trees in the neighbourhood
of water are, indeed, their favourite haunts ; and if the covert is sufficiently
dense and abundant, they display no marked aversion to the near neighbour-

184 Rusine Group

hood of human beings and their habitations. When alarmed, the hinds
are the first to make a move, racing off with a number of long flying leaps,
very much after the manner of black buck ; after a short interval spent in
gazing, they are followed by the lord of the herd. In feeding, they both
graze and browse ; and after drinking sometime between eight and ten in
the morning, they repose in the deepest shade accessible till the sinking
sun proclaims it to be again time for wandering. As might be expected
from their partiality for water, chital are excellent swimmers, and in the
Sandarbans of Lower Bengal they may frequently be seen passing from one

|

Fic. 50.—Male and Female Chital. From a photograph by the Duchess of Bedford.

island or strip of land to another. In addition to a scream of alarm, these
deer utter a peculiar half-barking cry. As is the case with the sambar,
the shedding of the antlers is very irregular, stags with these appendages
fully developed being met with at all times of the year. This implies
corresponding irregularity in the breeding-season and the birth of the
fawns, and although most of the latter appear to come into the world
during the Indian cold season, some are produced at almost all times of the
year. In spite of the warmth of its native home, the chital flourishes
excellently in England, none of the deer at Woburn Abbey appearing in
better condition than the herd of this species. Here the time of shedding

Chital 185

the antlers is very irregular, and the fawns may be dropped at any season,
a single doe not unfrequently having two fawns in the same year. A buck
born in December will usually shed his antlers in October, but one which
came into the world in July will throughout life drop them in May or
June.

An observer writing under the pseudonym of ‘“ Hawkeye” has given
such a graphic account of chital in their native haunts that although it
has been already quoted, it will bear another repetition. ‘‘ Deep in the
solitude of the woods,” he writes, “see now the dappled herd, and watch
the handsome buck as he roams here and there in the midst of his harem,
or, browsing amongst the bushes, exhibits his graceful antlers to the lurk-
ing foe, who by patient woodcraft has succeeded in approaching his un-
suspecting victim ; observe how proudly he holds himself, as some other
buck of less pretensions dares to approach the ladies of the group ; see how
he advances, as on tiptoe, all the hair of his body standing on end, and
with a thundering rush drives headlong away this bold intruder, and then
comes swaggering back! But, hark—a twig has broken! Suddenly the
buck wheels round, facing the quarter whence the sound proceeded. Look
at him now, and say, is he not a quarry worth the hunter’s notice ?

“With head erect, antlers thrown back, his white throat exposed, his
tail raised, his whole body gathered together, prepared to bound away
into the deep forest in the twinkling of an eye, he stands a splendid
specimen of the cervine tribe. . . . A doe suddenly gives that imper-
ceptible signal to which I have formerly alluded, and the next moment the
whole herd has dashed through the bamboo alleys, vanishing from sight—
a dappled hide now and again gleaming in the sunlight as its owner
scampers away to more distant haunts.”

IncERT#A SEDIS
1. Cervus lepidus

Cervus lepidus, Sundevall, K. Svenska Vet. Ak. Handl. for 1844, p. 180 (1846) ;
Brooke, Proc. Zool. Soc. 1878, p. 902.

Rusa lepida, Gray, Knowsley Menagerie, p. 63 (1850), Cat. Ungulata Brit. Mus.
p. 212 (1852); Fitzinger, SB. 4k. Wien, vol. |xx. part 1: p. 302 (1874).

Sir Victor Brooke observes that ‘‘ this is a very doubtful species. Some years ago

2B

186 Rusine Group

I saw the type in the Museum at Frankfort ; and it appeared to me to closely resemble
Cervus sika. Upon the occasion of my last visit to Frankfort, I sought in vain for
the specimen, and much fear that it has been destroyed.”

2. Cervus caspicus

Cervus caspicus, Brooke, Proc. Zool. Soc. 1874, p. 47, 1878, p. 909 ; Lydekker,
ibid. 1897, p. 38.

This presumed species was named on the evidence of a frontlet and antlers obtained
from the districts lying to the south-west of the Caspian. The type specimen, which
was figured in 1874, was originally in the collection of the late Sir Victor Brooke
but could not be discovered after his death. The antlers measure 26 inches in length
along the curve, and are three-tined. In his original description Sir Victor Brooke
referred the species to the rusine group, comparing it to C. unicolor and C. hipp-
elaphus ; but in his memoir of 1878 it was placed provisionally in the sikine group, with-
out any reference to the previous determination. The ground of this redetermination
seems to rest upon another antler from the Karun valley, in the Luristan district of
Persia, which may or may not belong to the same species as the type specimen.
With regard to the latter, it appears, so far as can be determined from the figure,
that the original determination was correct, and that the specimen is really rusine, the
interval between the brow-tine and the terminal fork being much greater than the
one between the brow-tine and the first fork of a sika’s antler. From subsequent
investigations, it is now nearly certain that neither a rusine nor a sikine deer
inhabits the district of Persia in question; and it appears highly probable that the
type specimen belonged to the Indian or one of the allied races of sambar, and was
imported into Persia.

3. Other Names

The following names have been applied by Heude (Mém. hist. nat. emp. Chinois)
to specimens, mostly skulls, belonging to the present group. With regard to these
I cannot do better than quote the following observations of Mr. D. G. Elliot," who
writes as follows :—‘ The paper is really of little assistance in determining the various
species, and the author appears to have seen a new one in almost every example he
procured, and unless one has access to the material in his possession, it will be
practically impossible to recognise the animals upon which he has bestowed so great
a list of names.” The list runs as follows, viz.—

1. Ussa gorrichanus, vol. ii. p. 21 (1888) Philippines.

2. ” p- 22 ” ”

1 Publ. Field Columbian Mus.—Zool, vol. i. p. 157 (1897).

», barandanus,

Heude’s Names

3. Ussa crassicornis, vol. 11. p. 23 (1888) Philippines.
4 5 francianus, Bo Ups 247 4, Ah
G3) wublanus, ANOS Ea? he ae »
6. ,, tuasoninus, ae pe 2s # t

q 5 Spatharius, ee AP 25)! vy, a

8 59 ramosianus, oe Pe 2ON.” 55 ie

9 5 @MOFOSIGNUS, 5, D. 27 45 re
te) 5 Mmacarianus, 5) 28 es -
IG = NCLOTLZ ANUS. «Ox APS 2B) ys “3
12. ,, garcianus, su AP 20: 5 a
13. 5, guidoteanus, 3 P29) ~ 55 i
I4. 4, vosarianus, po PegOr 5, “
WR IGT AtSIAnUS, 5, P.52T “, Fs
ROE MAGUIGTGIGHUS, 9... Dx 32 sy -
Ie es CAL ZANINUS,, 5. PB. 33 55 3
OG HUN ONASLQIUS: son Det 38 R .
HOs 45) longicUspLS, BP ONS i er :
TOE TUICTOAONLES. 6 3, =D. 34 45, =
Dies ss I LONZ AILS, Sr loer es ¥ 3
DREN ICLESOMIGNUS.. 55 DP. BO” ~'5 <.
Q. 4, Orachyceros, Se DOR 55 =
24. 4, corteanus, ae ee aan i
25. 4, verZzosanus, RMD ERIE) Ty -
26. ,, rubiginosus, RENO) 355 *
Di aRhEPOlItTanys. 7.) "Pp. 3O 5, Ms
DBanl esse CHT SOLTICLOS a Din QO) 55
29. 4, qguevaranus, SDA a8 *
30. Sambur curvecornis, vol. i. p. 42 (1888) Cochin-China.
aie Sn OULICOL UTS eon DNA Den 5, ‘3
Be 56 outreyanus, 50 Pp: 42 53
23. 5» planidens, Ce ap ras, «55 r
Ba. Se COLIMILETUNUS a = Daag 55 -
Bie Wee ComUaILerHnES..., P43 5, 3
36:0. 43°) “gnarids, en PAA. "55 3
205. 4g lemeanus, a Deen 55 53
Boe) bol. CGrarsianus. 4 plas 43 is
39. ye fOUleriianUs,. 3 Dp. 45 re ss
AOn es | Janaeis, es AG i} es
Atel 4, planiceps, pees > Fs, 3

188 Rucervine Group

42. Sambur officialis, vol. ii. p. 46 (1888) Cochin-China.

43. 4,' simontnas, “G5 pal; a

44. 4, brachyrhinus,,, p.46 ,, 4

45.9 64,9 verutes: oo pede? = S, -

46. Melanaxis masbatensis, vol. 11. p. 47 (1888) Masbate.

Flag fi * breviceps, sr Upp ees 5 is

48. - elegans, ee oe 5. Jteneo:

49. ‘ basilanensis, ,, pp. 49 ss Bastlan:

50. Hyelaphus calamianensis, vol. ii. p. 49 (1888) Calamianes.
si. ee annamiticus, >». PicSO» .5,4/) annem
52. Hippelaphus hamiltonianus, vol. iil. p. 49 (1896) Sundakan.
Ea: a macassaricus, ct PeshOe 430 -Vlacassar.
54. * menadensis, » pifOr4,. 6 Menade:
ae Pal floresianus, 3 PHO2 -.. Hieres.
56. " buruensts, 5 P2908) css, UEU:

We s hoevellianus, es: PEO 4a | aes 35

v. THE RuceERVINE Grovep—Sus-GEnus Rucervus

Recurvus, Hodgson, Ann. Mag. Nat. Hist. ser. 1, vol. i. p. 154 (1838).

Panolia, Gray, List Mamm. Brit. Mus. p. 180 (1843), Cat. Ungulata
Brit. Mus. p. 202 (1852), Cat. Ruminants Brit. Mus. p. 75 (1872);
Fitzinger, SB: Ak. Wien, vol. \xviii. part 1. p. 351: (1973), ixix. pare
Pp. 591 (1874).

Rucervus, Gray, List Osteol. Brit. Mus. p. 65 (1847), Cat. Ungulata
Brit. Mus. p. 303 (1852), Cat. Ruminants Brit. Mus. p. 75 (1872) ;
Fitzinger, SB. Ak. Wien, vol. \xvili. part i. p. 355 (1873), xx: part a.
p- 324 (1874) ; Brooke, Proc. Zool. Soc. 1878, p. 905.

Characters.—Antlers flattened or rounded, with both the bez (second)
and trez (third) tines wanting, the beam dichotomously forking, and one
or both branches again forked, so that the namber of tines is at least four,
and may be many more, the brow-tine forming either a right angle or a
continuous curve with the beam. Pelage of adult generally almost or
quite uniformly coloured, without a light caudal disk; young usually
spotted. Neck maned ; ears large ; face long ; tail short. Naked portion
of the muzzle of the rusine type, but decidedly narrower just above the
upper lip. The face-gland moderate, and not capable of complete eversion.

Swamp- Deer 189

Metatarsal gland small or wanting. Upper canines small ; upper molars
with a small additional column on the inner side. Size large.

The skull is of very much the same type as that of the preceding group,
but relatively narrower, with the auditory bulle on the under surface
only moderately inflated.

The group is evidently nearly allied to the rusine division, from
which the more specialised form of the antlers suggests that it is a late
development. Remains referred to this group have been described from
the later Tertiary deposits of India, but beyond this nothing is known as to
its paleontological history. Probably, however, it was developed in the
Oriental region, and from its absence from the Malayan Islands not
improbably at a date subsequent to the insulation of the latter.

Distribution—The mainland of the Oriental region, but unknown in
the islands with the exception of Hainan.

1. THE Swamp-DEER—CERVUS DUVAUCELI

Cervus duvaucelt, Cuvier, Ossemens Fossiles, ed. 3, vol. iv. p. 505 (1825).

Cervus bahraimja, Hodgson, Proc. Zool. Soc. 1834, p. 99.

Cervus elaphoides, Hodgson, “fourn. As. Soc. Bengal, vol. iv. p. 648
(1835).

Cervus (Recurvus) elaphoides, Hodgson, Ann. Mag. Nat. Hist. ser. 1, vol. i.
p- 154 (1838).

Cervus dimorphe, Hodgson, ‘fourn. As. Soc. Bengal, vol. xii. p. 807
(1843).

Axis (?) duvaucellu, Gray, List Mamm. Brit. Mus. p. 178 (1843).

Cervus (Hippelaphus) duvauceli, Sandevall, K. Svenska Vet. Ak. Hand.
for 1844, p. 178 (1846).

Rucervus duvaucellu, Gray, List Osteol. Brit. Mus. p. 65 (1847), Cat.
Ungulata Brit. Mus. p. 203 (1852), Cat. Ruminants Brit. Mus. p. 76 (1872) ;
Jerdon, Mamm. India, p. 254 (1867).

Rusa dimorpha, Gray, Knowsley Menagerie, p. 62 (1850); Fitzinger,
SB. Ak. Wien, vol. \xviii. part i. p. 355 (1873).

Cervus euceros, Gray, Knowsley Menagerie, p. 40 (1850).

Cervus eucladoceros, Falconer, Pal. Mem. vol. i. p. 587 (1868).

Cervus duvaucelh, Sclater, Trans. Zool. Soc. vol. vii. p. 346 (1871).

Igo Rucervine Group

Rucervus duvauceli, Fitzinger, SB. Ak. Wien, vol. Ixviii. part i. p. 356
(1873), lxx. part i. p. 324 (1874).

Axis dimorpha, Fitzinger, op. cit. Ixx. p. 272 (1874).

Cervus duvaucel, Brooke, Proc. Zool. Soc. 1878, p. 905; W. L. Sclater,
Cat. Mamm. Ind. Mus. part ii. p. 179 (1891); Blanford, Fauna Brit.
India—Mamm. p. 538 (1891) ; Ward, Records of Big Game, p. 13 (1896).

Rucervus duvaucell, Sterndale, Mamm. India, p. 510 (1884).

Plate XIV

Characters.—Height at shoulder from 3 feet 8 inches to 3 feet 10 inches ;
build stout and tall; hair moderately fine, and somewhat woolly ; muzzle
long and slender. Antlers smooth and
flattened, with a long brow-tine rising
almost at right angles to the beam ; above
the brow-tine the beam remains undivided
for about half its length, when it splits
into a regular fork, of which each branch
is usually again simply forked, although,
as in the specimen shown in Fig. 51, the
outer branch may be much longer than
the inner one, and bear three or more
tines ; small snags are frequently developed
on the upper surface of the brow-tine,
although sports at its junction with the
beam are seldom seen, and the brow-tine
is never forked. Metatarsal gland and tuft
wanting. General colour of pelage in
summer bright rufous brown, frequently,
or usually, with a broad brown line down
the middle of the back, bordered by a line

of white spots on each side, and more or

Fic. 51.—Side view of Frontlet and Antlers
of Swamp-Deer. From a specimen in
the British Museum. less faint traces of other spots ; the throat,

inner side of the thighs, and under-parts
white or whitish, and the lower surface of the tail invariably pure white ;
in winter the upper-parts yellowish brown, and the under-parts paler ; in
hinds the colour is lighter at all seasons, and the young are fully spotted

Tue Deer or Aut Lanps. PiatTE XIV.

SWAMP-DEER.

Published, by Rowland Ward Lia

Swamp-Deer IgI

with white. The ears are filled internally with long white hairs ; and
the naked portion of the muzzle is slaty.

The type specimen of Hodgson’s Cervus dimorphe has been identified
with this species by Messrs. Blanford and Thomas. The distinctive feature
of the antlers is the distinct angle formed by the junction of the brow-tine
with the beam, and the length of the undivided portion of the latter.
Any antlers over 30 inches in length may be regarded as good specimens.

The following are some of the largest measurements recorded by Mr.
Rowland Ward :—

Length along Number of

Cee onB Circumference. Tip to Tip. Widest inside. oe edit:
41 54 ? 6-6 Central Provinces
394 5 355 372 5-6 India
384 64 P 5 6-5 Central Provinces
38 : 43 : 6-6 is
37% 54 238 285 7-6 !

364 5 20 P 2 Mavalla District
36 5 294 33% 6-5 Nepal

354 St 224 ? 7-6 :

358 45 27 294 ; ?

342 52 384 ? 5-5 Central Provinces
345 Se 24% 285 6-5 Nepal

Distribution.—India, not extending to the eastward of the Bay of Bengal
or to Ceylon. In India the range extends along the foot of the Himalaya
from Upper Assam in the east to the Kyarda Dun west of the Jumna,
throughout Assam, thence in a few localities in the Indo-Gangetic plain
from the Eastern Sandarbans to Bahawalpur, and Rohri in Upper Sind, as
well as here and there throughout the extensive area lying between the
Ganges and Godaveri valleys as far eastwards as Mandla, the species being
abundant in portions of the upper Narbada valley, as well as the neighbour-
hood of Bastar to the southward. As pointed out by the late Captain
Forsyth, to whom we are indebted for the best account of its habits, the
range of the swamp-deer in the Central Provinces corresponds with that
of the red jungle- fowl, both species being exclusively confined to the
tracts covered with forest of sal trees. The absence of the species from
Ceylon confirms the opinion expressed above as to its being a comparatively
modern type.

1g2 Rucervine Group

Habits.—Throughout the plains of India this species is commonly
known as the barasingha, that is, twelve-tined deer, but as the same name
is, incorrectly, applied in Kashmir to the hangul, it is preferable to employ
the title swamp-deer, although, as pointed out by Mr. Blanford, that name
is not entirely free from objection. As regards food, the swamp-deer is

Fic. 52.—Front view of Skull and Antlers of Swamp-Deer.
(Rowland Ward, Records of Big Game.)

mainly a grazing species, which makes it the more remarkable that its
distributional area in Central India should be restricted to the limits of one
particular kind of forest tree. Its favourite haunts are the outskirts of
woods, or grassy plains dotted with trees; but in thick forest it is never
seen, although it may be met with in open woods. As the cold season
comes on, it collects in herds, which generally number from thirty to fifty

Schomburgk’s Deer 193

head, but in certain districts may include several hundred individuals. In
the spring these herds break up, and as in Assam single stags may be found
among long grass with their antlers in velvet at the end of March, it is
presumed that the shedding must take place at least as early as February.
Compared with the sambar, the swamp-deer is decidedly a more diurnal
animal, as it may frequently be observed grazing late in the forenoon and
again early in the afternoon, although it invariably retires to the shade for
a mid-day rest. The pairing-season appears to commence about the latter
part of October. Although this species has bred in the London Zoological
Gardens, I have no information as to the usual breeding-time, or whether
more than a single fawn is ever produced at a birth.

A writer quoted by Jerdon thus speaks of these deer in Central India :
“The plain stretched away in gentle undulations towards the river, distant
about a mile, and on it were three large herds of barasinghas feeding at
one time ; the nearest was not more than five hundred yards away from
where I stood ; there must have been at least fifty of them, stags, hinds,
and fawns, feeding together in a lump, and outside the herd grazed three
most enormous stags. . . . Then the herd went off in earnest, showing a
perfect forest of antlers, and the clatter of their hoofs on the hard ground
was like the sound of a squadron of cavalry going to water.”

This deer has received its specific name in honour of the French
naturalist Duvaucel, and it is not a litthke remarkable that such an amount
of error should have crept into the spelling of the Latinised version.

2. SCHOMBURGK’s DEER—CERVUS SCHOMBURGKI

(?) Cervulus cambojensis, Gray, Proc. Zool. Soc. 1861, p. 138.

Cervus (Rucervus) schomburgki, Blyth, Proc. Zool. Soc. 1863, p. 155,
F807, p. 835.

Cervus schomburgki, Sclater, Trans. Zool. Soc. vol. vii. p. 349 (1871) ;
Brooke! F706. Zoo). Soc. 1870, p. 304, 1878, p. go5; W. L. Sclater, Caz.
Mamm. Ind. Mus. part 11. p. 180 (1891) ; Ward, Records of Big Game, p. 15
(1896).

Rucervus cambyensis, Gray, Cat. Ruminants Brit. Mus. p. 76 (1872).

Rucervus schomburgku, Fitzinger, SB. Ak. Wien, vol. \xxix. part i. p. 64
(1879).

Za

194 Rucervine Group

Characters.—Height at shoulder about 3 feet 5 inches ; hair in winter
rather long and coarse. General colour of pelage uniform brown, darkest
on the nose and the upper surface of the tail, and lightest on the cheeks
and flanks; under-parts, lower surface of tail, and lower lip whitish ; a
tinge of rufous on the upper lip, the back of the head, and limbs ; the hair
on the front of the lower part of the fore-leg elongated to form a fringe.
Antlers large, complex, smooth, and polished ; the brow-tine very long,
frequently forked, and arising nearly at a right angle from the beam ; the

Fic. 53.—-Front view of Frontlet and Antlers of Schomburgk’s Deer. From a specimen in the
British Museum.

beam very short and more or less laterally compressed, then forking dicho-
tomously, with each of the main branches about equally developed, and
again forking in a similar manner, to terminate in long cylindrical tines ; in
immature males the hinder branch of the main fork is less developed than
the front one.

This handsome deer is very rare in collections, although several fine
pairs of antlers are exhibited in the British Museum. Never having seen
an entire specimen myself, the foregoing characteristics are taken from Sir
Victor Brooke’s description of a mounted male skin in the Paris Museum,

Thameng 195

brought from Siam in the year 1868. The chief point of distinction from
the preceding species is the shortness of the beam of the antlers below the
bifurcation, added to which is the frequent forking of the brow-tine, and
the generally greater complexity of the antlers.

The following dimensions of antlers of this species are recorded by Mr.
Rowland Ward, all the specimens except the fourth being in the British
Museum :—

Perec Circumference. Tip to Tip. Widest inside. pe eile
30¢ 5 152 29 10-10
298 sd 238 26% 9-8
292 42 282 312 1O-II
284 st iii 292 11-9
27% 53 14 24 9-8
274 43 185 245 7-8

With regard to the name C. cambojensis, Dr. Gray wrote as follows :
“On re-examination I am inclined to believe that the forehead and horns
brought by M. Menhot from Cambojia, and named Cervulus cambgensis, are
only irregularly developed horns of this species. The horns are short, thick,
nearly straight, with a short, thick, recurved branch on the upper part of
the front side near the base; and on one of them there is a somewhat
similar callosity on the hinder side at the same level.” The name “ cambo-
jensis” is the earlier, but since the species is said to be an inhabitant of
Northern Siam, and to be unknown in Southern Siam and Cambodia, this
appears to afford a sufhcient bar to its employment.

Distribution.—The northern districts of Siam.

Two specimens only, both of which were stags, have been exhibited in
the London Zoological Gardens, one received in 1873 and the other in 1880.
The female appears to be unknown ; and there seems to be no record of the
habits of the animal in the wild state.

3. THe THAMENG-——CERVUS ELDI

(?) Cervus smithn, Gray, Proc. Zool. Soc. 1837, p. 45.
Cervus eld, Guthrie (see Blyth, Proc. Zool. Soc. 1867, p. 837), Calcutta
‘fourn. Nat. Hist. vol. 11. p. 417 (1842).

196 Rucervine Group

Cervus (Rusa) frontalis, M‘Clelland, Calcutta ‘fourn. Nat. Hist. vol. i11.
P. 539 (1843). ,

Panolia acuticornis, Gray, List Mamm. Brit. Mus. p. 180 (1843).

Cervus lyratus, Schinz, Synopsis Mamm. vol. ii. p. 395 (1845).

Dama _ acuticornis, Reichenbach, Naturgesch. Saugethiere, vol. iil. p. 16
(1845).

Cervus (Hippelaphus) frontalis, Sundevall, K. Vet. Ak. Hand/. for 1845,
p- 320 (1847).

Panolia eedu, Gray, List Osteol. Brit. Mus. p. 66 (1847).

Panolia eldu, Gray, Cat. Ungulata Brit. Mus. p. 202 (1852), Cat.
Ruminants Brit. Mus. p. 75 (1872), Hand-lst Ruminants Brit. Mus. p. 144
(1873) ; Swinhoe, Proc. Zool. Soc. 1869, p. 652; Sterndale, Mamm. India,
p. 511 (1884).

Cervus eldi, Beevan, Proc. Zool. Soc. 1867, p. 759; Sclater, Trans. Zool.
Soc. vol. vil. p. 348 (1871) ; Brooke, Proc. Zool. Soc. 1878, p. go6; W. L.
Sclater, Cat. Mamm. Ind. Mus. part i. p. 180 (1891) ; Blanford, Fauna Brit.
India—Mamm. p. 541 (1891) ; Evans, ‘fourn. Bombay Nat. Hist. Soc. vol. ix.
p- 326 (1895) ; Ward, Records of Big Game, p. 16 (1896).

Panolia frontahs, Fitzinger, SB. Ak. Wien, vol. |xviii. parti. p. 352 (1873),
Ixix. part i. p. 592 (1874).

Rucervus eldi, Garrod, Proc. Zool. Soc. 1877, p. 17.

Characters.—Height at shoulder about 3 feet g inches, hair very coarse,
shaggy in winter, and much elongated and thickened about the neck in the
males. Antlers rounded and rugose, with a long curved brow-tine, forming
a continuation of the curve of the beam, which is set at right angles to the
pedicle ; the beam unbranched for a considerable distance, curving back-
wards, then outwards and finally forwards, after which it is dichotomously
forked ; the outer branch of the terminal fork more complex than the inner
one, the number of terminal points varying from two or three to at least
eight or ten ; one or more prominent snags usually developed at the junction
of the brow-tine with the beam ; the curve of the two antlers usually more
or less unsymmetrical. Colour of winter pelage of males typically dark
brown on the upper-parts, and white on the under-parts, sometimes with a
white mark above the eye ; in winter fawn-coloured above and pale brown
beneath ; the winter fringe on the neck much darker, and sometimes with

a white gorget; does paler rufous fawn, and the very young generally

Thameng

py /

spotted on the rump with white; a more rufous variety is, however,

known which is fully spotted in the adult.

The peculiar curvature of the antlers at once serves to distinguish the

thameng, or brow-antlered deer, as it has been called, not only from all

other members of the group, but like-
wise from all species of the genus. On
account of this marked peculiarity it
has been referred by many writers to a
distinct sub-genus, or genus, under the
name of Panolia, but there can be no
hesitation in regarding it as a member
of the present group. This view was
first taken by the late Mr. Edward
Blyth, who has been followed by Pro-
fessor A. H. Garrod, and most later
writers. The skull is of the same
general type as in the other two living
members of the group, although the
premaxillary bones, which form the ex-

Fic. 54.—Frontlet and Antlers of Burmese
(Rowland Ward, Records of

Thameng.
Big Game.)

tremity of the muzzle, are shorter and extend a smaller distance up the face.

The following dimensions of antlers are taken from Mr. Rowland Ward’s

Records of Big Game :—

Length along

Outer Curve. Circumference. Tip to Tip.
42 5 29
39% L 313
38% 6+ Be
372 Z 20L
3i7 64 ?
366 z 283
35% 43 193
348 Se 274
342 4 art
34% 5 24
34 58 one

Widest inside.

24
358
30%

3
294

Number of

Points.
=
20

16-19
ES)

P
4-3
aS)
6-6
6-6
6-6

10-10

Locality.

Burma

Siam

Distribution —F lat alluvial districts in the countries on the eastern side

of the Bay of Bengal, extending from the valley of Manipur in the north,

198 Rucervine Group

southwards in suitable localities throughout Burma and the Malay Penin-
sula, and eastwards to Southern Siam, Cambodia, and the island of Hainan.

Habits —This very distinct type of deer was discovered by Captain Eld
about the year 1840, and papers on its habits were contributed both by that
gentleman and by Captain R. C. Beavan ; the following account being taken
from the latter officer’s paper. ‘‘In habits,” he writes, “these deer are
essentially gregarious, and associate with no other species, although hog-deer
abound in the grass and jungle along the edges of the plain ; nor will they
allow the tame buffaloes to come nearer to them than about a hundred yards.
They are very wary and difficult of approach, especially the males. They are
also very timid and easily startled ; the males, however, when wounded and
brought to bay with dogs get very savage and charge vigorously. In fact
the thameng is essentially a plain-loving species ; and although it will fre-
quent tolerably open tree-jungle, for the sake of its shade, it will never
venture into dense or matted underwood, that is bush-jungle in contradis-
tinction to tree-jungle.

‘« When first started the pace of the thameng is great. It commences
by giving three or four large bounds like the axis or spotted deer, and after-
wards settles down into a long trot, which it will keep up for six or seven
miles on end when frequently disturbed. This is where the vegetation on
the plain is comparatively short. In the rains they do not go far before
they find a hiding-place in the paddy [rice]. Their powers of leaping are
highly developed. . . . They are least gregarious in the rainy weather. The
females have mostly then retired in twos and threes into quiet spots, and the
herds are altogether more scattered, owing to the increased density of the
vegetation.

“They feed both during the day and night, chiefly in early morning
and evening. Their food consists chiefly of jungle paddy ; during the night
they doa great deal of damage to the cultivated variety, treading down
more than they eat. They also feed on grass, and the leaves of two jungle-
trees called in Burmese the deay and the thameh. In a tamed state they will
eat plantain-leaves. The call of the female, uttered when disturbed, is a
short barking grunt, that of the males is louder and more prolonged. It is
most frequently heard in the rutting-season, during which the males have
frequent and severe battles. A pair are known to have been captured
whilst so engaged with their antlers interlocked.

Thameng 199

“About the end of January the first jungle fire sweeps over the plain and
destroys the dry herbage, leaving small patches here and there about the
edges of swamps. The second burning takes place about the end of March,
and leaves scarcely a blade of grass behind it ; the plain is then almost
entirely bare, and the deer, having no covert, congregate in large herds.
They are then to be seen on all sides, and, the buffaloes having been with-
drawn to the tree-jungle, are left alone ; they become at this time exces-
sively wary. From the middle of February until the first showers fall at

Seek fj

Fic. 55.—Group of Burmese Thameng. From a photograph by the Duchess of Bedford.

the end of April they apparently subsist without water ; they lie in the
salt swamps during this period, and get the benefit of heavy dews at night.”

It is added that the pairing-season lasts from the middle of March to the
middle of May ; and that the hinds bring forth their fawns in October and
November among the rice-fields, the rice being then either in flower or in
ear, and at its greatest height, so that it affords abundant covert. One fawn
is produced at a birth, and often remains with its dam till the second year.
Breeding may commence at the end of a year and a half. It is specially
stated that while the very young are spotted, all the adults in Burma are
of a uniformly brown colour. In Manipur the stags begin to shed their
antlers in June, but in Lower Burma they are not lost till about September.

200 Rucervine Group

a. BuRMEsE Race—CERVUS ELDI TYPICUS

Characters.—Antlers rounded throughout, with few or no additional
snags on the terminal tines, and the brow-tine usually very long ; colour
generally uniform umber-brown, as described above, or with a very few
spots along the middle of the back.

Distribution —The valley of Manipur, and thence southwards through
Burma and the Malay Peninsula. It has been shown by Mr. Blyth that
antlers from the Malay Peninsula and Mergui are smaller than those from
Manipur and Burma, and have also very frequently two or three vertical
snags on the brow-tine which are wanting in the latter. This difference is,
however, scarcely sufficient to justify a sub-specific separation, especially as
there is in all probability a complete gradation between the two types.

5. SIAMESE RAcE—CERVUS ELDI PLATYCEROS

Panolia platyceros, Gray, List Mamm. Brit. Mus. p. 181 (1843), Cat.
Ungulata Brit. Mus. p. 203 (1852).

Dama platyceros, Reichenbach, Naturgesch. Sdugethiere, vol. iil. p.16 (1845).

Panolia platycercus, Gray, Cat. Ruminants Brit. Mus. p. 75 (1872), Hand-
list Ruminants Brit. Mus. p. 144 (1873).

Plate XV

Characters.—The main termination of the antler much flattened, with a
large number of small snags on the sharp hinder edge ; and the brow-tine
relatively short. Pelage apparently always spotted along the middle line of
the back, and sometimes on the sides as well, reddish at all seasons.

In the British Museum this deer is represented by a fine pair of adult
antlers from Southern Siam presented by the late Sir R. Schomburgk, which
are so different from those of the typical race as to merit sub-specific dis-
tinction. In addition to the snags on the summit, they bear a large
number of small points near the junction of the comparatively short brow-
tine with the beam.

I can find no description of the coloration of Siamese or Cambodian
examples. Mr. Swinhoe! has, however, described and figured antlers from

1 Proc. Zool. Soc. 1869, pp. 652 et seq.

Tue Deer or Aut LAnps. PuatEe XV.

SIAMESE THAMENG IN WINTER PELAGE.

Published. by Rowland Ward Led

Thameng 201

the island of Hainan which he regarded as intermediate between those of
the Burmese and Siamese races, but which were identified by Mr. Blyth in
the same paper with the latter. As they show the same flattened extremi-
ties, with extra snags, there appears no sufficient justification for separating
them from the present race. Mr. Swinhoe describes the Hainan specimens
to which some of these antlers belonged as showing two lines of white spots
on each side of the back. The British Museum possesses three flat skins
from Hainan, sent by Mr. Swinhoe. One of these is an adult in summer
pelage, and displays faint traces of spots all over the back and sides. The
second belonged to a half-grown animal in the shaggy winter coat, and has
little or no trace of spotting. The third is that of a very young fawn, in
which faint indications of spots are discernible on the hind-quarters.

The male specimen from Hainan forming the subject of plate xv was
presented to the London Zoological Society in 1895 by Mr. Julius
Neumann,! and was fully spotted when I first saw it, soon after the
assumption of the winter dress, as it also was in December, when its colour
was foxy red, quite unlike that of the Burmese race. In February 1898 the
fringe on the throat had become very jong and dark-coloured, with a distinct
white gorget. The head-keeper of the Gardens informed me that the
summer pelage was also spotted, with the ground-colour of a bright rufous.

It therefore appears that, with present material, the Hainan representa-
tive of this deer cannot be separated from the Siamese race. And as the
Hainan form frequently shows more or fewer spots on the skin, it is presumed
that the same will hold good with specimens from the mainland. The
slight degree of spotting on the very young skin in the British Museum is
a notable feature. A thameng at Woburn Abbey with a bright red,
although unspotted, winter coat, would seem to belong to this race.

Distribution — Southern Siam, Cambodia, and the island of Hainan.

4. THE StwaLik Swamp-DEeER—CeERVvus sIVALENSIS (Extinct)

Cervus sivalensis, Lydekker, Paleontologia Indica (Mem. Geol. Surv.
Ind.), ser. 10, vol. i. p. xvii. (1880), Cat. Foss. Mamm. Brit. Mus. part il.

perko4; (1885).

1 See Proc. Zool. Soc. 1895, p. 520; the specimen is there stated to be from South China, but it
is labelled Hainan in the Gardens.

2; 1D)

202 Muntyacs

Characters.—This species was named on the evidence of molar teeth,
which present a considerable resemblance to those of C. duvauceli. ‘The
skull provisionally assigned to it is likewise very similar to that of the
latter, while the antlers probably belonging to the same form are apparently
intermediate between those of C. duvauce/t and C. e/d. Even if the skull
and antlers belong to a species distinct from that to which the teeth
pertained, they serve to indicate the existence in the Siwaliks of a member
of the rucervine group.

Distribution —India during the Pliocene period.

IV. Tue Muntyacs—Genus CeErvutus

Cervulus, de Blainville, Bu//. Soc. Philom. Paris, 1816, p. 77 ; Gray,
Cat. Ungulata Brit. Mus. p. 217 (1852), Cat. Rumimants Brit. Mus. p. 93
(1872) ; Brooke, Proc. Zool. Soc. 1878, p. 898 ; Rutimeyer, 4d/. schwerz.
pal. Ges. vol. vill. p. 23 (1881).

Stylocerus, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 319 (1827),
as a sub-genus.

Prox, Ogilby, Proc. Zool. Soc. 1836, p. 135; Fitzinger, SB. 4k. Wien,
vol, lxvili. part i. p. 361 (1873), Ixxix. part 1. p. 40 (1879).

Muntjacus, Gray, List Mamm. Brit. Mus. p. 173 (1843).

Characters.—Lateral metacarpal bones as in Cervus, that is represented
only by their upper extremities. Antlers not exceeding half the length of
the head, with a short brow-tine and an unbranched beam, supported on
long bony pedicles, continued downwards as convergent ridges on the
forehead of the skull; in the female tufts of bristly hair and small projec-
tions in the position of the antlers. Typically a small skin-gland on the
inner side of each frontal ridge. Muzzle with a large naked portion,
extending up between the nostrils, with the upper border of the portion
between the same slightly convex; ears rather small; tail long, thin, and
pointed ; face long. Pelage of adult uniformly coloured above, spotted in
young. No metatarsal or tarsal glands and tufts. In the skull the gland-
pit very large and deep, and the unossified vacuity of moderate extent ;
face-gland large. Median hoofs moderate, lateral hoofs generally small
and rudimentary, and the phalangeal bones of their digits entirely wanting.

Indian Muntyac 2.0 3

Upper canines of male forming long recurved tusks, not growing from
persistent pulps, those of female smaller ; cheek-teeth tall-crowned.
Stature small, and rump elevated.

The muntjacs, or rib-faced deer, as they are expressively called, form a
small but very natural group. In the presence of a brow-tine to the antlers
and the structure of the lateral metacarpal bones they resemble the typical
deer ; but in the characters of the antlers and their long pedicles, as well as
in the retention of tusk-like upper canines, they agree with extinct forms
noticed below, so that the divergence between Cervus and Cervu/us must
evidently have taken place at a remote epoch. In the uniformly coloured
coat of the adult, as well as in the rudimentary condition of the lateral
hoofs and the total loss of the lateral phalangeal bones of the feet, they
display highly specialised characters. ‘They may be divided into two
groups, according to whether the coloration of the adult pelage is of a
rufous or yellowish tinge, or of a deep sepia-brown ; the members of the
second group forming a connecting link between the more typical members
of the genus and the tufted deer.

Distribution—The Oriental and adjacent districts of the Holarctic
region.

1. Tue Inptan Munryac—CervuLus MUNTJAC

Cervus muntjac, Zimmermann, Geogr. Geschichte, vol. 11. p. 131 (1780).

Cervus vaginalis, Boddaert, Elenchus Anim. p. 136 (1785).

Cervulus moschatus, de Blainville, Bull. Soc. Philom. Paris, 1816, p. 77 ;
Gray, Cat. Ungulata Brit. Mus. p. 218 (1852), Cat. Ruminants Brit. Mus.
Pp: 93 (1872).

Cervulus subcornutus, de Blainville, /oc. cit. (1816).

Cervulus muntjac, de Blainville, /oc. cit. (1816) ; Brooke, Proc. Zool.
Soc. 1874, p. 38, 1878, p. 899; Anderson, Yunnan Expedition, p. 337
(1878) ; Sterndale, Mamm. India, p. 500 (1884); W. L. Sclater, Caz.
Mamm. Ind. Mus. part ii. p. 173 (1891); Blanford, Fauna Brit. India—
Mamm. p. 532 (1891) ; Ward, Records of Big Game, p. 1 (1896).

Cervus (Stylocerus) muntjac, H. Smith, in Griffith’s Animal Kingdom, vol.
iverps 44ev..p. 319 (1827). .

Cervus (Stylocerus) aureus, H. Smith, of. cit. vol. iv. p. 148 (1827).

204 Muntyacs

Cervus (Stylocerus) moschatus, H. Smith, op. cit. vol. iv. p. 149, v. p. 320
(1827).

Cervus (Stylocerus) subcornutus, H. Smith, of. cit. vol. v. p. 320 (1827).

Cervus albipes, F. Cuvier, Hust. Nat. Mamm. fasc. \xv. part vii.
(1824-42).

Cervus ratwa, Hodgson, As. Researches, vol. xviii. part Ul. p. 139
(1833).

Stylocerus muntjak, Jardine, Naturalists Library—Mamm. vol. iii. p. 185
(1835).

Stylocerus aureus, Jardine, op. cit. p. 185 (1835).

Stylocerus subcornutus, Jardine, /oc. cit. (1835).

Prox moschatus, Ogilby, Proc. Zool. Soc. 1836, p. 135.

Cervus melas, Ogilby, in Royle’s Himalayan Botany, p. \xxiii. (1839).

Stylocerus ratwak, Hodgson, “fourn. As. Soc. Bengal, vol. x. p. 914
(1841).

Muntjacus vaginalis, Gray, List Mamm. Brit. Mus. p. 173 (1843).

Prox ratva, Sundevall, K. Svenska Vet. Ak. Handl. for 1844, p. 185
(1846) ; Fitzinger, SB. 2k. Wien, vol. \xvill. part 1. p. 302°(1873), xxx
part 1. p. 41 (1879).

Prox albipes, Sundevall, /oc. cit. (1846) ; Fitzinger, op. cit. Ixviii. p. 362,
Ixxix. p. 42 (1879).

Prox stylocerus, Sundevall, /oc. cit. (1846) ; Fitzinger, of. cét. xviii.
p. 362, Ismix. p. 48/(1679).-

Prox muntjac, Sundevall, foc. cit. (1846); Fitzinger, op. cit. Ixviil.
p. 362, Texik po Gaao7on:

Prox melas, Sundevall, /oc. cit. (1846).

Styloceros muntjac, Cantor, “‘fourn. As. Soc. Bengal, vol. xv. p. 269
(1846).

Cervulus vaginalis, Gray, Proc. Zool. Soc. 1850, p. 234, Ann. Mag. Nat.
Hist. ser. 2, vol. v. p. 425 (1850), Cat. Ungulata Brit. Mus. p. 217 (1852) ;
Adams, Proc. Zool. Soc. 1858, p. 538.

Cervulus aureus, Jerdon, Mamm. India, p. 264 (1867).

Cervulus curvostylis, Gray, Cat. Ruminants Brit. Mus. p. 94 (1872),
Hand-list Ruminants Brit. Mus. p. 165 (1873).

Cervulus tamulicus, Gray, loc. cit. (1872).

Characters.—Height at shoulder varying from 20 to 22 inches. Ears

Indian Muntyac 205

narrow and pointed; tail short (length, inclusive OM haite7 sinches)/; no
tuft on crown of head ; lateral hoofs very small; hair comparatively short
and fine. General colour deep chestnut-red, darkest on the back, and
paler on the under-parts ; face and limbs brownish, a black line on the
inner side of the pedicles of the antlers continued for some distance down
the side of the ridges on the face ; tufts of black bristly hair in the female
on the spots where the pedicles of the antlers of the male are situated ;
chin and upper portion of the throat, the hinder part of the abdomen, the
inner side of the thighs, and under surface of the tail white ; anda small
whitish mark on the front of each foot above the
hoof. There is normally no black band on the
nape of the neck, although a faint indication of
such a line has been recorded by Dr. J. Anderson
in one Nepalese skin. The maximum recorded
length of the antlers is 74 inches.

Distribution. —Much discussion has arisen as
to whether there is more than one species of

Indian muntjac, and also as to whether the =
Malayan forms are identical with the Indian. py, 56.—Frontlet and Antlers
(hatalieethese belong «te yone ‘species, may, 1 °f Indian Muntjac. (Row:
i : ome land Ward, Records of Big
think, be regarded as certain, although it is Game)
quite probable that fuller material might enable
one or more of the Malayan races to be separated as distinct sub-species.
On this subject Sir Victor Brooke wrote as follows in 1874 :—‘‘In a
large collection of the skins, skulls, and horns of this species, which I have
received from all parts of India and Burma, and in a considerable number
of living specimens which I have examined, I have observed among adult
animals so much difference in size and intensity of coloration that I have
found it impossible to retain the muntjac of Java and Sumatra as a distinct
species. The muntjacs from the south of India are, as a rule, smaller than
those from the north, as is also the case with the axis and Indian antelope.
But even this rule is subject to many exceptions. I have received from
Northern India perfectly adult, and even slightly aged, specimens of both
muntjac and axis inferior in size to the average as presented by these
species in Southern India. These small races are always connected with
particular areas, and are doubtless the results of conditions sufficiently

206 Muntyacs

unfavourable to prevent the species reaching the full luxuriance of growth
and beauty of which it is capable, though not sufficiently rigorous to
prevent its existence.” .

In 1878 the same writer made the following additional observations :—
“This species appears to attain a larger size in Java, Sumatra, and Borneo
than it does on the mainland ; and I think it not improbable that per-
sistent race characters may eventually be found distinguishing the muntjac
of these islands from that of British India.”

This latter paragraph indicates the probability of the island races being
sub-specifically distinct, but says nothing as to the form from Burma and
the Malay Peninsula as compared with the one from India.

In 1878, Dr. J. Anderson, who has devoted special attention to the
question, wrote as follows :—‘* Hodgson separated the Nepal form as
distinct, having described it as Sty/ocerus ratwak, and Sundevall adopted this
opinion, but separated another from the same locality under the name of
Prox stylocerus. This latter naturalist also considered the Central and
Southern Indian barking-deer as a distinct species, and identified it with
the Prox albipes of Wagner. This latter race Sykes had considered as
C. muntjac, and more lately Gray renamed it under the designation of
C. tamulicus. I have examined the types of Sty/oceros ratva and C. tamuticus,
but I cannot detect that they differ specifically from Cervulus muntjac of
other parts of India, and the specific name applied by Sundevall to the
Malabar race seems to indicate that it is also the same. They appear to
me to be only local races of one widely distributed species which ranges
over the Himalaya, India, and Ceylon through Arracan and Burma to the
Malayan Peninsula, Sumatra, and Java, spreading from the Himalaya east-
wards to the seaboard of China, and, according to Swinhoe, stretching to
the island of Hainan, where, he says, C. reeves? is replaced by the allied
Indian form.” ‘To this distributional area Borneo has to be added. Examples
from the latter island, Sumatra, and Java, are, as already said, of somewhat
larger dimensions than those from the Indian mainland, and should for this
reason probably be regarded as sub-specifically distinct, but further material
is required before the splitting up can be attempted.

Habits.-The names of this species, both scientific and popular, are
remarkably numerous. In Hindustani it is known by the title of kakar,
while among English names we have, in addition to muntjac and rib-faced

Indian Muntyac 207

deer, the appellations of barking-deer, jungle-sheep, and red hog-deer ;
the latter being apparently confined to Ceylon, while jungle-sheep, which
is also applied to the four-horned antelope, is employed in Southern
India.

The muntjac, like so many of the smaller deer, is a non-gregarious
species, generally found solitary, although associating at certain seasons of
the year in pairs. Very rarely have four, or even three, been seen in
company. Its movements in covert are cautious and stealthy, the animal
carrying its head low, and creeping among tangled thickets or under fallen
timber, with the hind-quarters elevated. In running, the same postures are
adopted, but when walking quickly the legs are slowly lifted in a peculiarly
deliberate manner. These deer are seldom seen outside thick covert, which
they only leave to feed on the grass in the immediate outskirts or in
artificial clearings. A peculiarity, probably shared by the other members
of the genus, is the great length of the tongue, which can be protruded to
such an extent that the whole face is capable of being licked. In spite of
their very small size the antlers of the bucks appear to be shed annually,
falling in May, and being replaced by August. Although in some districts
at least the fawns appear to be produced at any time, the regular pairing-
time in Northern India is mainly during January and February, six months
after which the young make their appearance in the world; either one or
two being produced at a birth.

The hoarse, barking cry from which the species derives one of its
popular names is loud, and uttered both during the pairing-season, and
under the influence of alarm. Morning and evening are the usual times when
this call is heard echoing through the stillness of the jungle, but it 1s some-
times uttered after dark, and in the cold season in Burma Mr. Blanford has
heard it in the afternoon. In fighting, the bucks make use of their long
upper canine teeth as their principal, if not sole, weapons of attack and
defence. In captivity Mr. Sterndale states that muntjacs are coarse feeders,
eagerly devouring any kind of meat that may be offered them. Neverthe-
less, their flesh is of excellent quality, far superior to that of most Indian
deer.

At Woburn Abbey muntjacs are allowed to run wild in some of the
coverts, where they thrive well, although of course they are but seldom
seen except during a beat.

208 Muntyacs

2. THe Tiperan MuntyAc—Cervulus LACHRYMANS

Cervulus lacrymans, Milne-Edwards, Arch. Mus. Paris, vol. vii. p. 93
(1871), and Recherches Mamm. p. 348 (1872-74) ; Brooke, Proc. Zool. Soc.
1874, p. 40, 1878, p. 899 ; Anderson, Yunnan Expedition, p. 338 (1878).

Cervulus sclater1, Swinhoe, Proc. Zool. Soc. 1872, p. 814; Brooke, sid.
1874, p. 40; Gray, Hand-lst Ruminants Brit. Mus. p. 165 (1873); W. L.
Sclater, Cat. Mamm. Ind. Mus. part 11. p. 173 (1891).

Characters.—Nearly allied to the last, but slightly smaller, the height
at the shoulder being about 1g inches. General colour of pelage bright
rufous brown, with the hairs on the back speckled, the neck and head
bright yellowish brown; the face-markings and under-parts as in the
preceding species.

Sir Victor Brooke’s description of this species is as follows :—

“The entire forehead, occiput and outer side of ear-conchs yellow.
The intensity of the yellow varies in different specimens, but is always
sufficiently strong to form a very conspicuous character in the appearance of
the males of this species. A line running up the inside of the horn-
pedestals, starting from the frontal glands, jet-black, this marking contrasting
strongly with the yellow of the forehead; cheeks, anterior parts of neck and
throat, belly, and upper surface of tail foxy red. Chin, a line running down
the anterior surface of the tibial portion of the hind-limbs and under surface
of tail white. The rest of the body bluish brown speckled with red.”

In addition to its inferior size, the chief external characteristic distin-
guishing this species from the last appears to be the brighter and more
yellow tint of the head and neck. The skulls of the two forms are distinguished
sufficiently by their size, that of C. muntjac measuring 84 inches in length,
and that of the present species only about 7 inches. ‘The type specimen is
in the Paris Museum.

Distribution —Moupin, in Eastern Tibet, and the hills near Hangchau,
China ; probably also in some of the intermediate districts.

3. Tue CuinesE MunryAc—CervuLus REEVESI

Cervus reeves, Ogilby, Proc. Zool. Soc. 1838, p. 105.
Prox reevesii, Sundevall, K. Svenska Vet. Ak. Handl. for 1844, p. 185

THe DEER or Att Lanps. Pirate XVI.

TENASSERIM (1) anp HAIRY-FRONTED (z) MUNTJACS.

Lhiahial bry Rovdand Ward. Lta

~~

Tenasserim Muntyac 209

(1846) ; Fitzinger, SB. 4k. Wien, vol. \xviil. part i. p. 362 (1873), xxix.
pattitn p: 6o*(1879).

Cervulus reeves, Gray, Knowsley Menagerie, p. 65 (1850), Cat. Ungulata
Brit. Mus. p. 220 (1852), Cat. Ruminants Brit. Mus. p. 94 (1872) ; Brooke,
Proc. Zool. Soc. 1878, p. 899 ; Anderson, Yunnan Expedition, p. 338 (1878).

Cervulus reevest, Swinhoe, Proc. Zool. Soc. 1862, p. 361; W. L. Sclater,
Cat. Mamm. Ind. Mus. part ii. p. 174 (1891).

Cervulus micrurus, Sclater, Proc. Zool. Soc. 1875, p. 421, 1876, p. 696.

Characters.—The smallest member of the group, the size being about
one-third less than that of the Indian species. Face shorter and broader,
the pedicles of the antlers less divergent, and the face-glands considerably
larger than in the two preceding species. General colour of pelage reddish
chestnut speckled with yellowish gray, a distinct black band running down
the nape of the neck, limbs blackish brown, under-parts and face-markings
much as in the two preceding species. Length of skull 54 inches ; its
gland-pit very large.

The distinctive points in this species are the small size, the large face-
gland, and the black nape-streak. Dr. Anderson gives minute details as to
differences between the skull and that of C. reevesz.

Distribution.—Southern China, as far north as Ningpo, and Formosa.

Habits—In Formosa Mr. Swinhoe states that this muntjac affects the
lower ranges of hills, which are covered with long coarse grass and tangled
thicket. “It is there usually found in small herds, basking in the sun, or
lying in hidden lairs. They are very seldom approached near, except by
stealth. The least noise startles them, and they dash away with bounds
through the yielding grass, occasionally showing their rounded backs above
the herbage. They have, however, their regular creeps and passes through
the covert, near which the natives lie when stalking them, while others
drive them. The little startled creatures hurry from danger along those
beaten tracks, and are thus picked off with the matchlock.”

4. Tue TENAsseERIM MuntyAc—CERVULUS FE

Cervulus fee, Thomas and Doria, dun. Mus. Genova, ser. 2, vol. vii. p. 92
(1889) ; Blanford, Fauna Brit. India—Mamm. p. 534 (1891); Thomas,
Ann. Mus. Genova, vol. x. p. 945, plate x (1892). _

Plate XVI, fig. i
2E

210 Muntyacs

Characters.—Allied to the next species, with which it agrees in its sepia-
brown, instead of reddish or yellowish chestnut, general coloration, and its
black and white, instead of chestnut and white, tail. Distinguished from
C. crinifrons by the absence of the tuft of hair on the forehead, and the
consequent clearly defined face-markings, the presence of a white line on
the lower part of the hind-leg, and the much shorter tail. Size about equal
to that of the Indian muntjac.

The following is the original description. General colour of body
uniform dark brown, the centre of the crown, the pedicles of the antlers,
occiput, and the region round the bases of the ears bright yellow ; a black
line running up the inner side of each pedicle ; neck uniform brown ; fore-
legs brown superiorly, darkening to black on the metacarpals, with the
terminal inch next the hoof white all round, and a line of scattered white
hairs running up the front to the carpal joint (knee) ; hind-legs similarly
coloured, but with a distinct white line on the front. Tail short, black
above, white below and at the sides, the two colours sharply contrasted.
Under-parts brown, mixed with whitish on the chin and inner surfaces of
thelimbs. Face-glands large and prominent, but those on forehead wanting.
This species, which is known only by a single specimen preserved in the
museum at Genoa, forms a connecting link between the three preceding
forms and C. crinifrons.

Distribution —Mountains south-east of Maleyit, Tenasserim, and probably
other parts of the country.

5. THe Hatry-FRonTED MuntyAc—CERVULUS CRINIFRONS

Cervulus crinifrons, Sclater, Proc. Zool. Soc. 1885, p. 1, plate 1.; Styan,
ibid. 1886, p. 267.
Phate XV, jigs 2

Characters—The largest species, the height at the shoulder being about
244 inches. Forehead and crown of head tufted, so as to conceal the
pedicles of the antlers ; hair longer and coarser, ears shorter, more rounded
and more thickly haired externally, the tail much longer, and the lateral
hoofs larger than in either of the chestnut-coloured species. General colour
of pelage dark sepia-brown, with a tinge of purple, and the back finely
speckled with rufous ; head-tuft, ears, forehead, and cheeks bright orange

Hairy-Fronted Muntyac STE

chestnut ; a V-shaped faint dark mark near the pedicles ; limbs and under-
parts dark blackish brown, except the abdomen, which is whitish ; inner
side of hind-legs rufous chestnut ; inner surface of thighs, and sides and
lower surface of tail pure white ; upper surface of tail and a line extending
on to the rump black. Length of tail about g inches. Frontal glands
probably absent.

The above description is taken from a male specimen forming the type,
which was purchased by the Zoological Society in 1884 from Mr. Michie,
and is now mounted in the British Museum. From the small size of the
antlers and tusks, it is not improbably immature, so that the height may
really be greater than stated above.
The skin of a female was procured
in 1886 by Mr. F. W. Styan, who
gives the following description of its
colour : — ‘‘ Upper-parts rich dark
glossy brown, mixed with black,
darkest along the middle of the
back. Belly and inner sides of thighs
pure white, the line of demarcation
very distinct, except in the lower
part of the thighs, where it merges

gradually into dark brown. Tail
Fic. 57.—-Head of Hairy-fronted Muntjac.

(Rowland Ward, Records of Big Game.)
pale patch under the forearm; rest Whe tusks are not shown.

of the under-parts dark brown. The
brown of the neck gradually passes into reddish brown on the sides of the

above black, below pure white; a

head ; the nose-ridge dark brown; the longitudinal slits form a black V,
the colour between them isa bright reddish brown, which turns into a fery
orange-brown in the tuft of bristly hair, rather more than two inches long,
which rises from the top of the forehead. On each side of this is a very
small bony pedicle surmounted by short tufts of hair of the same colour.
Ears at the base of the same hue, but not quite so bright, and fading into
brown on the upper part.”

In the tuft on the forehead, the small rounded ears, the coarse hair,
general coloration, and perhaps also the small size of the antlers, this species
serves to connect the more typical muntjacs with the tufted deer.

212 Tufted Deer

Distribution.—Eastern China, in the neighbourhood of Ningpo.

Nothing is known as to the habits of this species, but Mr. Styan states
that it is apparently very rare, his collector refusing to believe in its existence
until he came across the female specimen described above. The only other

known example is the type male.

V. Tue Turrep DrEER—GENUS ELAPHODUS

Elaphodus, Milne-Edwards, Arch. Mus. Paris, vol. vii. p. 93 (1871),
Recherch. Mamm. p. 353, plates Ixv-Ixvii (1872-74) ; Garrod, Proc. Zool. Soc.
1876, p. 757; Brooke, zd. 1878, p. 899; Riitimeyer, AdA. schwerz. pal.
Ges. vol. vill. p. 28 (1881).

Lophotragus, Swinhoe, Proc. Zool. Soc. 1874, p. 452.

Characters.—Lateral metacarpals as in Cervus, but still more rudimentary,
and occasionally wanting. Antlers very small, supported on long slender
pedicles, which diverge on the face, and do not send down long ridges on
the forehead of the skull ; scarcely projecting above the large frontal tufts
of hair. No glands on the forehead. Other characters generally as in
Cervulus, but the long tusks of the male not everted at the tips, the hair
always very shaggy and coarse, the ears broad, rounded, and thickly haired
externally, and the tail of moderate length. The young are spotted only
along the middle line of the back.

The genus is very closely connected with the preceding by means of the
hairy-fronted muntjac, and it may even be a question whether the two
should not be merged in one; in which case the two members of the
present group should be regarded as sub-species rather than species. Both
Cervulus and Elaphodus present a remarkable peculiarity in the structure of
the ankle-joint, or tarsus, found in no other Old-World deer. In common
with other ruminants, all the Cervide have the navicular and cuboid bones!
of the tarsus welded together into a compound bone known as the naviculo-
cuboid ; but in the two genera mentioned a further solidification of this
joint is brought about by two other bones, respectively termed the middle
and outer cuneiform bone, uniting with the naviculo-cuboid. Further evid-
ence of their affinity is afforded by the structure of the skull, the large

’ For explanation of these terms, see Flower, Osteology of the Mammalia.

Tibetan Tufted Deer oa

tusks, and the absence of the metatarsal tufts. It was once thought that
the presence of frontal cutaneous glands formed a point of distinction, but
these are now known to be wanting in Cervulus fea, as they probably are in
C. crinifrons.

Distribution —The south-eastern portion of the Eastern Holarctic region.

1. THe TIBETAN TurTED DEER—ELAPHODUS CEPHALOPHUS

Elaphodus cephalophus, Milne-Edwards, Arch. Mus. Paris, vol. vil. p. 93
(1871), Recherch. Mamm. p. 353 (1872-74) ; Garrod, Proc. Zool. Soc. 1876
Pp. 757, in part ; Brooke, sid. 1878, p. 899.

Characters. —General proportions much the same as in the Indian munt-
jac, the height at the shoulder being from about 22 to 23 inches. Hair
very coarse and pithy. General colour of pelage deep chocolate-brown ;
the hairs in front of a vertical line drawn through the shoulder-joint being
whitish at the base, and gradually becoming dark brown towards the tip,
quite close to which is a narrow white ring, thus producing a speckled
appearance in this region ; posteriorly to the line mentioned the white on
the hairs absent, each hair gradually darkening from the white base to
become rich brown at the tip over the sides and back, the colour being
deeper on the middle line of the latter, and also darkening on the legs,
which near the hoofs become almost or completely black ; in some cases a
white line on the front of the legs above the hoofs. Hair of frontal crest
forming a horse-shoe on the forehead of a nearly black colour, bordered by
a gray line above the eye; ears whitish internally, the tip and the greater
part of the inner edge nearly pure white, and a transverse black band about
three-quarters of an inch in depth extending across the lower part of the
inner surface ; under surface of tail and inside of thighs and buttocks white.

This species, which is the type of the genus, is only known to me by the
descriptions of Messrs. Milne-Edwards and Garrod, the type specimens
being in the Paris Museum, and the British Museum possessing no examples.
It was one of the animals first discovered by Pere David, in his journeys
through Tibet.

Distribution.—Eastern Tibet, in the neighbourhood of Moupin. Nothing
appears to be ascertained as to the habits of this species.

214 Tufted Deer

2. Micuie’s Turrep DrER—ELAPHODUS MICHIANUS

Lophotragus michianus, Swinhoe, Proc. Zool. Soc. 1874, p. 452; Garrod,
ibid. 1876, p. 757, plate lxxvi.
Elaphodus michianus, Brooke, Proc. Zool, Soc. 1878, p. goo ; Styan, sid.

1886, p. 268.
Plate XVII, fig. 1

Characters— Very closely allied to the last, from which it is chiefly
or entirely distinguished by small differences of coloration. General hue

Silos

Fic. 58.—Head of Immature Male of Michie’s Tufted Deer. From Garrod, Proc. Zool. Soc. 1876.

more decidedly iron-gray, or pepper-and-salt colour, the individual hairs
being whitish with brown tips, and only a few of those occupying a very
small area in front of the ear being ringed with white ; lips, eye-brows,
and cheeks paler ; forehead and crest of male brown, of female lighter ;
under-parts slightly lighter, and a little white showing on the front of the
thigh. Young very similarly coloured, but no pale eyebrow, and a row of
not very distinctly marked white spots on each side of the back, followed by
very faint traces of a second outer row.

The history of this species is somewhat remarkable. It was originally
described under the name of Lophotragus by the late Mr. Swinhoe, who

THe Deer or Att Lanps. Puate XVII.

MICHIE’S TUFTED DEER (1) AND CHINESE WATER-DEER (2).

Published by Rowiand Ward Ltd

Extinct Muntyacs 215

appears to have been unacquainted with Professor Milne-Edwards’ descrip-
tion of E/aphodus published a few years previously. In 1876 the late Pro-
fessor A. H. Garrod demonstrated the unity of these two genera, and also
came to the conclusion that the Tibetan and Chinese animals were not specific-
ally distinguishable. This latter view was disputed by Sir Victor Brooke,
who was subsequently supported by Mr. O. Thomas. As already said, the
two forms are so similar that it is quite likely they should be regarded as
sub-species rather than species, but so long as they are placed in a genus
apart by themselves, this is a matter of little importance.

Distribution.—Eastern China, in the neighbourhood of Ningpo, and
probably in other districts.

Habits.—Beyond the statement that it is fond of the neighbourhood of
water and is found abundantly in the reed brakes bordering the rivers of
Eastern China, there seems nothing recorded of the habits of this little deer.
Ten specimens have been exhibited alive in the gardens of the London
Zoological Society between the years 1876 and 1884, and there was one
some time ago at Woburn Abbey. The British Museum possesses mounted
skins of the male, female, and young.

VI. THe Mrocens Munryacs—Genus Dremoruerium (Extinct)

Dremotherium, Geoffroy, Rev. Enclyclop. vol. lix. p. 82 (1833) ; Filhol,
Ann. Sci. Géol. vol. xi. art. 1, p. 40 (1881).

Paleomeryx, Meyer, Foss. Knochen und Zahne Georgesmiind, p. 92 (1834) ;
Ritimeyer, 4h. schweiz. pal. Ges. vol. x. p. 79 (1883) ; Lydekker, Caz.
Foss. Mamm. Brit. Mus. part ii. p. 119 (1885) ; Schlosser, Morphol. “fahro.
1886, pp. 68 and 294.

Micromeryx, Lartet, Notice sur la Colline de Sansan, Pp On To 5m).

Amphitragulus, Pomel, Cat. Methodique, p. 100 (1853).

Propalaomeryx, Lydekker, Pateontologia Indica, ser. 10, vol. ii. p. 173
(1883).

Characters.—Lateral metacarpals in certain cases complete, but in some
of the later forms apparently entirely absent ; antlers wanting in the earlier
species, but present in some of the later ones; when present, mounted on long
pedicles, from which they are imperfectly differentiated by a burr ; simply

216 Extinct Muntyacs

forked, covered with skin, and non-deciduous. Molars short-crowned,
those of the lower jaw with a characteristic fold in the enamel ; tusk-like
canines developed at least in those species in which antlers are wanting.

Following Dr. Max Schlosser, I regard Amphitragulus, in which there
are four lower pre-molars, as not generically separable from Dremotherium
proper, in which there are only three of these teeth. The earlier forms of
the genus (from the Oligocene beds of St. Gérand-le-Puy, France) are all
without antlers, and as many of them appear to retain both ends of the
lateral metacarpals, they may doubtless be regarded as the ancestral types
of the modern Old World deer, and probably also of those of North America.
It will be unnecessary to refer to these Oligocene species by name, and
specific mention is made only of two of those in which antlers are present.
These are of such a type as might be expected to occur in the common
ancestor of the brow-tined and fork-antlered deer of the present day.

In North America the Miocene muntjacs were represented by the
closely allied genera Blastomeryx and Cosoryx. Writing of these, Messrs.
Scott and Osborn? observe that “ B/astomeryx is, so far as we can at present
determine, almost identical with the type variously named in Europe
Paleomeryx and Dremotherium, about the only difference of importance
being the absence of the characteristic Pa/eomeryx-fold on the lower
molars. Cosoryx is very closely allied to B/astomeryx, and is distinguished
from it by the much more hypsodont (tall-crowned) molars.” Cosoryx is
admitted to be the ancestor of the American prong-buck (Antilocapra), and
has also been regarded as having given rise to the American deer, but
doubts as to the correctness of the latter theory are suggested in the sequel.
With regard to the antlers of these forms, Messrs. Scott and Osborn state
that they were almost certainly covered with skin, their smooth surface
showing that they could not have been bare. They likewise state that
their own observations show ‘‘ nothing which can be opposed to the view
expressed by Professor Cope that B/astomeryx should be placed in the
ancestral line of the distinctly American deer. A/ces, Rangifer, and Cervus
are really immigrants from the Old World, and do not belong to this
category ; but the truly American types, of which Cariacus (= Mazama) is
the chief example, have a peculiar skull-structure, first pointed out by
Garrod, which seems to show that the American deer were separated from

1 Bull. Mus. Harvard, vol. xx. p. 82 (1890).

Extinct Muntyacs Daley

those of the Old World at a comparatively early date, although it is very
questionable whether both series could have independently acquired the
extraordinary peculiarity of the deciduous antler.”

When treating of Cosoryx, they observe that ‘‘both Blastomeryx and
Cosoryx are probably to be derived from the species referred to the former
genus which occur in the John Day beds, but there is no form yet known
in the White River which could have given rise to the John Day
ruminants. The latter are most probably descended from some Pa/eomeryx
(= Dremotherium) of the Old World which migrated to this continent.” We
have thus, even on this view, an original Old World origin for all the
American deer.

1. Hensev’s MunrjAc—DREMOTHERIUM FURCATUM (Extinct)

Paleomeryx scheuchzert, Meyer, Neues ‘fahrb. Min. 1838, p. 413, 1n part.

Prox furcatus, Hensel, Zeitschr. deutsch. geol. Ges. vol. xi. p. 278 (1859).

Paleomeryx furcatus, Lydekker, Cat. Foss. Mamm. Brit. Mus. part 11.
p. 120 (1885) ; Schlosser, Morphol. ‘fahrb. vol. xii. p. 68 (1886).

Characters.—This is a large species from the Miocene beds of Bavaria
and France characterised by possessing antlers divided into a long fork
immediately above the incipient burr. Apparently the lateral metacarpals
are wanting. Dr. Schlosser remarks that this species is the earliest antlered
deer, the simply forked antlers being never shed and covered during life
with skin. Owing to the apparent absence of the lateral metacarpals, the
species cannot be the ancestor of any of the existing deer.

2. Kaup’s Munryac—DreMoTHERIUM ANOCEROS (Extinct)

Cervus anocerus, Kaup, Archiv. Mmeral. vol. vi. p. 217 (1833).

Cervus dicranoceros, Kaup, op. cit. p. 219 (1833).

Cervus trigonoceros, Kaup, op. cit. p. 221 (1833).

Dicrocerus anocerus, Gaudry, Les Enchainements, etc. p. 84 (1878).

Cervulus (?) dicranoceros, Lydekker, Cat. Foss. Mamm. Brit. Mus. part il.
ps 17 (1885).

Characters.—This species, of which the remains occur typically in the
Pliocene strata of Eppelsheim, Hessen Darmstadt, appears closely allied to

2F

218 Extinct Muntyacs

the last, but the beam of the antler does not divide till after a considerable
distance from the incipient burr, and the fork is short. The limbs are

unknown.

Fic. 59.—Frontlet and Antlers of Hensel’s Muntjac. After Riitimeyer.

VII. Tue Tuskress Munryacs—Genus Dicroceros (Extinct)

Dicrocerus, Lartet, Notice sur la Colline de Sansan, p. 34 (1851) ;
Schlosser, Morphol. ‘fahrb. vol. xii. p. 68 (1886).

Characters.—According to Dr. Schlosser, the one known species of this
genus differs from Dremotherium by the slight development of the fold in
the enamel of the lower molars, and by the antlers exhibiting a stronger
development, with the commencement of a distinct burr ; thus, at any rate,

W ater-Deer 219

foreshadowing a periodical replacement. According to Filhol, the upper
canines, if present at all, are small.
Distribution.—Europe in the Miocene epoch.

THE SAnsAN Munryac—Dicroceros ELEGANS (Extinct)

Dicrocerus elegans, Lartet, Notice sur la Colline de Sansan, p. 34 (1851),
in part ; Gaudry, Les Enchainements, etc. p. 84 (1878) ; Schlosser, Morphol.
‘fahrb. vol. xi. p. 68 (1886).

Cervus dicroceros, Gervais, Zool. et Pal. Francaises, 1st ed. p. 86 (1848-52) ;
Filhol, Mamm. Foss. de Sansan, p. 268 (1891).

Characters.—The remains of this species, which have often been con-
founded with Dremotherium furcatum, occur in the Miocene strata of Sansan,
in the Department of Gers, France, and probably in the contemporaneous
deposits of other parts of the Continent. The antlers have the same
general form as those of D. furcatum, but the cheek-teeth resemble those
of Cervus, and in M. Filhol’s figure no upper tusks are shown.

VIII. THe Watrer-DEErR—Genus Hypretapuus’*

Hydropotes, Swinhoe, Proc. Zool. Soc. 1870, p. 90; Gray, Cat. Ruminants
Brie Mase pa o5 (1572); Brooke Proc. Zool. Soc. 1872, p. 522, 1878,
p- 916; Garrod, sid. 1877, p. 789; Riitimeyer, AA. schwerz. pal. Ges. vol.
vill. p. 21 (1881); Forbes, Proc. Zool. Soc. 1882, p. 636; nec Hydropota,
Rond, 186r.

Characters—Lateral metacarpal bones as in Rangifer, that is to say,
represented only by their lower extremities. Antlers wanting. Muzzle
with a large naked portion surrounding the nostrils ; ears moderate, narrow,
and pointed ; tail very short ; face rather long. Pelage uniformly coloured
in the adult, spotted in the young. No metatarsal or tarsal glands and tufts.
In the skull, the gland-pit small but deep; face-gland small. Lateral
hoofs of moderate size. Upper canines of male forming very long, curved,
and slightly convergent tusks, which grow from semi-persistent pulps, those
of the female smaller ; cheek-teeth tall-crowned. Size small. In very old
animals the pulp-cavity of the upper canines obliterates.

1 New name.

220 Water-Deer

As additional characters of the skull, it may be noticed that the auditory
bulle on its lower surface are greatly inflated, and that the hinder angle of
the lower jaw is much produced backwards, forming a compressed semi-
circular process projecting behind the level of the condyle by which the
jaw is articulated with the skull. The vomer does not divide the aperture
of the posterior nostrils. The hind-feet have deep glands between the
hoofs, but in the front pair the corresponding glands are small and shallow.

In his Catalogue of Ruminants the late Dr. Gray classed the Chinese
water-deer with the musk-deer in a separate family. ‘The resemblance
between the two is, however, entirely superficial ; Professor Garrod and
others having shown that their internal anatomy is quite distinct. From
the structure of the ankle-joint and the lateral metacarpal bones, as well as
from the inflated auditory bulle of the skull, there is probably no very
close affinity with the muntjacs and their allies. Sir Victor Brooke, who
at first thought the water-deer might be related to the rusine deer, sub-
sequently came to the conclusion that its affinities were rather with the
roes, which it resembles in the structure of the lateral metacarpals and the
form of the lower jaw. On this subject the late Mr. W. A. Forbes, after
a study of certain parts of its internal anatomy, wrote as follows :—“ Sir
Victor Brooke has been led, from a consideration of other points, to
associate Hydropotes and Capreolus with Alces, as a group per se, with affinities
in some points in the direction of the Old World, in others in that of the
New World forms. It appears to me that the additional evidence in this
paper, especially that derived from the resemblance of the generative
organs, is strongly in favour of this association, so far, at least, as Hydropotes
and Capreolus are concerned. The general similarity in appearance of
Capreolus to Hydropotes has often struck me, and has even, I believe, led
others into the error of mistaking one for the other! That Hydropotes is in
no way intimately related to Moschus has been already amply demonstrated.”
I can fully endorse the external resemblance between the water-deer and
the roes, when seen at a distance in a park; although the difference in
their movements affords a mode of discrimination. Probably Hydrelaphus
is an ancient type, although no fossils have been referred to it. The
lower jaw of a deer from the Miocene of Sansan in France has, however, been
described under the name of Strongylognathus,' which has the same peculiar

| Filhol, Mamm. Foss. de Sansan, p. 265, plate xxx (1891) ; originally named P/atyprosopus.

Chinese Water-Deer 221

hinder angle, and may indicate an allied form, but as a similar peculiarity
occurs in the roes it cannot certainly be assigned to the present genus.
Distribution.—The eastern extremity of the Eastern Holarctic region.

THe CHINESE WaATER-DEER—HyYDRELAPHUS INERMIS

Hydropotes inermis, Swinhoe, Proc. Zool. Soc. 1870, p. 89; Hamilton,
igene ip eso. 1873, p. 473 3 Brooke, ibid. 1872, p. 522, 1878, p. 916 ;
Gray, Cat. Ruminants Brit. Mus. p. 9§ (1872); Garrod, Proc. Zool. Soc.
G7 7a pa 700s POTbes, 2074. 1382, p. 636.

(?) Hydropotes argyropus, Heude, C. R. Ac. Paris, vol. xcvill. p. 1017
(1884), imperfect description.

Plate XVII, fig. 2

Characters—Height at shoulder about 20 inches. Hair very coarse
and thick, longest on the neck and rump ; the individual hairs on the back
and sides flattened, and undulated from side to side. General colour of
upper-parts light rufous chestnut, stippled with blackish, the rufous tinge
most marked on the head and back of the ears ; the individual hairs grayish
white from the base for the greater part of their length, then blackish
brown, and finally light chestnut, the dark rings giving the stippled appear-
ance to the pelage; neck paler than back, with its under surface plain-
coloured ; shoulder, limbs, and tail brownish chestnut ; under-parts, front
of thigh, chin, throat, a narrow band on the muzzle, a mark above the eye,
and the inner surface of the ear white or whitish. The fawn is somewhat
sparsely and indistinctly spotted with white ; the spots running in longi-
tudinal lines from the neck to the tail, leaving a space of about an inch and
a half in width uniformly coloured. The upper line of spots is fairly
distinct and defined, but the two lower ones are shorter and more irregular.
The indistinctness of the spots is due to the circumstance that they are
formed, not by completely white hairs, but by hairs which are reddish for
the greater part of their length and only tipped with white.

Distribution.—North-Eastern China, typically from the islands in the
Yang-tsi-kiang, and perhaps Corea. The form stated to come from the
latter locality was considered to be a distinct species and named H. argyropus,
but no sufficient description of it seems to have been published, although
it is stated to be lighter coloured than the ordinary form.

222 Roes

Habits—Between the years 1873 and 1883, inclusive, the Chinese
water-deer was represented by seven examples in the London Zoological
Gardens, although none have been received between the latter date and
1896. A female is now (1898) living in the park at Woburn Abbey. A
female in the possession of M. Cornély bred at Tours in 1877, the number
of fawns produced being three ; and it appears that either three or four is
the usual number of young at a birth, the female having four nipples.
This prolific reproduction is quite unique among the Cervida, and is one of
the reasons for regarding the genus as a very ancient type.

The Chinese name of the animal at Shanghai is fe. In his original
description Mr. Swinhoe writes that these small deer “ crouch in the reeds
and long grass, admitting pretty close approach, and then, rising with a
bound, spring away. They were generally put up singly or in twos and
threes. In running, they cock their ears, round their fore-legs, bend up
their hind-legs, hog their rumps, and scurry away with little quick leaps,
very much after the manner of a hare.” These observations are fully
confirmed by the habits of the female at Woburn Abbey. This animal is
kept in a large paddock, where the grass is rather long and tussocky.
Among these tussocks it lies so completely concealed, that hours may be
spent without catching a glimpse of it. When disturbed, it springs off
with the short quick leaps mentioned by Swinhoe. After running for some
distance it drops among the grass with a suddenness which is almost startling.
The grass in its paddock being of only a moderate height, the head and
neck of the little deer are fully visible, and its restless eyes may be seen
watching the intruders. Presently the head and neck are lowered, and the
creature becomes completely hidden. Among its native reeds it would
doubtless become quite invisible from the moment of dropping.

IX. Tue Rors—GeEnvus CAPREOLUS

Capreolus, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 313 (1827) ;
Gray, Cat. Ungulata Brit. Mus. p. 221 (1852), Cat. Ruminants Brit. Mus.
p. 80 (1872) ; Fitzinger, SB. 4k. Wien, vol. |xviii. part i. p. 352 (1873), Ixx.
part i. p. 239 (1874) ; Brooke, Proc. Zool. Soc. 1878, p. 917; Ritimeyer,
Abh. schwerz. pal. Ges. vol. vill. p. 41 (1881).

Characters Dip he

Characters.—Lateral metacarpal bones as in Rangifer. Antlers compara-
tively small, rising very close together and almost vertically from the crown
of the head, without a brow-tine, the beam dichotomously forking at a point
usually about two-thirds of the total length, and the posterior, or upper
prong of this fork, which is the larger, again dividing ; the normal number
of tines being thus three. Muzzle with a large naked portion extending
up between the nostrils, the upper border of the portion between the same
straight, and the part below the nostrils narrow; ears large ; tail very short
and rudimentary ; face short, with the muzzle blunt. Pelage of adult
uniformly coloured, with a pure white patch in the region of the tail in
winter ; young spotted. A metatarsal gland and tuft, situated in the upper
half of the metatarsal segment. In the skull, the gland-pit extremely small
and shallow, the face-gland itself being almost obsolete. Lateral hoofs well
developed. Upper canines wanting ; upper molars tall-crowned, without
distinct additional columns on the inner side. On the under surface of the
skull the vomer not dividing the inner aperture of the nostrils into two
portions. Size rather small ; build tall.

The lower jaw has the same contour posteriorly as in the Chinese water-
deer ; but the auditory bulle on the under surface of the skull are not
markedly inflated, and the unossified vacuity near the gland-pit is reduced
to a narrow slit. The foot-glands are very similar to those of Hydrelaphus,
being rudimentary in the fore-feet, and deep in the hinder pair. With
regard to the systematic position of the genus, the late Professor Garrod’
wrote that “‘ Capreolus caprea is one of the most difficult of the deer tribe to
localise ; and I have placed it not far from Cervu/us on account of the con-
figuration of” certain organs. In some degree this reference is confirmed
by the extinct Dremotherium, in which the antlers are to a certain extent
intermediate between the brow-tined and the forked types. But if Capreo/us
is close to Hydrelaphus and the latter is remote from Cervu/us, the connection
between the first and third cannot be very intimate. In the structure of
the lateral metacarpal bones Capreo/us resembles Mazama ; and by Sir Victor
Brooke Alces, Hydrelaphus, and Capreolus were placed in a group by them-
selves next the American deer. The late Dr. Gray was so fully convinced
of the intimate relation between the present genus and the American deer,
that he termed some of the latter ‘““ American roes”’; and probably this

1 Proc. Zool. Soc. 1877, p. 18.

224 Roes

latter relationship is the closest. The under-mentioned fossil species shows
that the genus is an ancient type. The existing roes are so similar to one
another that they might well be regarded as sub-species instead of species, but
in a restricted genus the matter is of no importance, one way or the other.

Distribution—The Eastern Holarctic region, dating from the early part
of the Pliocene period.

1. THE EurRoPpEAN RoE—CAPREOLUS VULGARIS

Cervus capreolus, Linn. Syst. Nat. ed. 12, vol. i. p. 94 (1766) ; Jenyns,
Brit. Vert. Animals, p. 35 (1835); Bell, British Quadrupeds, p. 407 (1837) ;
Owen, Brit. Foss. Mamm. p. 487 (1846).

Cervus capreolus fossilis, Cuvier, Ossemens Fossiles, ed. 3, vol. iv. p. 103
(1825) ; Meyer, Paleologica, p. 95 (1832).

Cervus (Capreolus) capreolus, H. Smith, in Griffith’s Animal Kingdom, vol.
iv. p. 124, v. p. 314 (1827) ; Blasius, Saugethiere Deutschl. p. 457 (1857).

Capreolus vulgaris, Fitzinger, Beitr. Landesk. Osterreichs, Vol. Lipset si
(1832), SB. 2k. Wien, vol. Ixviui. parti. pi 353 (1873)5 lxx. partiape 230
(1874).

Capreolus europaeus, Gloger, Handbuch Naturgeschichte, p. 140 (1841) ;
Sundevall, K. Svenska Vet. Ak. Hand. for 1844, p. 184 (1846).

Capreolus caprea, Gray, List Mamm. Brit. Mus. p. 917 (1843), Cat.
Ungulata Brit. Mus. p. 222 (1852), Cat. Ruminants Brit. Mus. p. 81 (1872) ;
Brooke, Proc. Zool. Soc. 1878, p. 917.

Capreolus fossils, Owen, Rep. Brit. Assoc. for 1843, p. 238 (1844), Brit.
Foss. Mamm. p. 487 (1846).

Capreolus caprea, Bell and Alston, British Quadrupeds, p. 362 (1874) ;
Flower and Garson, Cat. Ost. Mus. Coll. Surg. part ul. p. 318 (1884) ;
Ward, Records of Big Game, p. 53 (1896) ; Satunin, Zool. “fahrb. Syst. vol.
ix. p. 310 (1896).

Plate XVIII

Characters.—Height at shoulder about 26 inches or rather more (a speci-
men in the British Museum from Dorsetshire measures 27 inches, as
mounted). Ears long, pointed, and comparatively thickly haired, both ex-
ternally and internally. Antlers only moderately rugose. Pelage in summer
thin, and at the early part of the season uniformly foxy red above, with-

Tom Deezer or Att LAnps Piuatre XVII.

EUROPEAN ROE IN SUMMER (1) AND WINTER (2) PELAGE.

Pablished by Rowland Ward Ltd

European Roe Db

out any white on the rump, but paler on the under-parts; a blackish
streak across the hairy portion of the muzzle, cutting the hinder part of the
nostril and running to the angle of the mouth, and some small whitish
markings in this region; front of face grizzled gray, darkest below the
eyes ; late in the season a bufish rump-patch produced, apparently by fading
of the hair. Winter pelage very much thicker and coarser ; general colour
speckled yellowish gray-fawn, the hairs
being gray at the base and rufous at
the tips ; a patch of pure white on the
rump, including the region of the tail,
but not extending markedly on to the
flank ; head, with the exception of the
markings on the face, uniform grizzled
gray. The metatarsal tuft is darker
than either summer or winter coat.
Albino varieties of the roe have been
recorded from Scotland and Germany.
Ten or eleven inches is about the
normal length for fair-sized horns, the
maximum recorded by Mr. Rowland
Ward being 13 inches. A pair from
Austria measuring 154 inches in length
are, however, mentioned in Bell and
Alston’s British Quadrupeds. Occasion-
ally specimens are met with in which

the front, or lower prong of the main

: a. ; Fic. 60.—Head of Siberian Roe.
fork is divided, thus producing four (Rowland Ward, Records of Big Game.)

tines; and malformed antlers are of
very frequent occurrence. The first antlers form a simple spike, and the
second pair are forked once, with the hinder prong the longer ; the third
pair being three-tined. Young roes are fully spotted with white.
Distribution.—In suitable localities over the greater part of Europe as far as
and including the Caucasus, and probably Asia Minor, but the exact easterly
limits not determined. Although unknown in Ireland, the species occurs in
England and Scotland, the south of Sweden, France, Germany, Austria,
Hungary, Spain, Tuscany, Greece, Turkey, and Northern Palestine. In the
Ze

226 Roes

British Isles wild roe are found in Scotland and some of the northern English
counties, although they have long since been killed off from the south of
England. Their remains are, however, abundant in the fens of Cambridge-
shire and Lincolnshire, and have also been discovered in several English
bone-caverns, as well as in the older ‘‘ forest-bed” of the Norfolk coast,
which belongs to the early part of the Plistocene period. In the early
part of the century they were reintroduced into Dorsetshire, where they
now flourish in the woods of the south side of the Blackmoor Vale ; and
some have been turned loose into Epping Forest and other districts in the
home counties. Dr. Satunin states that the roe of the Caucasus is C.
pygargus, but a small herd from that district at Woburn Abbey belong to
the European form.

The roe is doubtless the animal known to Aristotle as prox, although in
scientific literature that name has been applied to the muntjacs.

Habits —The ‘bounding roe,” as this deer is often termed, describes
one of its most characteristic habits, as, at least when first starting, it
always progresses by a series of long bounding leaps, with the head carried
high in the air. Roe are generally found in pairs or small family portions,
but never in herds. In Argyllshire Messrs. Harvie-Brown, and Buckley '
state that a birch-clad glen of three or four miles in length rarely contains
more than from two to four at any one time. Large woods with plenty of
under-covert adjacent to grass or arable land are the haunts where they are
most generally found. From these they sally forth to feed as evening
approaches with great punctuality, usually making regular tracks to and
from their feeding-grounds. In Scotland they are more abundant in
plantations than in natural copses. They take readily to the water, both of
their own accord and when pursued. After the bounding gallop in which
they start off, roe generally settle down to a kind of trot, and at no time is
their pace very great. Their leaping powers are, however, almost un-
rivalled for their size. ‘The cry has been described as a kind of harsh bleat.
Like most of the deer tribe, roe will both graze and browse, and they are
also stated to eat fungi in autumn. The antlers of the old bucks are shed
about the end of the year, and the new pair generally free from the velvet
towards the latter part of February. The pairing-season does not, however,
take place till July or August, the fawns being born about the following

| Vertebrate Fauna of Argyll and the Hebrides, p. 33 (1892).

Siberian Roe ZI lig,

May. ‘There may be either one or two of these latter produced at a
birth ; and the older writers state that in the case of twins they are
always male and female. The buck strictly confines his attentions
to a single doe; and it appears that the union is renewed for several
seasons, if indeed it does not last for life. In defending her helpless
young, which are carefully concealed among thick covert, the doe displays
great boldness.

When their curiosity is specially excited, roe exhibit even more bold-
ness than the generality of deer, as is well illustrated by the following
anecdote vouched for by Messrs. Harvie-Brown and Buckley in the volume

b)

already cited. ‘‘For nearly twenty minutes,” they write, “‘a buck, close
to Glenborrodale Castle, Ardnamurchan, stood within fifteen yards of
where three gentlemen were seated. They carefully abstained from mov-
ing in the slightest degree. ‘The buck stamped his feet, and went round
them in half circles, and not until he at last got wind of them did he fly.
Then he rapidly wheeled round, and with a snort of alarm disappeared in
the thick wood.”

Nomenclature—W ith regard to nomenclature, the European roe by
many modern naturalists would be termed Capreolus capreolus. Not
admitting this combination, the earliest post-Linnean name applied to the
living form has been selected ; but this is antedated by the name /fossi/is
given to fossilised remains, and this accordingly has the right of priority,
although its application to a living species would be unsatisfactory, even if
it really rank as a specific title.

2. THe SIBERIAN RokE—CAPREOLUS PYGARGUS

Cervus pygargus, Pallas, Reise Russ. Reichs, vol. i. p. 453 (1771) ;
Noack, Humboldt, vol. viii. p. 7 (1889).

Cervus ahu, Gmelin, Reise Russ. vol. ili. p. 496 (1780).

Cervus (Capreolus) pygargus, H. Smith, in Griffith’s Animal Kingdom,
MOliv. Pwl22, 7. p, 2h4 (0827):

Capreolus pygargus, Gray, List Mamm. Brit. Mus. p. 176 (1843), Cat.
Ungulata Brit. Mus. p. 223 (1852), Cat. Ruminants Brit. Mus. p. 82 (1872) ;
Gloger, Handbuch Naturgeschichte, p. 141 (1841) ; Fitzinger, SB. 4k. Wien,
Pole yeieepatt tap. 25 9) ( 1079),,Ixxe part 1. p. 248 (1874) ; Brooke, Proc.

228 Roes

Zool. Soc. 1878, p. 917; Ward, Records of Big Game, p. 54 (1896) ; Satunin,
Zool. “fahrb. Syst. vol. ix. p. 310 (1896).

Characters.—Closely allied to the last, but of larger size, the height at
the shoulder, according to Pallas and Brandt, varying from about 28 to 34
inches. Ears relatively shorter, wider, less pointed, and much more
thickly haired, both externally and internally. Antlers larger, more
divergent, and much more rugose on the inner border, where they form a

series of small irregular, nodular snags. Winter pelage much thicker and

S WA TYPE

Fic. 61.—Antlers of Siberian Roe. From a photograph lent by Mr. Carl Hagenbeck.

rougher than in the European species, being quite shaggy on the sides and
lower portion of the head, chest, and under-parts; the back, which is
coloured a mixture of yellowish and grayish brown, is more distinctly
speckled with blackish, and the white rump-patch extends in a short V on
to the flanks. The summer coat is a brighter and lighter rufous, with the
hairs lying more smoothly ; when first donned it shows little or no signs of
a light rump-patch, but, apparently by fading, a yellowish white disk
gradually develops in this region as the season advances. ‘The face-mark-

ings are generally similar to those of the European species. Other features

Siberian Roe Z210

are the slightly larger size of the face-glands, the more distinct development
of the tail, and the shorter and broader hoofs.

In referring to this form, Sir Victor Brooke writes as follows :—‘‘ Most
authors have considered C. pygargus to be a large variety of the common
roe. There are, however, in the British Museum two specimens purchased
from Brandt, and said to be from Siberia, which, in their very much larger
size, larger anal disk, and much more hairy ears, differ widely from all
specimens of the common species. Specimens of roe from Mantchuria in

Fic. 62.—Abnormal Antlers of Siberian Roe. From a photograph lent by Mr. Carl Hagenbeck.

the British and Paris Museums are very much smaller than the above-
mentioned specimens ; they also differ from the larger specimens and the
common roe in some details of coloration. It is not improbable that the
Mantchurian roe may prove to be a third modification of the form.”

With regard to Brandt’s specimens in the British Museum, one formerly
exhibited in the mammal gallery is not, as mounted, so tall as a roe
from Dorsetshire in the Museum, as it measures only 26 inches at the
shoulder. This is, however, very probably due to faulty taxidermy, as the
skin is evidently that of a much larger animal, and appears set up in too
bulky a form. In all its characters it agrees with the descriptions given by

2.30 Roes

Pallas, Brandt, and Noack, and undoubtedly indicates a distinct form,—
whether a species or sub-species being immaterial.

The antlers, which are always recognisable by their great rugosity and
the rugged processes on the inner edge of the beam, are subject to great
variation. In some cases, as shown in Fig. 62, they develop a kind of
palmation on the hinder branch of the main fork, the front branch of
which may be forked. They then exhibit a decided approximation to the
American deer of the Blastoceros group of Mazama, to which genus there is
reason to believe the roes are related.

A large, long-legged hind of this species from Siberia living at Woburn
Abbey in 1897 showed a very bright summer coat, with no trace of the
white caudal patch at the beginning of the season. Roughly speaking,
the size of this hind might be compared with that of a fallow deer.

The following dimensions of antlers which may be safely assigned to
this species are recorded by Mr. Rowland Ward, all being apparently from

Siberia :—
Length along Outer Curve. Basal Circumference. Tip to Tip.
3
15 44 ?
3
1476 4 ?
3
132 2 6
3 i
I 38 4 93
2 4 8f

Distribution.—In suitable localities from the Altai range and the
mountains of Turkestan to Siberia; probably also the mountains of the
Caspian provinces of Persia, where it is known as the ahu. In Siberia not
extending so far north as some of the species of Cervus, the northern limit
being about the 53rd or 54th parallel of latitude, and not ranging as far as
the mouth of the Amur river. During winter, migrating south into
Manchuria, and apparently Corea. The alleged occurrence of this roe in
the Caucasus is referred to on page 226.

As regards the above-mentioned localities, a skin of a big female roe
from Corea recently obtained by the British Museum appears too large for
the next species, and is accordingly referred to the present form. The
pelage is that of winter; the general colour being a pale brownish fawn,
with an irregular broad darkish band running down the middle of the back,

Manchurian Roe oie

and expanding as it joins the pure white tail- patch, the under-parts being
decidedly lighter than the back.

Habits.—The following particulars of the habits of this species in
Siberia, partly furnished by Herr Dorries, have been published by Professor
Noack.! According to these notes, the replacement of the winter by the
summer pelage generally takes place about the latter part of April, although
dependent to a certain extent on the season ; and by the end of September
the winter dress is once more resumed. The pairing-season is somewhat
later than in the European species, not taking place till September ; and
during this period the bucks utter a louder and deeper toned cry than the
former. Indeed, as might have been expected from its superior size and
weight, the Siberian roe is an altogether more courageous animal than its
Western cousin, and will not hesitate to charge dogs. In summer it affects
copses and open meadows, where it can wallow in the marshes or swim in
the neighbouring lakes ; but as winter comes on it retires to the protected
mountain forests, and is fond of consorting with the herds of larger deer
which then seek the same asylum. When the snows of November fall, the
roes themselves commence to collect in herds, which may number from
300 to 500 head, and soon after migrate southwards into Manchuria,
whence they return about the end of March or beginning of April. On
the Ussuri, which they must cross, they are at this season slaughtered in
thousands by the hunters, without regard to age or sex. In the year 1882,
however, when snow only fell locally, no such migrations took place, and
the roes only collected in small parties, and not in large herds.

3. Tue MAnNcuuriAn RoE—CAPREOLUS MANCHURICUS

Cervus pygargus mantschuricus, Noack, Humboldt, vol. viii. p. 9, fig. 12
(1889).

Characters. —Considerably smaller than the last, and more like the
European species, but distinctly red in winter. Antlers more slender than
in the Siberian roe, although with the same general form ; and the winter
pelage shorter. According to Professor Noack, the coloration (? in winter)
is as follows :—Forehead whitish yellow, nose umber-brown, region
round the eye very light, cheek reddish yellow, neck yellowish red umber-

1 Der Weidmann, 21st August 1891, p. 419.

23% Roes

brown ; ear more yellowish gray than in the European roe, but the colora-
tion of the lips very similar, with the exception that the dark spot on the
lower one is very small, and the white band on the upper one broader.
Specimens at Woburn Abbey were distinctly red in the middle of winter,
and thus quite unlike the European roe at the same season. The probable
distinctness of this form was first pointed out by Sir Victor Brooke,
although I have been unable to identify the skin in the British Museum to
which he refers (p. 229), and have had no opportunity of seeing the Paris
specimen, or specimens. A roe from Manchuria (from the same district
as the Pekin sika) living in the menagerie at Woburn Abbey in 1897 was
a much smaller animal than the Siberian species. Two pairs of antlers
in the British Museum from Manchuria recorded by Mr. Rowland Ward
respectively measure 134 and 11% inches in length ; the former being only
a little larger than fine specimens of the European roe.

Distribution.—The mountains of Manchuria. According to Professor
Noack, these roe never come down into the plains, neither do they make
the southerly winter migrations undertaken by the preceding species in
Siberia ; and they never collect in large herds.

4. Tue PriioceNne Rozr—Capreo us cusanus (Extinct)

Cervus cusanus, Croizet and Jobert, Oss. Foss. Puy-de-Déme, Cervide, plate
viii. (1828) ; Dawkins, Quart. “fourn. Geol. Soc. vol. xxxiv. p. 404 (1878).

Cervus (Capreolus) cusanus, Depeéret, Bull. Soc. géol. France, ser. 3, vol.
xii. p. 270 (1884).

Cervus (Capreolus) neschersensis, Deperet, op. cit. p. 272 (1884).

Capreolus cusanus, Lydekker, Cat. Foss. Mamm. Brit. Mus. part i. p. 75
(1885).

Characters.—Apparently of the approximate size of the European roe,
but with the antlers more flattened (? partly the effect of crush), rugose
only at the base, with the main fork nearer the middle of the length, and
the two tines of the second fork less divergent. The length (apparently in
a straight line) of an antler figured by Professor Dawkins is 11.4 inches,
and its basal circumference 2.8 inches. The same writer considers that this
species is the direct ancestor of the European roe, and that it 1s itself the

descendant of a still earlier species (C. matheroni) from the lower Pliocene

Milou Deer 223

deposits of Greece and France. In the latter kind the main fork of the
antler is placed still lower down, so that the undivided portion of the beam
is less than a third the total length of the antler; the front prong of the
main fork being relatively longer. I can find no sufficient characters for
regarding C. neschersensis, which occurs in the same beds, as distinct from
the present form.

Distribution.—F rance during the later part of the Pliocene period, the
typical localities being Perrier and Ardé in the Department of Puy-de-Déme.

X. THe Mitov DEER—GENUS ELAPHURUS

Elaphurus, Milne-Edwards, Ann. Scr. Nat. ser. 5, vol. x. p. 380 (1866) ;
Gray, Cat. Ruminants Brit. Mus. p. 82 (1872) ; Fitzinger, SB. 4k. Wien, vol.
Ixx. part i. p. 328 (1874) ; Brooke, Proc. Zool. Soc. 1878, p. go6, as a sub-
genus of Cervus.

Characters.—Lateral metacarpal bones as in Cervus, that is to say,
represented by their upper extremities. Antlers large and cylindrical, with
the beam making a marked angle with the plane of the face ; no brow-tine
or sub-basal snag, the beam dichotomously forking at a comparatively small
distance above the burr ; the front, or upper prong of the main fork curving
forwards and again dividing once or more, the posterior, or lower prong long,
straight, simple, and projected backwards. Muzzle with a large naked
portion, which is deeper and broader below the nostrils than in the elaphine
group, but extends only a little on to the front of the face, where its upper
border is deeply concave ; ears small and narrow ; tail very long, cylindrical,
and bushy at the extremity ; neck maned in the male ; face long. Pelage
of adult uniformly coloured throughout the upper-parts ; in young spotted.
No tarsal tuft. Metatarsal tuft continuous and situated in the upper third
of the metatarsus. The gland-pit in the skull large, and the face-gland also
large. Main hoofs large and spreading ; lateral hoofs very large. Upper
canines small ; upper molars moderately tall, with a small additional column
on the inner side. ‘The vomer not dividing the inner aperture of the nostrils
on the under surface of the skull into two moieties. Size large; build
heavy, with the limbs stout. No foot-glands.

Much difference of opinion has existed as to the systematic position of

2H

2.34 Milou Deer

this genus, of which only a single specific representative is at present known.
By its describer, Professor A. Milne-Edwards, it was regarded as a very
distinct form ; and this view was maintained by the late Dr. Gray, who
placed it between the roes and the American deer. Professor Fitzinger
likewise regarded it as entitled to generic distinction, but located it in the
neighbourhood of the rusine deer. On the other hand, Mr. Sclater, while
separating the fallow deer as a genus apart, includes E/aphurus in Cervus ;

t fk
Ne!

‘n s
Stary ae))Z55

Fic. 63.—Pére David’s Milou Deer in the Park at Woburn Abbey. From a photograph by the
Duchess of Bedford.

and his view was followed by Sir Victor Brooke, with whom most other
subsequent writers have agreed. Mr. A. Gordon Cameron, basing his
conclusions on the structure of the antlers, has, however, revived the view
of Dr. Gray, and there is little doubt that he is correct in so doing. If
antlers count for anything in classification—and it is almost impossible to
deny that they do so—the genus has nothing to do with any of the living
Old World deer with the exception of the roes, while its alliance with the
American deer seems close. Probably it is still more closely connected with
the under-mentioned extinct types. That it differs from the American deer

Characters ae

in the structure of the vomer and the lateral metacarpal bones must be
admitted ; but the backward production of the former bone in the American
deer may probably be regarded as a specialised feature, and, as already said,
I attach only minor importance to the lateral metacarpals. All who have
seen the animal alive can scarcely fail to recognise how unlike it is, both in
appearance and gait, to all the Old World deer of the present epoch ; and it

Fic. 64.—Five successive pairs of Antlers of a Pére David’s Milou Deer at Woburn Abbey. In 1897
this stag shed its antlers twice. Since the last pair photographed, it shed a pair like those of the

second year, and by Christmas had grown a new pair like the last pair in the figure. Photographed
by the Duchess of Bedford.

is probably more or less intimately related to the following extinct genus,
which there seems good reason for regarding as also related to the ancestral
stock of the modern American deer. To some of the latter the present
genus approximates by the unusual length of its tail. Its geographical
distribution also harmonises with its apparent affinity to the American deer,
a large number of animals from North-Eastern Asia being more or less
closely related to North American types.

Distribution —The eastern portion of the Eastern Holarctic region.

rw

26 Milou Deer

id

Pkre Davip’s Mirovu DEER—ELAPHURUS DAVIDIANUS

Elaphurus davidianus, Milne-Edwards, Ann. Sct. Nat. ser. 5, vol. X, Px sooy
and Arch. Mus. Paris, vol. 11. p. 27 (1866) ; Gray, Cat. Ruminants Brit. Mus.
p. 82 (1872) ; Fitzinger, SB. 4k. Wien, vol. 1xx. part i. p. 329 (1874).

Cervus davidianus, Sclater, Trans. Zool. Soc. vol. vii. p. 333 (1871) ;
Brooke, Proc. Zool. Soc. 1878, p. 906; Ward, Records of Big Game, p. 17

(1896). Plate XIX

Characters.—Height at shoulder about 3 feet 9 inches, or nearly the size
of a large red deer. Head large, with small eyes and ears, and a long,
narrow muzzle; limbs stout. Hair generally short and smooth, but longer
on the middle line of the chest and under-parts, and on the neck and throat
of adult males forming a mane. General colour of upper-parts reddish
tawny with a tinge of gray, passing through an ill-defined darker band on
the sides to a more decided whitish gray on the under-parts ; the neck,
chest, and lower portion of throat dark brown ; on the neck and fore part
of the back a blackish brown longitudinal stripe, and a similar stripe on the
chest ; rump and inner side of thighs yellowish white, passing gradually into
the general colour of the body ; outer side of upper part of limbs similar to
the back, inner side and lower portion whitish yellow-gray ; tail like back,
except the terminal tuft, which is blackish brown ; the face brownish, with
a whitey brown patch round the eye, bordered on the inner side by a dark
brown streak; chin and inside of ears white, and a whitish oblique streak
on the sides of the muzzle, above the nostrils. Female somewhat lighter
coloured. Young reddish brown with a tinge of yellow, at first spotted
profusely with white.’

The following dimensions of antlers are recorded by Mr. Rowland
Ward :—

One Gis bicpuentea Dip tO Tibe aes tiore peat
324 62 138 184 8-8
304 5 3541's 351'6 TiS?
284 5% 265 nor 333
27 58 204 ? 6-5
22 42 ? P 4-3
1 Brooke suggested that the young were unspotted, but this is disproved by the specimens born at

Woburn Abbcy.

Tae Deer or Att Lanpos. Prater XIX.

PERE DAVID’S DEER.

ibiished. by Rowland Ward. Led

|

Pere David’s Milou Deer 207

Distribution —Northern China. The type specimen was obtained by
Pére David from the Imperial Park at Pekin, and is now preserved in the
museum at Paris. The London Zoological Society received a pair of
specimens from the late Sir Rutherford Alcock in 1869, and a second pair
were purchased in 1883. The male figured in plate xix is one living in
the park at Woburn Abbey.

Habits —By the Chinese this deer is known as mi-lou, a name which
may be adopted as the English title for the genus. Nothing appears to be
recorded of its habits in the wild state, and the following notes are from
observations on the specimens at Woburn Abbey. In his notice of this
species Mr. Sclater wrote as follows :—‘ The general aspect of the elaphure
is much more like that of the true Cerv than I had anticipated from the
description and figure of M. Milne-Edwards. The only two very noticeable
points of distinction, besides the horns of the male, which are not at present
shown in our animals, are the rather larger, heavier legs, the longer and
more expanding toes, and the long tail. The latter character, however,
seems to me to have been somewhat exaggerated in M. Milne-Edward’s
figures—the tail in our specimen not nearly reaching the hocks, and, though
of somewhat different form, being little, if any, longer than that of the
fallow deer and some of the American deer (such as the Virginian deer).”
With these conclusions it is very difficult to agree, the general appearance of
Pére David’s deer, when roaming at liberty in the park at Woburn Abbey,
being quite unlike that of any other member of the group. Its gait is a
kind of “lolloping” trot, recalling more the action of a mule than that of
a deer ; and its whole carriage, although almost impossible to describe, is
totally unlike that of any other deer. In the Woburn specimens the tail
reaches the hocks, and is decidedly longer than that of the fallow deer,
The large spreading hoofs proclaim that these animals are in the habit of
frequenting marshy ground, and at Woburn during summer they may
frequently be seen wading far into the lakes, or even swimming in the
deeper water. At this season their food consists chiefly of rushes and various
other water-plants. The stags call in June and July ; their cry being a kind
of bray, more like that of a donkey than the call of other deer. The antlers
are dropped during November or December.

Extinct Group

w)
(>
CO

XI. Genus Anoctocuis (Extinct)

Anoglochis, Croizet and Jobert, Oss. Foss. Puy-de-Déme, Cervide, descrip-
tion of plate 11 (1828), as a sub-genus'; Pomel, Cat. Méthodique, p. 107
(1853).

Polycladus, Gervais, Zool. et Pal. Frangaises, 2nd ed. p. 146 (1859), nec
Polyclados, Brandt, 1835.

Eucladoceros, Falconer, Pal. Mem. vol. ii. p. 472 (1868), as a sub-genus.

Characters.— Lateral metacarpals unknown; antlers large, without a
brow-tine, but with a sub-basal snag, above which the beam is dichotomously
forked, one or both prongs of the fork again dividing once or more, and
the whole antler forming a marked angle with the plane of the face ; vomer
unknown.

Although their conclusions have been rejected by M. Deperet, it
appears evident that Messrs. Croizet and Jobert were perfectly justified in
dividing the French Pliocene deer into a brow-antlered (Catoglochis) and
fork-antlered group (Azoglochis), and there seems no good reason for
separating Dr. Falconer’s Euc/adoceros from the latter of these, which was
typified by Cervus ramosus. In all these deer, what has usually been
described as the brow-tine of the antlers, is clearly comparable with the sub-
basal snag of many of the existing species of Mazama, as may be seen by a
comparison of the figures ; and it is not till a greater or smaller distance
above the origin of this snag that the antler makes its regular dichotomous
fork. That these deer are generically distinct from E/aphurus seems
probable from the conformation of the antlers ; but, although it is perhaps
unlikely that they are generically identical, yet until the nature of the
lateral metacarpal bones and the relations of the vomer are known, it seems
impossible to formulate a definition from the antlers alone by which they
can be satisfactorily separated from the American Mazama. ‘These deer
indicate the probability of Mazama having sprung from an Old World
type.

Distribution —Western Europe during the upper Pliocene and early
part of the Plistocene period.

' The text of this portion of the work was never published, and the generic and specific names

appear only on the original covers,

Ardé Deer 2.39

1. THE Arp& DEER—ANOGLOCHIS ARDEUS (Extinct)

Cervus ramosus, Croizet and Jobert, Oss. Foss. Puy-de-Déme, Cervide, plate
iv (1828) ; Deperet, Bull. Soc. géol. France, ser. 3, vol. xil. p. 256 (1884) ;
nec de Blainville, fourn. Physique, vol. xciv. p. 276 (1822).

Fic. 65.—Antlers of Ardé Deer in three stages of development. 4, sub-basal snag ; 4, lower prong
of main fork ; c, upper do. After Depéret.

Cervus ardeus, Croizet and Jobert, of. cit. plate ii (1828) ; Deperet,
Op.tit. p. 255 (1884).

Cervus polycladus, Gervais, Zool. et Pal. Frangaises, 1st ed. p. 82
(1848-52).

Cervus ambiguus, Pomel, Cat. Méthodique, p. 107 (1853).

Cervus cladoceros, Pomel, op. cit. p. 108 (1853).

Cervus cylindroceros, Dawkins (ex Bravard), Quart. ‘fourn. Geol. Soc.
vol. xxxiv. p. 414 (1878) ;, Lydekker, Cat. Foss. Mamm. Brit. Mus. part
be pe ULI (18's).

240 Extinct Group

Cervus borbonicus, Depéret (ex Bravard), Bull. Soc. géol. France, ser. 3,
vol. xii. p. 260 (1884).

Characters.—Antlers of medium size with the beam rising for a short
distance straight till it gives off a short sub-basal tine, then curving strongly
and continuing undivided for a considerable length, after which it forks
dichotomously, and each prong of the fork thus formed again divides. In
younger specimens the two prongs of the main fork are undivided ; but in
very old individuals the complexity is greater.

I believe that C. ardeus, which was founded on an antler in which the
beam palmates and divides into three tines, instead of forking dichotomously,
is merely a somewhat abnormal form of the antlers described as C. ramosus.
There appear no suthcient characters by which the C. borbomicus of M.
Deperet, which is the same as the C. cy/indroceros of Professor Dawkins,
differs from immature antlers of the present form. While C. ramosus and
C. ardeus are assigned by M. Deperet to a separate extinct group (Po/ycladus),
C. borbonicus (cylindroceros) is referred both by him and Professor Dawkins to
the rusine group, with which it has obviously no affinity. M. Deperet
has suggested some relationship between the present species and both the
American deer and Sedgwick’s deer.

By comparing the figures here given with those of the antlers of the
Virginian deer in Fig. 69, it will be seen that the two agree in the position
and size of the sub-basal snag, in the curvature and length of the beam
before forking, and also in the splitting of both prongs of the main fork ;
and they may therefore be regarded as occupying analogous positions in
their respective genera.

Distribution —France, and perhaps other parts of Europe, during the
latter portion of the Pliocene period, the chief localities being Arde,
Bourbon, and Perrier, in the Department of Puy-de-Déme.

2. SepGwick’s DrEER—ANOGLOCHIS SEDGWICKI (Extinct)

Cervus (Eucladoceros) sedgwickit, Falconer, Pal. Mem. vol. 11. p. 472
(1868) ; Depéret, Bu//. Soc. géol. France, ser. 3, vol. xii. p. 259 (1884).
Cervus dicranmius, Riitimeyer (ex Nesti), 46h. schweiz. pal. Ges. vol. vil.

plate 1 (1880) x. p. 106 (1883).

Sedgwick’s Deer 241

Cervus sedgwickt, Lydekker, Cat. Foss. Mamm. Brit. Mus. part i. p. 93
(1885).

Characters. — Antlers very large and complex, giving off a long,
upright, forked, sub-basal snag a short distance above the burr, the
beam soon after forking dichotomously, the upper prong of the fork
again dividing, with one branch simple and the other forked, and the

Fic. 66.—Skull and Antlers of Sedgwick’s Deer. The innermost upright forked prong is the sub-basal
snag, a short distance above the origin of which the beam splits into the main fork. In life the
lower branch of this fork probably curved forwards. From Riitimeyer.

lower prong of the main fork also splitting and both its branches
dividing.

In the shorter length of beam between the giving off of the sub-basal
snag and the main fork this magnificent species comes nearer to the
mule-deer type, although the sub-basal snag is much longer. If the
accompanying figure be compared with the front views of the antlers
of the Virginian and mule-deer, the homology of the various elements
will be clearly evident ; and it is difficult to believe that the two forms
are not related.

Distribution England and Italy during the latter part of the Pliocene
and early portion of the Plistocene periods, the British remains occurring in
the Norfolk forest-bed.

72M

2.4.2 Extinct Group

3. Tue Five-rinep DrrrR—ANOGLOCHIS TETRACEROS (Extinct)

Cervus tetraceros, Dawkins, Quart. ‘fourn. Geol. Soc. vol. xxxiv. p. 416
(1878) ; Lydekker, Cat. Foss. Mamm. Brit. Mus. part ii. p. 113 (1885).

Characters.—Antlers smaller and simpler than in the last species, giving
off a long, undivided sub-basal snag a short distance above the burr, the
beam soon after forking and probably curving forwards ; the lower prong
of the main fork simple, the upper divided, and its anterior division again
splitting.

This species was established upon the evidence of a series of shed
antlers in the British Museum which are crushed flat, and thus give the
impression of a straight beam carrying four upright tines on its upper
surface. The restoration is consequently somewhat difficult, but there
seems little doubt that the first tine represents the sub-basal snag of the
antlers of the Virginian deer, and that the beam was then suddenly bent
forwards. It would thus show four upwardly projecting tines, of which
the first two would be the prongs of the main fork. The annexed figure
has been formed by taking the adult antlers and drawing them with the
same curvature as those of the Virginian deer, and a comparison with the
figures of the latter will leave little doubt as to their essential similarity.
Indeed, they might well belong to the same genus. That these antlers
are also structurally similar to those of Sedgwick’s deer, will likewise be
apparent from a comparison of the respective figures, the difference being
that whereas in the present species each of the five tines is simple, in
Sedgwick’s deer all but the terminal one bifurcate; the one type being
evidently only an earlier and simpler form of the other. In younger
examples of the antlers of this species there are only four tines, and in still
younger ones three ; that is to say, each antler consists of a short sub-basal
snag and a simple dichotomous fork, as in young antlers of the Virginian
deer. As age advances the sub-basal snag increases in proportionate size
till it becomes nearly as large as the true tines. Very young antlers
are simply forked, and are then strikingly like those of young American
deer. In his original description Professor Dawkins rightly compared the
present form with the Virginian deer, remarking that the two were dis-
tinguished by the different position of the ‘ brow-tine,” that is to say, the

American Deer 2.4.3

sub-basal snag ; and adding that the fossil antlers curved forwards some-
what after the manner of those of the living species. In a restoration
subsequently attempted they are, however, represented as inclining back-
wards.! About 27 inches is the length of the largest antler.

Distribution —France, England, and probably other parts of Europe,
during the latter part of the Pliocene and the early portion of the
Plistocene period ; the type specimens being from Peyrolles in the Puy-
de-Déme, and the English examples occurring in the Norfolk forest-bed.

Fic. 67.—Restoration of Antlers of Five-tined Deer. a, sub-basal snag ; 4, lower prong of main fork ;
c, upper prong do, ; d, terminal tines.

XII. Tue American DEER—GeENus MazAMA

Mazama, Ratinesque, Amer. Month. Mag. vol. i. p. 44 (1817).

Characters.—Lateral metacarpal bones as in Rangifer, that is, with their
lower extremities remaining. Antlers very variable in size, the beam
forming a marked angle with the plane of the face, and without a
brow-tine ; when consisting of more than a simple undivided prong,
dichotomously forked, frequently with a sub-basal snag, and always with
the lower, or anterior prong of the main fork projected from the front
edge of the beam, in some cases the lower, in others the upper (posterior),
and in others both prongs again dividing. Muzzle with a large naked

" Dawkins, Early Man in Britain, p. 86, fig. 18 (1880).

24.4 American Deer

portion ; ears and tail variable. Pelage of adult uniformly coloured ; in
young usually spotted. A tarsal tuft present. Metatarsal gland and tuft,
when present, very variable in form and position.’ Gland-pit in the skull
and face-gland very variable. Main hoofs long and pointed ; lateral hoofs
well developed. Upper canines present or absent ; upper molars with or
without a small additional column on the inner side. The inner aperture
of the nostrils on the under surface of the skull divided by the thin
longitudinal plate-like bone known as the vomer into two distinct
chambers. Size medium or small. |

All naturalists are agreed that the American deer form a very natural
group, connected together by many common features, and differing in
several respects from all the existing Old World deer. Different opinions
are, however, held as to whether they should all (with the exception of
the pudus) be included in a single genus, divided into several sub-generic
groups, or whether such minor groups should be allowed the rank of
genera. Gray and Fitzinger adopt the latter alternative, and Sir Victor
Brooke the former. Mr. Sclater, in the List of Ammatls in the Zoological
Society's Gardens, follows a middle course, including by far the greater
number of the species in Carzacus, but separating the guemals as Furcifer.

To allow the minor groups the rank of distinct genera destroys to a
considerable extent the unity of the whole assemblage, and thus renders its
distinctness from all the existing Old World deer less apparent. And if, as
is done in this volume, the typical brow-tined deer of the Old World are
included in the single genus Cervus, then the groups into which the
American deer are divisible should likewise be brigaded together in the
single genus Mazama. If, on the other hand, it is thought advisable to allow
the various groups into which Cervus is divided, such as Pseudaxis, Dama,
Rusa, etc., to rank as genera instead of sub-genera, then the same course
should be adopted in the case of the American deer, and the various groups
here ranked as sub-genera should stand as genera. If this course be
adopted, it would be necessary to divide the sub-family Cervine into
several sectional groups, as otherwise the various genera would have very
different values, some being very nearly related, and others much more
widely separated.

! Brooke, Proc. Zool. Soc. 1878, p. 918, states that the metatarsal tuft is always situated in the
lower third of the cannon-bone ; whereas in M. co/umbiana it is solely in the upper half.

Characters 2.4.5

Including, then, all these exclusively American deer in one genus, the
next question is the name to be used. Except one, the earliest 1s Mazama
of Rafinesque, but as this name included several animals, Dr. Baird in his
North American Mammals (p. 665) urged that it should not be used at all,
although it undoubtedly included some of the American deer. It was
employed by Hamilton Smith in 1827 for the Virginian deer group, the
brockets being separated as Swbu/o. According to Dr. C. H. Merriam,’ the
deer included by Rafinesque was one of the brockets, which by elimination
becomes the type of the genus. If used at all, the genus must therefore
be typified by the brockets ; but if, as is done here, all the true American
deer, except the pudus, are regarded as forming one genus, Mazama,
as the earliest satisfactory name, may be employed for the entire group.

As already mentioned, there are at present no sufficient means of dis-
tinguishing from Mazama, in its widest sense, the Old World fossil deer
constituting the genus Anog/ochis, although if the whole organisation of the
latter were known, such might be found. Anoglochis, it may be observed,
antedates all names except Mazama and one other.

As noticed in the introductory chapter, and also on page 216, it has been
generally considered that Mazama traces its origin to the extinct North
American Cosoryx and Blastomeryx, but the extinct fork-antlered deer of
Europe suggest that its origin was rather in the Old World, as it is difficult
to realise the independent origin of such essentially similar types. More-
over, if the American deer originated in the New World from forms with
permanent antlers, they must have developed the deciduous antlers in-
dependently of the deer of the Old World, which, as remarked by Messrs.
Scott and Osborn, is somewhat difficult to believe.

Whether they were developed in the Old World or the New, it is evident
that the American deer originated in the northern hemisphere, and that
they are comparatively modern immigrants into South America, where they
now attain their maximum development. For a long period, during
which the isthmus of Panama was non-existent, the fauna of South
America was quite unlike that of North America, and it was not till about
the commencement of the Pliocene epoch that a land-connection was
established which permitted the deer and other northern types of mammals
to flow into South America. That the simple-antlered South American

1 Science, ser. 2, vol. i. p. 208 (1895).

246 American Deer

brockets are not the ancestral types of the American deer is evident from the
fact that remains of these are unknown in North America, and that species
with complex antlers of the type of those of the marsh-deer were already
in existence at or about the time communication was established between
the northern and southern halves of the New World; remains of such
species occurring in the Pliocene beds of Monte Hermoso, in Argentina.
Rather must the brockets, as has
already been pointed out by Mr. A.
Gordon Cameron, be regarded as
degraded, or arrested, types of the
group.

A large amount of misconception
has arisen with regard to the structure
of the antlers of this group. In 1872
the late Dr. Gray rightly termed the
single upright prong arising from the
inner side of the lower part of the
beam of the antlers of the Virginian
deer the ‘‘sub-basal snag,” but this
snag Sir Victor Brooke incorrectly
identified with the brow-tine of the
typical Old World deer. This error
has been pointed out by Mr. A.
Gordon Cameron in the following

Fic. 68.—Side view of Antlers of Virginian Deer, ney ;
from a specimen in the British Museum. words :—‘‘ These characteristic tines

The sub-basal snags are the long, upright have nothing in common with the

prongs on the extreme left, passing up be-
tween the beam of each side at its bend. true brows of Old World types, and
rise vertically from the inner side of
the beam, between the coronet and the main furcation, usually converg-
ing at the apex. They are subject, in common with the antlers that
produce them, to all kinds of eccentricities, are frequently forked and
sub-palmate ; and appear to bear an inverse relation to the calibre of the
posterior prong. They develop with the second antlers of C. /eucurus,
but only with the later antlers of C. macrotis.”
Regarding the different forms assumed by the antlers of the group, the

same observer writes as follows :—‘ In the dominant type of the north the

Characters 2.4.7

anterior prong tends to become heavier than the posterior one, and the
reduction of the calibre of the latter is accompanied by a proportionate
development of the sub-basal snag. In the dominant type of the south the
posterior prong tends to become heavier than the anterior one, and the sub-
basal snags are not developed at all.” The occasional forking of the sub-
basal snag is parallel by the condition normally prevailing in the extinct
Sedgwick’s deer ; and the occurrence of that snag in the latter species and
its allies, and its disappearance in the living Southern American forms is an
important indication of the apparent relationship between Anoglochis and
the northern types of Mazama. Moreover, it is only in these two latter
that the great preponderance in the development and forward curvature
of the anterior prong is noticeable. This ultra-development of the
front prong tends largely to obscure the main dichotomous forking
of the antlers, which is best displayed in those of the mule-deer and
marsh-deer.

The division of the hinder aperture of the nasal aperture in the dry
skull into two separate chambers by the vertical partition of the vomer
is, as was first pointed out by the late Professor Garrod, a character
possessed by the American deer and the reindeer, and one unknown
in all other members of the family; but it seems highly improbable
that it indicates any intimate relationship between the two genera
in question, being a peculiarity which might easily be developed
independently.

In the possession of a tarsal gland and tuft on the inner side of the hock,
the American deer differ from all the species of the genus Cervus. In such
of them as have a metatarsal gland and tuft situated on the outer side of
the metatarsus, this gland may be placed partially or wholly in the lower
third of that segment of the leg; and it is always of a more or less
elongated form, with the hairs forming a distinct fringe round the naked
gland, which may be exposed. In the mule-deer the fringe of hair
bordering this gland is stated by Mr. Caton to be capable of erection
during periods of excitement.

Distribution —The Western Holarctic, Sonoran, and Neotropical regions,
ranging from boreal North America to Patagonia.

248 American Deer

i. THE DorcELAPHINE GRoup—Sus-GEnus DorcELAPHUS

Mazama, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 314 (1827),
nec Rafinesque, 1817.

Dorcelaphus, Gloger, Handbuch Naturgeschichte, p. 140 (1841, title-page
dated 1842).

Cariacus, Lesson, Nouv. Table Hist. Nat. p. 173 (1842); Gray, Lust
Mamm. Brit. Mus. p. 175 (1843), Proce. Zool. Soc. 1850, p. 237,0Can
Ungulata Brit. Mus. p. 33 (1852), Cat. Ruminants Brit. Mus. p. 82 (1872) ;
Brooke, Proc. Zool. Soc. 1878, p. 918; Riitimeyer, 4bh. schweiz. pal. Ges.
vol. vill. p. 47 (1881).

Reduncina, Wagner, in Schreber’s Sdugethiere, vol. iv. p. 373 (1844), as a
sub-genus ; Fitzinger, SB. 4k. Wien, vol. Ixviil. part i. p. 357 (1873),
Ixxvill. part 1. p. 312 (1879).

Macrotis, Wagner, /oc. cit. (1844), as a sub-genus, nec Macrotis, De}. 1833.

Eucervus, Gray, Ann. Mag. Nat. Hist. ser. 3, vol. xviii. p. 338 (1866),
Cat. Ruminants Brit. Mus. p. 85 (1872).

Orelaphus, Fitzinger, SB. Ak. Wien, vol. Ixviii. part 1. p. 356 (1873), to
replace Macrotis.

Gymnotis, Fitzinger, op. cit. Ixxvili. part 1. p. 343 (1879).

Characters.—Antlers generally large and complex, with a sub-basal snag,
and the lower, or anterior, prong of the main fork more or less developed
at the expense of the upper, or posterior prong. Metatarsal gland and tuft
usually present. Tail long or moderate, and hairy below. Face very long
and narrow ; the face-gland very small, and the gland-pit in the skull of
moderate extent. Upper canines wanting. Size relatively large. Fawns
normally spotted.”

Distribution —The Western Holarctic, Sonoran, and northern portion of
the Neotropical region.

1 The priority of this name is pointed out by Mr. Thomas in Aan. Mag. Nat. Hist. ser. 6, vol. xv.
p. 193 (1895), but Mr. Bangs, Proc. Soc. Washington, vol. x. p. 25 (1896), observes that “as
Dorcelaphus is undoubtedly also antedated by other names, it seems advisable to keep the well-known
name Cariacus until this point is definitely settled.” The possibility of Anoglochis being the proper
term, is mentioned on page 238.

2 A Virginian deer hind at Woburn Abbey gave birth to a pair of fawns in 1897, one of which was
spotted and the other uniformly coloured.

Common American Deer 24.9

1. THE ComMMon AMERICAN DEER—MAzZAMA AMERICANA

Cervus dama americana, Erxleben, Syst. Regn. Anmatlis, p. 312 (1777).

Cervus virginianus, Boddaert, Elenchus Animalum, p. 136 (1785) ; Baird,
Mamm. N. America, p. 649 (1857) ; Caton, Antelope and Deer of America,
p- 100 (1877).

Cervus (Mazama) virginianus, H. Smith, in Griffith’s Animal Kingdom,
VOlmverpe 27, V. Pp. 315 (1827).

Cervus (Mazama) clavatus, H. Smith, of. cit. iv. p. 132, v. p. 315 (1827).

Mazama virginiana, Jardine, Naturalists Library—Mamm. vol. iii. p. 176
(1835).

Dorcelaphus virginanus, Gloger, Handbuch Naturgeschichte, p. 14.0 (1841).

Cartacus virgimanus, Lesson, Nouv. Table Hist. Nat. p. 173 (1842) ; Gray,
List. Mamm. Brit. Mus. p. 175 (1843), Cat. Ungulata Brit. Mus. p. 228
(1852), Cat. Ruminants Brit. Mus. p. 83 (1872), Hand-list Ruminants
Brie Visep. 05 (1873); Brooke, Proc. Zool. Soc. 1878, p. 919; Alston,
Biol. Centr. Amer.—Mamm. p. 115 (1879) ; Merriam, Mamma!s of the Adiron-
dacks, p. 107 (1884) ; Ward, Records of Big Game, p. 55 (1896).

Cervus clavatus, Sundevall, K. Svenska Vet. Ak. Handl. for 1844, p. 183
(1846).

Cervus capricornis, de Saussure, Rev. Mag. Zool. ser. 2, vol. xii. p. 252
(1860).

Reduncina virginiana, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 357
(io72) exvine pate it. 313 (1870).

Cervus (Cartacus) virginianus, Herrick, Mamm. Minnesota, p. 281 (1892).

Cartacus americanus, Bangs, Proc. Soc. Washington, vol. x. p. 25 (1896).

Dorcelaphus americanus, Rhoads, Proc. Ac. Philadelphia, 1897, p. 208 ;
Mearns, Proc. Soc. Washington, vol. xii. Pmz3 (1878).

Characters.—Build light and graceful, both body and limbs being long.
Height variable, ranging from about 3 feet 1 inch at the shoulder to 26
inches. Antlers with a long sub-basal snag, above the origin of which the
beam is abruptly curved forward, and soon after forks dichotomously, the
posterior prong of the main fork upright and generally undivided, the
anterior, or lower prong again forking, with its lower division also divided ;
the whole antler in advance of the sub-basal tine thus having the appear-
ance of a horizontal beam with three nearly vertical tines arising from its

Oak

250 American Deer

upper surface (Fig. 68). Ears relatively small and sparsely haired externally ;
tail long and pointed ; muzzle very long and slender; face-gland very
small, and almost hidden by a fold of skin. Metatarsal gland, when present,
small, sub-circular, and usually situated in the lower third of the cannon-
bone, its centre being bare and black, the surrounding fringe of hair pure
white, and this followed by an outer ring of fawn; tarsal tuft large and
whitish. General colour of summer pelage varying from bright rufous
chestnut to yellowish fawn or gray on the upper-parts, in winter some shade

Fic. 69.—Front view of Frontlet and Antlers of Virginian Deer. From a specimen in the British
Museum. The sub-basal snag is the innermost upright prong of each antler. (Rowland Ward,
Records of Big Game.)

of yellowish leaden gray, faintly speckled, and often with a tinge of rufous ;
under-parts, inner side of thighs and buttocks, part of inner and hinder
side of fore-legs, lower surface of tail, chin, throat, lips, a ring round
the eyes, and a band round the muzzle white. Fawn normally fully
spotted with white. The case of the numerous forms allied to the
typical Virginian deer is very similar to that of the sambar group, so
that there is an equal difficulty in deciding whether they should be
regarded as species or sub-species. In 1878 Sir Victor Brooke’ wrote
as follows in regard to this question :—“’The number of specimens from
well-authenticated localities at present existing in European collections is far

1 Proc. Zool. Soc. 1878, p. 920.

Common American Deer 251

too small to enable me to form any decided opinion respecting the degree of
persistency of modification in the above-mentioned forms. Mr. Allen* has
conclusively proved that a decrease in size and characteristic development
takes place in all mammals, the centre of whose area of distribution is in
North America, as they spread southwards ; and, so far as the materials at my
command enable me to judge, I think it probable that the characters upon
which Cariacus leucurus, C. mexicanus, C. savannarum, and C. peruvianus have
been established as species may be in large measure the result of this law.”
So far as specimens in the British Museum are concerned, the question
scarcely admits of a more decisive answer now than then. There can,
however, be no question but that all these forms are modifications of what
is essentially one type of animal, and they are accordingly here regarded as
sub-species rather than species ; an opinion shared by Mr. Rhoads, so far at
least as the Mexican form is concerned, and adopted by Mr. Caton for most,
and by Mr. A. Gordon Cameron for all the modifications. With regard to
their relationship and distribution Dr. P. Matschie writes as follows :—
“The two other kinds of Mazama deer found in South America are sufh-
ciently well distinguished from Cariacus peruvianus ; C. savannarum, which
probably inhabits the whole extent of the Orinoco area, is pure grayish
brown with white speckling; C. gymmnotis, from the Magdalena River dis-
trict, yellowish brown with grayer speckling. To the northwards, in
Central America, C. nemoralis replaces C. gymnotis ; then follows in Yucatan
and South Mexico C. ¢foltecus, C. mexicanus in Texas and Mexico, C.
leucurus in the Western, and C. virgimianus in the Eastern United States.”
Although several previous writers have referred to the decrease in bodily
size and the diminution of the antlers in the members of this group as they
are traced from north to south, they do not appear to have noticed two
other features. These are a gradual darkening, and the tendency to the
assumption of a permanently dark coat (gymmnotis and peruvianus), and also to
the disappearance of the metatarsal gland and tuft, which become rudi-
mentary in wemoralis, and are quite lost in éo/tecus, gymnotis, and peruvianus.
Both these are undoubtedly specialised peculiarities, the former being an
example of that tendency to blackness in many animals inhabiting humid,
and especially warm regions, to which reference has already been made
in the introduction. Glands are frequently found in animals living in

1 Bull. U.S. Survey, 1876, p. 304.

259 American Deer

herds, and serve by their scent to enable the members to keep together.
Virginian deer are certainly found in herds, and if some of the southern
forms should prove to be less gregarious, a reason for the abortion or
disappearance of the gland would be apparent.

Whether regarded as species or sub-species, the various members of this
group are best recognised by the peculiar characters of the antlers, the small
ears, long tail, and, when present, by the small white metatarsal tuft. The
antlers are subject to an extraordinary amount of individual variation, and
in some of the smaller southern forms are so modified from the normal type
as to be a little dificult of recognition, although they apparently always
retain the large sub-basal snag. The specimen of which a side view is
shown in Fig. 68 is a very characteristic one, and displays the great length of
the latter snag and the undivided upper prong of the main fork. In the
one of which the front view is represented in Fig. 69 the sub-basal snag is
shorter, and the upper prong of the main fork divided. Should the numer-
ous local variations be regarded in the light of distinct species, it would be
necessary, in order to preserve some degree of equivalence in the divisions
employed, to raise the mule-deer and the Columbian black-tail to the rank
of a sub-genus (Eucervus of Gray), an alteration which would entail further
modifications in the scheme of classification adopted throughout this volume,
which is to minimise divisions so far as possible.

Distribution. —North America to Peru, Bolivia, and Guiana.

a. VIRGINIAN Race—MazaAMA AMERICANA TYPICA
Plate XX

Characters.—Size large, the height at the shoulder reaching to about
3 feet 1 inch. Antlers large. Usual colour of pelage of upper-parts in
summer bright rufous chestnut, with black markings on the face and tail ;
in winter speckled yellowish gray ; a transverse black band on the chin ;
tail chestnut or gray above, white beneath, with more or less black at the
tip on the upper surface. Metatarsal gland well developed, and situated
low down on the leg. Even in this race there appears to be a considerable
degree of colour variation. Mr. Caton observes that while in some examples
there is no appreciable difference in the colour of the upper-parts and of
the outer sides of the limbs between this race and the Columbian black-tail,

Tee Deer or ALi LANDS. Puatm XX.

VIRGINIAN DEER IN WINTER PELAGE.

Published by Rowland Ward Ltd .

—

Virginian Race 2253

yet the rich russet shade of the latter is not common in the former. ‘In
general there is a bluish shade observed on the common deer, which is so
prevalent as to have given the winter coat the general appellation of the
blue among hunters, who say the deer is in the red or the d/ue, as it may be
in the summer or winter coat. But the difference in the depth of this
colour is so very great, as well as the different shades of colour, as to surprise
any one who will examine thirty or forty together. As the winter advances,
all become appreciably of a lighter colour.

“On this deer the white which universally prevails on the under side
of the head terminates with the throat, or just after it reaches the upper
part of the neck. Thence the under side of the neck has no white, but is
of the prevailing colour of the rest of the neck, until we reach the lower
extremity. There commences a black, or, on some specimens, a brown
stripe, which is always constant, and extends along the brisket to a line even
with the posterior part of the fore-legs. On either side of this black stripe
all is white, which extends down the inner side of the fore-legs to the
knees. All of the belly is also a very pure white, embracing also the inside
of the thighs and hind-legs to the hocks, and up to the tail.” It is added
that the white of the under-parts gradually widens from between the fore-
legs to the abdomen. The extent of white on the lower part of the legs
displays considerable individual variation ; some examples exhibiting a gray
mark four or five inches in length on the inner front side of the fore-leg,
terminating inferiorly in a point, below which it is separated by a tawny
stripe from the white.

The following dimensions of antlers belonging to this and the next race

are recorded by Mr. Rowland Ward :—

Gaerne isartcende Dip De Wire mae path
275 43 14 19 6-6
253 4 9 19 1
255 43 105 19 Li)
254 43 12% 174 11-9
244 44 123 183 6-6
244 4h EEE 194 6-6
24 42 P 18
224 4 P 12
215 & 134 133 4-4
193 4 ° 194 12

254 American Deer

Distribution.—Eastern North America, namely from Maine over much
of the United States east of the Missouri river, typically from Virginia
and Carolina, and probably ranging as far south as Louisiana.

Habits —One of the most striking peculiarities in the habits of this
beautiful deer is the manner in which it runs when first disturbed. This it
does in a kind of rush, with the head carried very low, and the tail elevated
over the back ; it never bounds after the manner of the mule-deer. By
elevating the tail its pure white under surface is displayed ; and this, with
the white of the inner surface of the buttocks, produces a conspicuous
recognition mark. A full account of the habits of this deer in the Adiron-
dack Mountains of New York has been published by Dr. C. H. Merriam,
while Mr. Caton has contributed much important information as to its
habits in captivity, and especially in regard to the periodical changes of coat.
In spite of incessant persecution, a few years ago it was still the most
abundant of the larger mammals in the Adirondacks, where it is found both
high up on the mountains and in the valleys below. And it resorts alike to
the thickest covert and to the open grass lands. During the warmer months of
the year its diet is a varied one, comprising leaves and shoots of trees, various
wild fruits, such as blackberries, grass, and aquatic plants ; beech-mast being
very largely consumed in the autumn. So long as food of this nature is
obtainable the deer thrive and keep in good condition ; but when the snow
is on the ground, where it frequently lies for half the year, they are often
hard pressed for provender. When beech-mast is abundant, and can be
reached by scraping away the snow with their hoofs, they do fairly well ;
but when the supply of this food is small, or the snow unusually deep, they
are driven to browse on the twigs and shoots of various trees and shrubs,
including both the deciduous kinds and conifers. A few mosses and lichens
are also cropped when opportunity offers. Thin as they become towards
the end of a hard winter, no sooner does the succulent marsh grass spring
up in the lowland meadows than they rapidly regain good condition.

Although after the snow has melted in the forests and new vegetation 1s
sprouting, the deer forsake the open meadows for the thickets, a large portion
of their sustenance is obtained during the summer in the neighbourhood of
water. And from June till the latter part of August herds of these deer
visit the streams and lakes of the Adirondacks during the night, and return
at daybreak to the shelter of the forests. To obtain the leaves and seed-pods

Virginian Race BUS

of the water-lilies, which form a very favourite article of food, they con-
stantly wade long distances into the shallow waters, and will sometimes even
venture out of their depth and swim. Here they suffer great persecution
from flies and mosquitoes, which they apparently submit to as an unavoidable
evil. Occasionally in the Adirondacks, and more commonly farther south,
they enter the water when not in search of food, and will then stand with
scarcely more than their eyes and nose exposed ; but whether they do this
in order to rid themselves of insect torments, or merely to enjoy a bath, is
very difficult to decide. Early in September, when the marsh plants in the
Adirondacks begin to wither and die down, the deer forsake the neighbour-
hood of the water-courses for the forests, where a plentiful supply of food
awaits them.

In places much frequented by man the deer in the Adirondacks feed
chiefly or entirely by night, but in more sequestered districts they may be
seen abroad at all hours of the day, and apparently repose at night. Their
favourite resting-places are often under the low boughs of an evergreen
shrub in some dense thicket, or beneath the protecting branches of a fallen
tree. In the deep snows of severe winters the deer frequently collect in
companies, and keep more or less open a limited space, or “ yard,” by
constant trampling, in the same way as the American elk.

In the Adirondacks the new antlers of the bucks make their first
appearance about May, and are generally free from the velvet by the com-
mencement or middle of September. The pairing-season in the same
district commences in the latter part of October and continues till the
beginning of December. At this season the necks of the bucks swell in a
remarkable manner, and their whole demeanour alters. They rush wildly
through the forest in search of the does, fiercely attacking any members of
their own sex with which they may happen to meet, and occasionally even
charging human beings. From their peculiar shape, the antlers are
exceedingly likely to become interlocked during such combats; and not
only have there been several instances of two bucks thus caught, but one
case of three with their antlers interlocked is on record. In fighting, while
the antlers form the chief weapons of offence, the front hoofs are also called
into requisition, and inflict terrific wounds.

‘““T was once sitting quietly in a deer park,” writes Dr. Merriam,
‘“‘when a buck approached, and, making a sudden spring, dealt me such a

256 American Deer

powerful blow on the head with the hoofs of his fore-feet, as to render me
unconscious. No sooner was I thrown upon the ground than the vicious
beast sprang upon me, and would doubtless have killed me outright had it
not been for the intervention of a man who rushed at him with a club and
finally drove him off. Both my father and myself have been knocked flat
upon the ground by being struck in the abdomen by the fore-feet of a very
harmless-looking doe.”

The fawns are born in the Adirondacks between the latter part of April
and the beginning of June ; two being the normal number at a birth and
one the exception. The spots usually persist till September, when both
young and old assume the winter pelage. The birth of an unspotted fawn
at Woburn Abbey has been already mentioned.

With regard to the change of pelage, Mr. Caton writes as follows :—
“The change from the summer to the winter coat is gradual, the new dis-
placing the old by dislodging the hairs promiscuously, till they become so
thin that the new coat is seen through the old. This is not simultaneous
over the whole animal, for the neck and shoulders may be clothed entirely
with the new dress, while the old still prevails on the thighs and rump ;
or the winter coat may have replaced the old on the back, while the belly
still shows only the summer pelage. When the winter has replaced the
summer garb, the hairs are short, fine, and soft ; but they rapidly grow in
length and diameter, and undergo the change of colour peculiar to the
species. At first they lie down smoothly, but presently the diameter
becomes so great that they force each other up to a mere vertical position,
or at right angles to the skin. As the diameters increase, the cavities
within enlarge and become filled with a very light pith ; and the hairs
become brittle and lose their elasticity, so that the integrity of the walls is
destroyed when sharply bent, and they remain in the given position.”

In the Adirondacks Dr. Merriam states that the time of changing the
coat varies somewhat according to the season, and that it is not even
constant in all the deer of any one locality in any particular season. The
difference seems to depend largely upon the degree of severity of the
winters, and the consequent good or bad condition of the animals. After
an unusually hard winter, when the deer are half-starved, the change is
delayed, and the red summer dress is not assumed till late in June or even
the beginning of July; the reassumption of the blue in autumn being

Western Race D7

correspondingly retarded. On the other hand, after a mild winter, or one
in which the supply of beech-mast has been plentiful, the summer change
is early, the gray being soon shed, and the red coat covering the greater
part of the animal by the middle of June, and sometimes even sooner. In
such years the blue dress is likewise donned at an early date, the change
being not unfrequently well advanced by the end of August. Whether the
red summer dress is retained for a longer period in the hotter States, I have
been unable to ascertain.

Although naturally timid in the extreme, Virginian deer, when taken
very young, become exceedingly tame, and thrive well in confinement,
if allowed sufficient space, both in the United States and in England.
They breed readily, but in the United States, at least, display weakness of
reproductive power after a few generations. Mr. Caton observes that if
bucks are raised by hand, they are apt to become dangerous during the
pairing-season, and are consequently unsafe to be allowed to run at large ;
but if reared by their own mothers they never lose their instinctive fear of
man, and consequently never show the least disposition to attack him, or to
take food from his hand. This does not, however, accord with my experi-
ence of the numerous individuals kept at Woburn, where all exhibit
extreme familiarity, and will readily allow themselves to be handled.
Although allowed to run at complete liberty, they do not seem to associate
at all with other deer, and only to a limited extent among themselves.
One pair, for instance, frequent the neighbourhood of the house and stables,
where they will come up unhesitatingly to every visitor to be fed or
fondled ; while other pairs, or solitary individuals, take up their quarters in
some particular portion of the park, from which they seldom wander.

5. WeEsTERN RaceE—MaAzAMA AMERICANA MACRURA

Cervus macrourus, Rafinesque, Amer. Month. Mag. vol. i. p. 436 (1817).

Cervus (Mazama) macrourus, H. Smith, in Grifhth’s Animal Kingdom,
WOlNIV. Ps 134,.V..p. 316 (1827).

Cervus leucurus, Douglas, Zool. “fourn. vol. iv. p. 330 (1829) ; Baird,
Mamm, N. America, p. 649 (1857) ; Caton, Antelope and Deer of America,
p. 100 (1877), as synonym of wrginianus.

2a

258 American Deer

Dorcelaphus macrurus, Gloger, Handbuch Naturgeschichte, p. 140 (1841).

Cartacus leucurus, Lesson, Nouv. Tabl. Hist. Nat. p. 173 (1842) ; Gray,
Cat. Ungulata Brit. Mus. p. 230 (1852), Cat. Ruminants Brit. Mus. p. 83
(1872), Hand-list Ruminants Brit. Mus. p. 155 (1873) ; Brooke, Proc. Zool.
Soc. 1878, p. 919.

Cervus (Mazama) leucurus, Sundevall, K. Svenska Vet. Ak. Handl. for
1844, p. 181 (1846).

Reduncina fleucura, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 357
(1873), lxxviil. part ip. 329 (1879):

Dorcelaphus virginianus macrourus, Allen, Bull. Amer. Mus. vol. vii. p. 257
(1895).

Dorcelaphus macrourus, Mearns, Proc. Soc. Washington, vol. xii. p. 26
(1898).

Characters. — Very similar to the last, but of slightly smaller size,
rather paler colour, and no black on the face and tail. Antlers rather
small and much bowed. General colour of summer pelage on upper-parts
and outer side of limbs reddish brown, in winter bright grayish fawn with
black specklings ; tail fawn-colour, passing into rusty brown, above, pure
white beneath and at the tip’; tarsal gland yellowish brown.

Whatever may be the case with some of the under-mentioned forms,
there can be no doubt as to the present one being merely a local race of
M. americana, with the type form of which it probably intergrades. Mr.
Caton, who includes both /ewcurus (macrurus) and mexicana under the
heading of americana, observes that “in their northern range and in the
mountainous regions of the west, the white portions cover a larger surface
of the body than in other regions, where (in the west ) they have been
ranked by many naturalists as a distinct species under the name of Cariacus
/eucurus. By hunters these have been called the long-tailed, or white-tailed
deer, the latter name having been used by Lewis and Clark, but in truth
their tails are no longer than in those found in other regions. From the
larger extent of white frequently, if not generally, found on them, we might
possibly be justified in assigning them the distinction of a variety, though
this peculiarity is by no means universal, for many individuals cannot be

! In Gray’s Catalogue of Ruminants (1872) the tail of the Virginian race is stated to be white all
round, with the tip brown above, and that of the present race reddish above ; but this is not altogether

correct.

Florida Race 2.59

distinguished from those found in Illinois or Wisconsin. I have one
specimen from North-Western Minnesota with all the legs entirely white to
several inches above the hocks and knees, with occasionally a tawny hair
interspersed among the white. The white on the belly, too, extends
further up the sides than is usually observed. This is exceptional, though
not very uncommon in the north-west, but I have never seen it in their
middle or southern range. I have never found any black on the tails or
faces of the northern variety, while it is very common in more southern
and eastern varieties.”

The name long-tailed deer was doubtless applied to this race in contra-
distinction from other species, and not as distinctive from the eastern race.
This has, however, misled Fitzinger, who speaks of the present race as
differing from the eastern race by the greater length of the tail.

Distribution.—Western United States, including North California,
Oregon principally east of the Cascade Mountains, Washington, Dakota
west of the Missouri, and Nebraska.

c. FroripA RacE—MazAMA AMERICANA OSCEOLA

Cariacus osceola, Bangs, Proc. Soc. Washington, vol. x. p. 26 (1896).

Dorcelaphus osceola, Mearns, ibid. vol. xii. p. 25 (1898).

Characters.—Scarcely more than half the size of the common ™.
americana typica, of the Eastern United States, with the pelage darker at all
seasons, and differing less in summer and winter. There is a difference
in the form of the maxille and nasals, and the cheek-teeth are larger.
Mr. Bangs characterises this form as follows :—‘“‘Size small; general
colour dark ; hair short and fine at all seasons. Upper parts of back, neck,
and head a mixed dark and light brown, each hair banded, dark brown at
the tip, then yellowish brown, then dark brown, and Isabella colour at the
base. The dark brown colour predominates in a narrow median band along
the back, and is most intense on the neck and between the ears. On the
flanks and along the sides the hairs are not banded, but are Isabella colour
at base and cinnamon at tips; sides and under surface of neck cinnamon ;
throat, belly, inside of legs white ; ears sparsely haired ; upper surface dark
brown, many of the hairs tipped with yellow ; inside surface white ; the

260 American Deer

hairs of the upper side of tail are dark-red brown at base and cinnamon at
tips ; under side of the tail white, the hairs very long ; eyelashes jet black.
An old male, in worn midsummer coat, has lost the banding of the hairs
and is a bright russet cinnamon above, which extends to the front of the
eyes. The muzzle is very sparsely haired, and of a grizzled hair-brown
colour, with a black spot behind each nostril. The tail is broadly edged
with black at the base and black above at the tip. An old male from
Blitches Ferry, Citrus County, Florida, in fresh autumnal pelage, is very
dark above, the lower dark band of the hairs extending to their base and

Fic. 70.—Florida Deer. From a photograph by Mr. Rowland Ward.

imparting to the whole upper-parts a rich dark-brown colour, variegated
by the yellow bands of some of the hairs; tail not edged with black, but
like that of the type. A half-grown female has the hairs of the back
unbanded and is clay-colour above, beautifully marked with small irregular
white spots.” The limbs are longer, the antlers longer, the teeth larger,
and the muzzle more elongated than in the next race.
Distribution.—Peninsula of Florida. Mr. Bangs writes that the Florida
deer is of very general distribution over the whole of peninsular Florida,
but in the more thickly settled and accessible parts of the State it has been
much reduced in numbers of late. Its northern range is unknown to me,
and I am therefore unable to state whether or not it overlaps the range of

M. americana typica.

Mexican Race 261

d. SONORAN RacE—MazAMA AMERICANA COUESI

Cervus mexicanus, Baird, N. Amer. Mamm. p. 653 (1857), nec Lichten-
stein, 1827-34.

Dorcelaphus couest, J. A. Allen, Bull. Amer. Mus. vol. vii. p. 200 (1895).

Characters.—A very small, pale-coloured form (smaller than the Texan
race), with relatively large ears, on the margins and tips of which there is
no black ; tail relatively long ; and cheek-teeth small. Antlers small, with
short tines.

Distribution —Sonora, typically from the Santa Cruz Mountains.

e. TEXAN RacE—MazAMA AMERICANA TEXANA

Dorcelaphus texanus, Mearns, Proc. Soc. Washington, vol. xii. p. 23 (1898).

Characters. — Size small; ears relatively small, with black on the
margins and tips; general colour pale; pelage fine and long; limbs
relatively short; and cheek-teeth large. Antlers small and strongly
incurved.

Distribution —Texas and North Mexico.

j) Mexican Race—MazaMaA AMERICANA MEXICANA

Cervus mexicanus, Gmelin, Syst. Nat. vol. i. p. 179 (1788) ; Lichtenstein,
Darstell. Thiere, plate xviii (1827-34) ; Caton, Antelope and Deer of America,
p. 100 (1877), as synonym of wrginianus.

Cervus (Mazama) mexicanus, H. Smith, in Griffith’s Animal Kingdom,
matey p. 715 (1827).

Mazama mexicana, Jardine, Naturalists Library—Mamm. vol. ii. p. 175
(1835) ; Gray, Knowsley Menagerie, plate xlvi (1850).

Cariacus mexicanus, Lesson, Nouv. Tab/. Hist. Nat. p. 173 (1842) ;
Gray, Cat. Ruminants Brit. Mus. p. 156 (1872), Hand-list Ruminants Brit.
Mus. p. 156 (1873) ; Brooke, Proc. Zool. Soc. 1878, p. 919; Alston, Bi/.
Centr. Amer. Mamm. pp. 82, 113 (1879); Ward, Records of Big Game,

p. §8 (1896).

262 American Deer

Reduncina mexicana, Fitzinger, SB. Ak. Wien, vol. xviii. part 1. p. 357
(1873), Ixxvili. part i. p. 328 (1879).

Cariacus virginianus mexicanus, Rhoads, Amer. Naturalist, vol. xxviii.
p- 525 (1894).

Characters.

Size very much smaller than that of the typical race, the
height at the shoulder in Lichtenstein’s example being 2 feet g inches.
Antlers smaller, but of the same general form as in the typical race. In
Lichtenstein’s example the coloration (season not stated) is described as
follows. Upper-parts rusty grayish brown, without any distinct mixture
of red, finely speckled with whitish, the individual hairs being whitish for
the greater part of their length, then ringed with reddish brown, next
with yellowish white, and the tips black ; chest reddish brown ; under-
parts white ; limbs above similar to back, inferiorly without speckling ;
lower jaw and throat whitish, without any dark markings on the former ;
tail yellowish brown speckled with whitish at the base of the upper
surface, then more uniform rusty yellowish brown, and white at the tip
and beneath ; tarsal tuft brownish with a white border. The British
Museum has recently acquired two skins of Mexican females, the one in
summer and the other in winter pelage. In the former the general colour
of the upper-parts is speckled foxy red, but the head and ears dark speckled
gray, becoming tawny behind and below the ears ; no white on the muzzle,
but the chin, lower jaw, throat, and under-parts pure white ; tail bright
foxy red above, white beneath and at the margin. Metatarsal tuft very
small. Possibly the gray head may be due to the assumption of the winter
coat. The specimen in winter pelage is dark brownish gray, speckled with
white, but the upper surface of the tail still bright rufous. The hair is
much longer than in the summer specimen. These were received before
the description of the Texan race, with which I have not had the
opportunity of comparing them.

In one of Lichtenstein’s examples the antlers measure 114 inches in
length, but a pair in the British Museum reach 133 inches along the
curve. Specimens with eight and ten, and in one instance with fifteen
tines, are recorded from Mexico and Texas by Sir Victor Brooke ; some
of these belonging to the preceding race. The smaller specimen from
Guatemala referred to by the same writer belongs to M. americana nemorats,
as do the Costa Rica examples described by Dr. Gray.

Yucatan Race 2.63

The specimens which must be regarded as the actual types are those
described by Lichtenstein, and preserved in the Museum at Berlin, since
Gmelin’s C. mexicanus is a medley. On this point Mr. Rhoads observes
that the Cervus mexicanus of Gmelin is based on the teuthlalmacame of
Hernandez’s History of Mexico (1651). The description of the latter does
not apply to the prong-buck, as asserted by Berlandier,’ and repeated by
. Alston. Hernandez’s figure of the teuthlalmacame, whether intended for
the deer or prong-buck (it partly fits both), cannot affect the description,
which applies to the deer.

Distribution —South Mexico.

g. YucATAN RaceE—MazAMA AMERICANA TOLTECA

(?) Cervus cariacou, Wiegmann, Isis, 1833, p. 965; Pucheran, Arch.
Mus. Paris, vol. vi. p. 366 (1852).

Cervus toltecus, de Saussure, Rev. Mag. Zool. ser. 2, vol. xii. p. 247
(1860).

Coassus toltecus, Gray, Cat. Ruminants Brit. Mus. p. 92 (1872).

(?) Reduncina cartacou, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 357
(1873), Ixxvill. p. 334 (1879).

Cervus yucatanensts, Hays, Ann. Lyc. N. York, vol. x. p. 218 (1874).

Cervus acapulcensis, Caton, Antelope and Deer of America, p. 113 (1877).

Cartacus toltecus, Brooke, Proc. Zool. Soc. 1878, p. 921; Alston, Biol.
Centr. Amer.—Mamm. p. 117 (1879).

Cartacus yucatanensis, True, Proc. U.S. Mus. vol. xi. p. 423 (1888).

Characters.—Size very small, apparently about one-third less than that
of mexicana, but the tail relatively longer than in the latter. Antlers short,
upright, nearly straight, and semi-palmate, with but little forward projection
of the lower prong of the main fork, and the number of tines reduced.
Metatarsal gland and tuft wanting. General colour of upper-parts dark
chestnut-brown ; face blackish ; under-parts white ; tail, which is truncated
at the tip, brown above and white beneath. No seasonal colour-change.

The aborted antlers (figured on page 224 of Mr. Caton’s book) and the
want of the metatarsal tuft might be considered as sufficient evidence to
raise this form—the acapulco deer of Mr. Caton—to. the rank of a distinct

1 See Baird, Mammals of N. America, p. 666 (1857).

264 American Deer

species. But the antlers are essentially only a degraded type of those of the
Mexican race, and too little is known of the functional importance of the
metatarsal gland to render its absence necessarily a specific character.
Moveover, its small size, or absence in the variety memoralis, indicates a
transition from the typical race to the present one.

Distribution.—Yucatan and part of Southern Mexico.

Habits.—Of the habits of this deer in a wild state I can find no record.
In captivity, according to Mr. Caton, it displays great pugnacity, the bucks
not hesitating to attack other deer of three times their size and weight, and
beating them by sheer courage and pluck. They are hardy and breed
readily, although it has not been ascertained for how many generations this
will continue. Although living naturally in the torrid zone, they will
withstand a temperature some degrees below zero, and thus present a
remarkable contrast to the mule-deer and black-tailh Mr. True has
suggested that de Saussure’s fo/fecus is the same as sartorit, in which case
yucatanensis will be the name of the present race.

A. CENTRAL AMERICAN RAcE—MazAMA AMERICANA NEMORALIS

Cervus (Mazama) nemoralis, H. Smith, in Griffith’s Animal Kingdom, vol.
IV. Pu. 137, Xp» 307 oe).

Mazama nemoralis, Jardine, Naturalists Library—Mamm. vol. ii. p. 175
(1835).

Cariacus nemoralis, Lesson, Nouv. Table Hist. Nat. p. 173 (1842) ; Gray,
Cat. Ungulata Brit. Mus. p. 232 (1852) ; Matschie, Mitt. geogr. Ges. Liibeck,
1894, p. 130.

Cervus nemoralis, Pucheran, Arch. Mus. Paris, vol. vi. p. 336 (1852) ;
Brooke, Proc. Zool. Soc. 1878, p. g1g, as synonym of mexicanus.

Reduncina nemoralis, Fitzinger, SB. Ak. Wien, vol. Ixviil. part 1. p. 357
(1873), Ixxvill. part i. p. 338 (1879).

Characters.—Apparently nearly allied to the last, but with the metatarsal
gland usually remaining, although very small, with scarcely any white, and
situated near the middle of the cannon-bone. A female from Guatemala
mentioned by Victor Brooke (as C. mexicanus) measured 27} inches at the
shoulder ; and antlers from the same locality have six points. The follow-
ing description of the coloration is given by Dr. Gray on the evidence of a

Colombian Race 265

male and female from Costa Rica in the British Museum :—‘“‘ Fur brown,
punctulated with yellow ; forehead and crown blackish, with hair diverging
from a central longitudinal line ; legs gray ; tail same colour as the back
above, under side and tip white; upper lip and hinder part of lower lip
black ; spot on sides of nostrils and lower lip white ; a narrow white streak
behind the brown on the nose.” ‘The antlers measured by Sir Victor
_ Brooke have a length of 74 and a span of 84 inches. In two skins in the
British Museum from Costa Rica (referred to in Gray’s Catalogue of Rumu-
nants as C’. mexicanus) the metatarsal gland and tuft, although small, are
distinct ; but in an example from Honduras in the same collection they are
totally wanting. The slight value of this organ for classificatory purposes
is, therefore, apparent.

Distribution —Central America, from Honduras through Costa Rica and
Nicaragua to Panama.

7. COLOMBIAN RAcE—MazAMA AMERICANA GYMNOTIS

Cervus gymnotis, Wiegmann, Iss, 1833, p. 965.

(?) Cervus goudoti, Gervais and Gay, Ann. Sci. Nat. ser. 3, vol. v.
p- 93 (1846) ; Fitzinger, SB. 4k. Wien, vol. \xxix. part i. p. 67 (1879).

Cariacus gymnotis, Brooke, Proc. Zool. Soc. 1878, p. 921; Matschie,
Mitt. geogr. Ges. Liibeck, 1894, p. 130.

Gymnotis wiegmanni, Fitzinger, SB. Ak. Wien, vol. xxviii. part 1.
Pp: 344 (1879).

(?) Cervus columbicus, Fitzinger, op. cit. vol. Ixxix. p. 66 (1879).

Characters.—Height nearly the same as in the last (26 inches at the
shoulder in the type), but the metatarsal gland and tuft wanting, the ears
large, drooping, and almost naked externally, the head very narrow and
pointed, and the general form slender and delicate. General colour of
pelage of upper-parts yellowish brown speckled with gray at all seasons,
the individual hairs being gray at the base, then brown and ochry, with
black tips; a dark streak from the crown of the head along the neck ; a
grayish white ring round the eye; yellowish white and brown spots on
the forehead, a darker spot on the nose, and a patch on the upper lip,
followed by a white streak reaching to the angle of the mouth; chin
white, with a dark streak reaching to the lower lip, and thence extended

2M

266 American Deer

to the angle of the mouth; under-parts pure white; tail pale reddish
brown above, with the tip and lower surface white. ‘The antlers appear
to be very similar to those of the last, and measure just short of 7 inches in
the type.

This deer, of which the type specimen is preserved in the museum at
Berlin, seems to be one of the most distinct of the group, and was regarded
by Sir Victor Brooke as probably entitled to specific rank, which view may
be correct. It was raised to the rank of a genus by Fitzinger on account
of the absence of the metatarsal tuft, but fo/fecus shows that such a distinc-
tion cannot be maintained. If the latter be classed only as a race, it seems
almost necessary to follow the same course in the present instance.

Fitzinger’s C. columbicus, founded on antlers described by Pucheran
from Bogota, Colombia, is considered by its founder to be probably identical
with savannarum, but the locality suggests the present form.

Distribution.—Colombia.

J. SAVANNA Race—MazaMa AMERICANA SAVANNARUM

Cervus savannarum, Cabanis and Schomburgk, Rezsen Brit. Guiana, vol.
il. p. 785 (1848) ; Gray, Cat. Ruminants Brit. Mus. p. 84 (1872), as a
synonym of mexicanus.

Reduncina savannarum, Fitzinger, SB. Ak. Wren, vol. \xviii. part i.
p- 359 (1073), Ixxvul. part 1. p..242 (1870),

Cariacus savannarum, Brooke, Proc. Zool. Soc. 1878, p. 920; Matschie,
Mutt. geogr. Ges. Liibeck, 1894, p. 130.

Characters ——Another small, short-antlered race, of which the type is
preserved in the museum at Berlin. In many respects it is stated to be
intermediate between the Virginian and Colombian races, the general
colour of the pelage being clear grayish brown speckled with white.
From the former it differs by its greatly inferior size, smaller antlers, and
shorter tail; from the latter by the hairy external surface of the ears and
the presence of the metatarsal gland and tuft. The under lip is stated to
be peculiar in having only a single dark spot on each side, totally uncon-
nected with any other markings.

Distribution.—Guiana and probably the whole Orinoco basin.

Wrue’s Deer 267

hk. PERUVIAN RAcE—MazAMA AMERICANA PERUVIANA

Cervus (Coassus) peruvianus, Gray, Ann. Mag. iNet Eliseser. A. vol. Xiit.
Pp: 332 (1874).

Cariacus peruvianus, Brooke, Proc. Zool. Soc. 1878, p. 920 ; Matschie,
Mitt. geogr. Ges. Liibeck, 1894, p. 129.

Characters—Another nearly related small form, without a metatarsal
gland or tuft, the type specimen of which is in the British Museum.
General colour of pelage of upper-parts dark grayish brown, with whitish
speckling, but the upper surface of the tail uniformly dark brown ; the
individual hairs dark isabelline gray at the base, with a whitish band
below the black tip ; under-parts whitish.

Since several skins of this deer preserved in the British Museum are
all of the same colour, it is probable that the dark hue is persistent at all
times of the year. Fitzinger states that the hair on the hinder half of the
back is devoid of speckling, but this is not the case with the specimens in
the British Museum.

Distribution —Peru and Bolivia.

2. TRuE’s DEER—MAZAMA TRUEI.

Cariacus clavatus, True, Proc. U. S. Mus. vol. xi. p. 417 (1888), nec
Cervus clavatus, H. Smith, 1827.

Characters ——A small deer, ranging from Tehuantepec in Southern
Mexico to Costa Rica, has been described under the name of Carzacus
clavatus, with the following leading characteristics. Size relatively small ;
general appearance and colour very similar to M. americana typica ; antlers
in the form of simple spikes directed backwards nearly in the plane of the
face; a small metatarsal gland present; hoofs yellowish at the tip.
General colour of pelage of upper-parts in summer bright chestnut, grayer
on the head than on the back ; a white spot on each side of the nose,
followed by a dusky brown band extending from the nostril to the margin
of the upper lip, and continued by a spot on the border of the lower lip ;
behind the dusky band another of gray, passing into dark gray behind ; a

dusky brown streak down the middle of the face ; a crest of long reflexed

268 American Deer

hairs on the forehead, darker in general tint than the hair of the face.
Outer side of ear mostly gray, but a large white spot at the base ; upper
surface of tail tawny like back ; under surface of lower jaw, throat, under-
parts, upper half of inner surface of limbs, and under surface and tip of tail
white ; rest of limbs nearly like the back. In winter the general colour
speckled brownish gray (‘‘ pepper-and-salt.”)

Mr. True is perfectly correct in regarding this deer as allied to the
Virginian deer, and having no affinity with the brockets. And since it comes
from the same region as M. americana nemoralis, with which it agrees in the
presence of a small metatarsal gland, and apparently in the general colora-
tion, the suspicion naturally arises that it is not separable from that sub-
species. It is known that the Virginian deer has spike-like horns in the
first year, which in rare instances may persist till the second year, and
those of the present form might be an arrested development. But the
skull, of which Mr. True has sent me an adult example, is quite different
from that of all forms of the common American Deer. As c/avatus is
preoccupied by Hamilton Smith’s Cervus clavatus (see p. 249), which is a
synonym of M. americana, | propose the name of Mazama truet.

Distribution —South Mexico to Costa Rica.

3. CRooke’s BLACK-TAILED DEER—-MAZAMA CROOKEI

Dorcelaphus crooki, Mearns, Proc. U. S. Mus. vol. xx. p. 468 (1897).

Characters.—This form, to which specific rank is only provisionally
allowed here, was founded on a female specimen stated to present characters
intermediate between the western race of M. americana and the mule-deer.
The original description is as follows :—In the summer pelage. Colour
reddish fawn, darker from black annulations on the back, lightening to
grayish cinnamon on the sides, and grayish drab on the neck. The legs
are cream-buff, except where new clay-coloured hair is coming in on the
anterior border, the limbs being almost the last part to receive the summer
coating. The colouring of the head is very similar to that of the mule-
deer in corresponding pelage. It has the horseshoe, or arrow mark on the
forehead, and other dark markings of the head to correspond; and the ears
are relatively almost or quite as large, and as scantily [sc] coated with hair.
The bushy hair around the metatarsal gland, which agrees in size and

Mule- Deer 269

location with that of Mazama columbiana, is sooty at base, and white apically.
The tail is coloured much as in M. columbiana, but has a longer terminal
switch ; upper side and extremity of tail all black, lower side white
mesially, and naked towards the base. The pelage of this deer is short
and coarse in comparison with that of the white-tailed or Virginian deer,
or the black-tailed deer of the Columbia River region; and, as would
naturally be expected, is not so red as that of the latter. The skull has
very nearly the same conformation as that of M. co/umbiana, the lachrymal
pit being deeper than in the Virginian deer, but shallower than in the
mule-deer. The same intermediate condition obtains with respect to the
vomer, in the relationships of the nasal and premaxillary bones, in the form
and size of the teeth; and, in short, the whole animal appears to be a
compromise between the characteristics of the white-tailed and mule-deer.
Is it possible that it may be a half-breed between the two?
Distribution.—Dog Mountains, Grant County, New Mexico.

4. THe Mure-DEER—MazaAMA HEMIONUS

Cervus hemionus, Rafinesque, Amer. Month. Mag. vol. i. p. 436 (1817).

Cervus auritus, Desmarest, Mammalogie, vol. 1. p. 443 (1822).

Cervus macrotis, Say, in Long’s Exped. Rocky Mountains, vol. ii. p. 83
(1823) ; Caton, Antelope and Deer of America, p. 93 (1877).

Cervus (Mazama) macrotis, H. Smith, in Griffith’s Animal Kingdom, vol.
iv Deekeo=y. p- 910 (1827).

Mazama macrotis, Jardine, Naturalist’s Library—Mamm. vol. iii. p. 175
(1835).

Dorcelaphus macrotis, Gloger, Handbuch Naturgeschichte, p. 140 (1841).

Cariacus macrotis, Lesson, Nouv. Tab/. Hist. Nat. p. 173 (1842) ; Gray,
Proc. Zool. Soc. 1850, p. 239, Cat. Ungulata Brit. Mus. p. 234 (1852) ;
Brooke, Proc. Zool. Soc. 1878, p. 921; Alston, Biol. Centr. Amer.—Mamm.
p- 114 (1879) ; Ward, Records of Big Game, p. 60 (1896).

Eucervus macrotis, Gray, Ann. Mag. Nat. Hist. ser. 3, vol. xviii. p. 339
(1866), Cat. Ruminants Brit. Mus. p. 86 (1872), Hand-list Ruminants Brit.
Wasp. 157 (1873).

Orelaphus macrotis, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 356

(574), xxvin. part i. p. 303 (1879).

270 American Deer

Dorcelaphus hemionus, Allen, Bull. Amer. Mus. vol. vii. p. 257 (1895).

Characters.—Build heavier and coarser than in the common American
deer. Size somewhat larger than the Virginian race of the latter, the
height at the shoulder reaching to 3 feet 3 or 4 inches. Antlers with a
very short sub-basal snag, beyond which the beam is directed outwards for
a short distance and then curves upwards to form a regularly dichotomous

Fic. 71.—Front view of Head of Mule-Deer. From a specimen in the possession of Mr. E. S. Cameron.

fork, of which both prongs are normally nearly equal, and again divide, the
normal number of points thus being five on each side. Lars very large and
thickly haired ; tail moderately long, terminating in a brush-like tuft of hair,
naked on the under surface at the base; muzzle rather short ; face-gland
rather large. Metatarsal gland and tuft long and straight, occupying the
whole of one side of the upper half of the outer surface of the cannon-bone,

its hair nearly the same cinnamon tint as that of the leg, as is likewise that

THE Deer or Atu Lanps. Puate XX].

MULE-DEER IN WINTER PELAGE.

Published by Rowland Ward Lid.

Mule- Deer pias

of the tarsal gland. General colour of pelage of upper-parts some shade of
reddish or yellowish tawny in summer, in winter dark brownish or rufous
gray, more or less speckled with whitish, darkest on the withers ; forehead
usually with a dark brown patch, extending nearly to the eyes, and
commonly known as the “horseshoe,” a brown patch on each side of the
nose, rest of face and throat white; inner side of buttocks and legs,
abdomen, and most of inner surface of ears white or whitish, as is also the
tail, with the exception of its terminal tuft, which is black ; rest of under-
parts blackish brown. The front border of the ear is black. On the first
assumption of the winter coat the colour, according to Mr. Caton, becomes
for a short time almost black.

The large hairy ears, the elongated metatarsal gland, the tufted tail,
with its lower surface naked at the base, and the shape of the antlers, form
the leading characteristics of this well-marked species.

Distribution.-—The greater part of North America westward of the
Missouri River, extending, according to Mr. Caton, over about thirty
degrees of latitude, from Cape St. Lucas, at the southern extremity of
California in the south, into British Columbia in the north. The range
thus includes suitable localities in parts of Dakota, Nebraska, Kansas, Texas,
Colorado, Wyoming, Montana, Idaho, Nevada, California, Oregon,
Washington, British Columbia, and probably some other States.

Mr. Caton writes that “the original range of this deer has not been
very much restricted since it was first discovered, although it has been
driven back from the Missouri River, and has deserted other limited local-
ities, where the miners or settlers have driven it away. . . . West of the
Rocky Mountains it is met with almost everywhere, though much more
abundantly in some places than in others. In the coast range of Northern
California it is almost entirely replaced by the Columbian black-tailed deer,
while in the coast range of Southern California scarcely any other deer 1s
met with. . . . In all of Oregon, in Washington Territory, and in British
Columbia, this deer is met with, though much less abundant than the true
black-tailed deer, or even the Virginian deer.”

Mr. J. Turner-Turner! has published a map showing the present dis-
tribution of the mule-deer in British Columbia, from which it appears that
the range is now limited to a narrow strip of country running to the north-

1 Three Years’ Hunting and Trapping (1888).

ye oh American Deer

west of Fort George, a second running southwards from that point for some
distance down the Fraser River and then to the eastward of the same, and a
third patch lying some way to the west of the lower part of that river.
Antlers.
with regard to the cranial appendages :—‘ The first antlers are usually forks,

Mr. E. S. Cameron has supplied the following information

but I have obtained an eighteen months’ old fawn with perfect spikes, and
twice shot deer bearing for their first antlers a spike on one side and a fork
on the other. In the second antlers the beam lengthens, but is still simply
forked. In the third antlers one of the prongs, either anterior or posterior,
is again forked, while the other remains simple. In the fourth antlers both
prongs are forked. ‘The sub-basal snag is not, apparently, developed until
the assumption of the fifth antlers. The antlers are clean by the end of
October and are shed in March.”

The following dimensions of antlers belonging to this species and to the

black-tailed Columbian deer are recorded by Mr. Ward :—

Length along Basal : : Widest Number :
Outer Curve. Circumference. Pipe ey: inside. of Points. BEET E
30 53 41 v7 Colorado
283 43 134 17% 5-5 Wyoming
284 P e 244 ? Colorado
273 6 15i 19+ 9-6 Brit. Columbia
27 54 195 215 6-5 °
26% 5 ? P 35 Dakota
3 3 5 1
263 43 192 204 5-5 Wyoming
264 52 P 174 12
264 44 154 22% 6-6 P
254 54 e 214 8
254 43 P 28 Montana
1 3 1
254 43 18 244 5-5 Colorado

Habits —For the following information regarding the habits of the
mule-deer, I am indebted to Mr. E. S. Cameron, who has had unusual
opportunities of observing them in Montana and Dakota. These deer
possess great speed, and can maintain it for a long time; I have seen them
run right away from greyhounds in a broken country. Their ordinary gait
is a succession of stiff-legged bounds, which they exchange for an even
gallop if wounded, or if pressed either by a greyhound or by a swift horse.

Mule- Deer 273

They also trot well. A mule-deer with a broken hind-leg is capable of
wonderful speed downhill, and should a fore-leg be broken, a fast horse
must gallop to catch it on level ground. In horizontal leaping these deer
surpass all animals with which I am acquainted, and for ability to traverse
a rough country there is little to choose between them and their neighbours
the mountain-sheep. On the other hand, when frequenting the fenced

Fic, 72.—Oblique view of Head of Mule-Deer. (Rowland Ward, Records of Big Game.)

pastures of deserted ranches, for which they have a great inclination, they
invariably crawl under, or squeeze through the wires instead of jumping
over them. ‘They are very similar to range-horses in their habits, and the
herds, small or large, will frequent a tract of about ten miles in diameter,
watering at the same place every day if left undisturbed. Like mountain-
sheep, they love to lie sheltered from the wind, on the narrow ledges of
the high buttes, commanding the view all round, except to windward, from
2N

274. American Deer

which side their noses sufficiently protect them. During the summer they
associate in small bands ; a doe with several of her fawns of different years
in one group, and two or three old bucks with a young buck, which acts
as sentinel, in another, while the medium-sized bucks are often quite alone.
At the end of October, or as soon as their antlers are clean, the bucks begin
to “round up” the does; and make no objection to the presence of two-
year, and even three-year-old, male deer in the harem, for I have killed a
master-buck and a three-year-old out of the same party. Although I have
seen a buck with does as early as 18th of October, from the first to the 15th
of November is usual for the commencement of the pairing-season, when
the old bucks grunt like a fallow deer; and I have seen a buck followed
by several does walk to the edge of a “bad-land” point, and challenge
repeatedly across the prairie in the still air.

The bucks do not appear to be able to keep many does, sixteen, to the
best of my recollection, being the most I have seen with one buck, while
eleven or twelve is a more common number ; nor do they appear to suffer
much from the effect of the rut, as immediately after they are more wary
and run greater distances than before. The duration of the pairing-season
is about six weeks. They now collect into small parties, from two to a
dozen, but the idea that they retire singly into solitude to shed their antlers
is, in my opinion, a fallacy.

Although the bucks are pugnacious among themselves during the
pairing-season, I have never known of an injury being inflicted by an
antler-thrust, but have twice come across cases of blindness from cataract.
The old bucks will stand at bay when severely wounded, but never in my
experience act upon the aggressive towards a man on foot, so long as he
keeps a few yards off, although a buck will wheel and charge a horseman
who is pressing it too closely. The does, which drop their spotted fawns
in May, and generally give birth to twins, are always watchful; and
when a party of deer are resting, the long ears of the recumbent does move
backwards and forwards with irregular jerks, one of them constituting her-
self a sentinel, and rising at intervals to listen on all sides before again lying
down. Mule-deer feed chiefly on grass, and keep their condition in spite
of the severe winters, as they can paw away the snow like horses,
supplementing their diet with cedar and buffalo berries, which are very

plentiful. Should the depth of snow be excessive, as a last resource they

Mule- Deer D7

eat the tops of the sage-brush. They only venture on to the prairie at
night to enjoy the superior grass there, but in a wooded or broken country
they may be seen feeding at any hour of the day, more especially if the
night has been stormy. Although their true home is in the “‘ bad-lands,”
they seek shelter from snowstorms in woods, leaving them again as soon as
the sky clears, and I have found the big pine- and cedar-thickets a certain
-haunt in wintry weather. ‘These they will not leave if they discover that
they are being followed, but circle round and round, returning to the
place from which they started and traversing impenetrable brakes through
which a horse cannot be led. At such times they chiefly trot, and the
noise of their feet can be heard in the frozen snow like the ticking of a
clock. I never heard of an unwounded mule-deer being killed by wolves,
which, in my opinion could not catch it. A wounded deer, however, is at
once devoured by them. On the other hand, I know several instances
of these deer being killed by pumas.

When I first went to Montana, in 1889, mule-deer were so exceedingly
plentiful as to excite surprise, but now, 1897, their numbers have been
much reduced by the professional hunters, who shoot them down in
hundreds, and evade the laws by shipping the hind-quarters out of the
State under fictitious names, calling them poultry, mutton, etc., and leaving
the fore-quarters for the wolves. This mercenary chase of mule-deer is
only pursued during the pairing-season, because the meat would not keep
before that period.

Several American-bred specimens of mule-deer have been from time to
time exhibited in the London Zoological Gardens, and from these a large
number of fawns have been produced in the menagerie. Some time ago
there was a pair at Woburn Abbey, but they survived only for a short
period.

a. TypicAL , RAcE—MAZAMA HEMIONUS TYPICA

Cervus macrotis montanus, Caton, Antelope and Deer of America, 2nd ed.
p. 94 (1881).

Characters.—General coloration that described above, of a full dark
tint ; tail yellowish white, with a jet-black terminal tuft.

Distribution —The greater part of the area of the species, exclusive of
that occupied by the other sub-species.

American Deer

hw
NI
Nn

45. CALIFORNIAN RacE—MaAzaMA HEMIONUS CALIFORNICA

Cariacus macrotis californicus, Caton, Amer. Naturalist, vol. x. p. 464
(1376); xix. poOsEL (P88 5).

Characters.—Kars still larger than in the typical race, and the colour of
the pelage, although variable, not unfrequently brighter ; a brown or
tawny line on the upper surface of the tail connecting the dark of the
upper-parts with the black tail-tip. After remarking that the tail is
slightly longer than in the typical race, Mr. Caton has the following
observations on this form :—‘ Its great distinction is in the colour of the
tail, which alone is sufficient to declare it a very distinct variety, where
other indicia, which are plainly declared, wanting. It has the naked
portion on the under side the same as in the eastern variety, but instead of
all being white above the black tuft at the end, a stripe of the colour of
the back above the tail, with which it unites, runs down the upper side of
the tail and unites with the terminal black tuft. On some specimens this
brown stripe grows darker towards the lower end, and on some the tawny
brown shade of the stripe invades the black tuft.”

Distribution.—California south of San Francisco, but the extreme

southern limit not determined.

c. La Paz Race—MazAMA HEMIONUS PENINSULZ

Characters—Much smaller than the last, and still more brightly
coloured, with the antlers in the form of simple spikes and a basal snag.
The winter pelage on the upper-parts is dark speckled iron gray, with an
irregular black band along the middle of the back, expanding towards the
hind - quarters, and continued on to the upper surface of the tail, where
it may be connected by a narrow line with the black extremity, or may be
separated by a broad ring of pale straw-coloured hair. The legs are bright
chestnut, and there is a patch of the same colour on the flanks, separating
the dark speckled gray of the back from the uniform blackish brown of the
under-parts.

This race is named on several skins of females and sub-adult males
recently acquired by the British Museum. Similar specimens from Cape

Mule- Deer 27a

St. Lucas are noticed by Mr. Caton,' who refers especially to the simple
form of the antlers, which are of a degraded type. Several of the other
mammals from the same district are distinguished from their representatives
to the northwards by their brighter coloration.

Distribution —The extreme south of the Californian Peninsula.

d. WESTERN DeEsERT RACE—MAZAMA HEMIONUS EREMICA

Dorcelaphus hemionus eremicus, Mearns, Proc. U. S. Mus. vol. xx. p. 470
(1897).

Characters—The following is the original description of this sub-
species :—The mule-deer of the Western Desert Tract, like all mammals
of that region, is remarkable for the extreme pallor of its coloration. An
adult male, taken in December 1895, in the Sierra Seri, near the Gulf of
California, in the most arid portion of Sonora, Mexico, is in full winter
pelage. The coat is short and glossy. Coloration above very pale drab-
gray, with a dark vertebral area, which begins as a narrow median stripe
on the upper side of the neck, broadens and becomes fainter on the back,
forms a blackish spot at the root of the tail, down which it descends for a
short distance. The buttocks, inguinal, and abdominal regions, and the
middle of the tail all around are white. The axille and hollows of the
thighs are entirely naked. The edges of the buttocks, posterior surface of
limbs, and the feet are washed with pale, muddy cinnamon. The chest is
light sooty drab. Tail with a heavy brush or short switch of black hair at
the end, the middle portion being white all around, the dusky colour
running down a short distance on the upper side from the blackish spot at
its base. While the general effect is to produce a pale drab-gray colour-
ing of the upper surface, there is the usual pepper-and-salt commingling
of colours, produced by light and dark annulation of the hairs, those in the
vertebral area being pointed with brownish black. It appears to be a
larger animal than the mule-deer of the Eastern Desert Tract, and, unless
the specimens are abnormal, its antlers are heavier and more divergent,
being remarkable for the great length of the beam before forking. Ina
youngish specimen from the Sonoyta Valley they are doubly dichotomous
throughout, having four points, besides a basal snag, on each. Another

1 Amer. Naturalist, vol. x. p. 468 (1876).

27 American Deer

pair belonged to a fully adult animal, killed at Black Butte, on the Colorado
Desert near the mud-geysers, Lower California. This was one of six
killed during the winter of 1893-94. The beam is unusually stout ; the
antlers being doubly dichotomous.

Distribution —Western Desert Tract of the Mexican border of the
United States.

5. Tue Biack-TaILep DEER—MazaMa COLUMBIANA

Cervus macrotis, var. columbtanus, Richardson, Fauna Bor. Amer. p. 257
(1820).

Cervus lewisit, Peale, Mamm. U. S. Explor. Exped. p. 39 (1848).

Cervus richardson, Audubon and Bachman, Quadrupeds of N. America,
VoL ‘i. pe 2a1 (185 3):

Cariacus punctulatus, Gray, Proc. Zool. Soc. 1850, p. 239, Cat. Ungulata
Brit. Mus. p. 232 (1852).

Cervus columbianus, Baird, N. Amer. Mammals, p. 659 (1857) ; Caton,
Antelope and Deer of America, p. 96 (1877).

Eucervus columbianus, Gray, Ann. Mag. Nat. Hist. ser. 3, vol. xviii.
p. 388 (1866), Cat. Ruminants Brit. Mus. p. 86 (1872), Hand-list Ruminants
Brat. Mas. pi 057. (a 872)

Otelaphus richardsoni, Fitzinger, SB. Ak. Wien, vol. \xviii. part 1. p. 357
(1873), Ixxvill. part 1. p. 308 (1879).

Reduncina punctulata, Fitzinger, op. cit. Ixvu. part 1. p. 357 (1873).

Cariacus columbianus, Brooke, Proc. Zool. Soc. 1878, p. 921.

Otelaphus punctulatus, Fitzinger, SB. Ak. Wien, vol. \xxviii. part 1.
p. 307 (1879).

Cariacus macrotis columbianus, Ward, Records of Big Game, p. 60 (1896).

Dorcelaphus columbianus, Mearns, Proc. U.S. Mus. vol. xx. p. 468 (1897).

Characters.—Smaller than the mule-deer, with relatively shorter ears
and finer hair, especially distinguished by the shorter metatarsal gland and
tuft, which occupy a considerable portion of the upper half of the seg-
ment of the cannon-bone, and also by the greater portion of the tail being
black, with only the basal third of the lower surface white. General
colour of pelage of upper-parts in winter speckled tawny brownish gray, the
individual hairs being dark brown for the greater part of their length, but

Black-Tailed Deer 279

near the extremity ringed with yellowish brown, and terminating in a black
tip ; hinder portion of under-parts and region of base of tail, as well as
upper throat and chin, white; face gray, darker on the forehead ; legs
dark cinnamon, without any admixture of white hairs ; tarsal and metatarsal
tufts only a shade lighter than the adjacent portions of the leg ; no naked
portion of gland visible in the elongated metatarsal tuft. In summer the
- general colour of the upper-parts reddish yellow. Face-gland rather small.
Antlers of the general type of those of the mule-deer, but relatively smaller.

The distinctive features of this species are the entirely black tail-tip, and
the medium length of the metatarsal gland and tuft, which are situated
entirely above the middle of the cannon-bone. Mr. Caton states that the
gland is intermediate between that of the mule-deer and that of the Virgin-
ian, and figures a specimen in which the naked portion is about an inch and
a half in length. But in a pair of this species in captivity at Woburn
Abbey in the summer of 1897, where they survived only a very short time,
the gland and its tuft were longer, and much more like those of the mule-deer.

Distribution.—W estern North America, from British Columbia, through
Washington and Oregon west of the Cascade Mountains, to California ; the
typical locality being the mouth of the Columbia River, in Washington,
where it was discovered by Lewis and Clarke in 1805. It also occurs in
the coast districts of Vancouver Island. Mr. Caton observes that “the
most extraordinary fact in connection with this deer is the extremely
narrow limits of its range, which is within a narrow belt along the Pacific
coast of America, in the temperate zone. In many parts of this district it
is the most abundant deer to be met with. Why it has never wandered
beyond these bounds, it is hard to say. It has never even reached the base
of the Rocky Mountains, except possibly in the extreme northern part of its
range. The mountain barriers could not restrain it ; for it ranges high up
on the Sierra Nevada, and is found on the eastern slope of that range. If
the deserts at the south would deter it from an eastern migration, the
valleys of the streams heading in the Rocky Mountains, and emptying into
the Columbia River, invited it to follow their banks, and would have led
it to the summit of the range, and to practicable passes.” A map of the
distribution of this species in British Columbia is given in Mr. J.
Turner-Turner’s Three Years’ Hunting and Trapping in America and the
Great North-West.

280 American Deer

Hadits.—In its general mode of life the black-tail is in some respect un-
like the mule-deer, although it resembles the latter in its bounding gait
when frightened. Such a fatiguing pace can, however, be maintained only
for a comparatively short distance, and the deer consequently soon become
blown when they start off in this manner. When starting without being
frightened, they run in a more ordinary way, and are then able to hold out
for a much longer time, as is also the case with the mule-deer. Unlike the
latter, the present species is a forest-loving animal, frequenting the dense
woods of conifers bordering the Pacific coast, whose deep shade affords ample
concealment. In the less disturbed districts these deer even venture down
to the sea-coast, where they feed upon the seaweed cast on the beach ; and
in their journeys to and fro not a few fall victims to prowling Indians.
Like the common deer, this species takes readily to the water, and may
often be seen crossing a river or lake. Although spending a large portion of
the year in the neighbourhood of swamps, in the pairing-season, which
takes place in October, the old stags often resort to higher ground. The
fawns are usually bornin May, their number being generally two, although
triplets have been recorded. They are more fully spotted than those of the
mule-deer, the spots themselves being more sharply defined, and arranged
in more definite longitudinal lines. In these respects the fawns are more
like those of the Virginian deer.

Although some of the adults kept by him in confinement throve fairly
well, Mr. Caton, in the first edition of his book, states that he was never
able to rear a fawn. He thinks that under more favourable circumstances
the fawns might survive for a year or two, but he doubts the probability
of their propagating if brought direct from their native haunts to an unsuit-
able climate. No examples of this species have hitherto been exhibited in
the London Zoological Gardens.

IncERTA SEDIS

1. Mazama spinosa

Cervus spinosus, Gervais and Gay, Aun. Sci. Nat. ser. 3, Pp. 93 (1846) ;
Fitzinger, SB. Ak. Wien, vol. Ixxix. part 1. p. 68 (1879) 5 Gray, Cat. Ruminants
Brit. Mus. p. 84 (1872), as synonym of mexicanus.

Cariacus (?) spinosus, Gray, Cat. Ungulata Brit. Mus. p. 236 (1852).

Marsh - Deer 281

Founded on a single immature antler from Cayenne, now in the Paris Museum
and very probably referable to M. virginiana savannarum.

2. Mazama similis

Cervus affinis, Pucheran, C. R. Ac. Paris, vol. xxix. p. 777 (1849), wec Hodgson,
1841.

Cervus similis, Pucheran, Arch. Mus. Paris, vol. vi. p. 357 (1852).

Reduncina similis, Fitzinger, SB. Ak. Wien, vol. |xviii. part 1. p. 357 (1873)
Ixxvili. part i. p. 321, lxxix. part 1. p. 62 (1879).

Cariacus similis, Brooke, Proc. Zool. Soc. 1878, p. 920.

Founded on a male specimen in the Paris Museum, of which the exact locality is
unknown, but which probably came from the Southern United States. Its general
coloration is very similar to the typical race of the common American deer in summer
pelage, but the upper half of the tarsal tuft is rusty red, and only the lower half
white.

3. Mazama leptocephala

Cariacus leptocephalus, Gray, Cat. Ruminants Brit. Mus. p. 85 (1872), Hand-list
Ruminants Brit. Mus. p. 157, plate xxxvii (1873).

Founded on an immature male skull in the British Museum (No. 52. 12. 26.
160), reputed to have come from South America. It is evidently of the mule-deer
type, and is probably of North American origin.

4. Mazama pusilla

Eucervus pusilla, Gray, Hand-list Ruminants Brit. Mus. p. 157 (1873).

Founded on an immature male skull from North America in the British Museum
(No. 68. 2. 13. 2), probably belonging to the mule-deer group. The antlers are
unbranched. Description apparently insufficient.

5. Mazama sp.

Cervus brachyceros, Philippi, 4u. Mus. Chile, 1895 ;' nec Gervais and Ameghino,
1880.

Cariacus sp., Nehring, SB. Ges. Nat. Berlin, 1895, p. 12.

Founded on a Chilian specimen referred by its describer to Xenelaphus, but
assigned by Dr. Nehring to the present sub-genus, without determination of the

species.

1 T have been unable to see copy of this volume, and therefore cannot give the page.

270

282 American Deer

i. THE Guazu Group—Sus-GENnus BLAsTocERos

Blastocerus, Sundevall, K. Vet. dk. Hand/. for 1844, p. 182 (1846), as a
sub-genus ; Gray, Proc. Zool. Soc. 1850, p. 237, Cat. Ungulata Brit. Mus.
p. 223 (1852), Cat. Ruminants Brit. Mus. p. 87 (1872) ; Brooke, Proc. Zool.
Soc. 1878, p. 922; Riutimeyer, 4d/. schwerz. pal. Ges. vol. viii. p. 49
(1881).

Blastoceros, Fitzinger, SB. 4k. Wien, vol. \xviii. part i. p. 358 (1873).

Paraceros, Ameghino, Mamm. Foss. Répub. Argent. p. 605 (1889) ;
see Lydekker, dn. Mus. La Plata—Pal. Argent. vol. i1. p. 80 (1893).

Antifer, Ameghino, op. cit. p. 610 (1889) ; see Lydekker, of. cit. p. 81
(1893).

Epzeuryceros, Ameghino, of. cit. p. 613 (1889) ; see Lydekker, of. cit.
p- 81 (1893).

Oxotoceros, Ameghino, Rev. Argent. Hist. Nat. vol. 1. p. 243 (1891), to
replace B/astocerus, Sundevall, on account of being preoccupied by Béastocera,
Gerstein, 1856, which is not the case.

Characters. — Antlers large and complex, without a sub-basal snag,
forking in a regularly dichotomous manner, with the upper, or posterior
prong more or less exceeding the lower, or anterior one in size. Metatarsal
gland and tuft absent. Tail short. Face moderately long ; face-gland well
developed, and the gland-pit in the skull rather large and deep. Upper
canines generally present in adult males. Size large or rather small.
Fawns uniformly coloured or spotted.

The absence of the metatarsal gland and of a sub-basal snag to the
regularly dichotomous antlers are the most easily recognised features of the
existing members of this small group. Extinct species occur in the
Plistocene deposits of the Argentine pampas, and apparently also in the
Pliocene beds of Monte Hermoso, in the same country. The occurrence
of such remains in the last-mentioned deposits is of importance as indicat-
ing that the brockets are not the ancestral type of the group ; the Monte
Hermoso beds having been deposited soon after the union of North with
South America, at which time deer of the present, or a nearly allied, type
were already in existence.

Distribution.—The Neotropical region.

Marsh- Deer 28 3

1. THE MarsH-DEER—-MAZAMA DICHOTOMA

Cervus dichotomus, Mliger, Abh. Ak. Berlin, 1811, pp. 108 and 117.

Cervus paludosus, Desmarest, Mammalogie, vol. 1. p. 443 (1822) ; Bur-
meister, Descript. Phys. Répub. Argent. vol. iil. p. 480 (1879) ; Goeldi,
. Mammiferos do Brasil, p. 106 (1893).

Cervus palustris, Desmoulins, Dict. Class. Hist. Nat. vol. il. p. 379
(1823).

Cervus (Mazama) paludosus, H. Smith, in Grifhth’s Animal Kingdom,
wale. p. 216 (1827).

Mazama paludosa, Jardine, Naturalists Library—Mamm. vol. ii. p. 173
(1835).

Dorcelaphus paludosus, Gloger, Handbuch Naturgeschichte, p. 140 (1841).

Cariacus paludosus, Lesson, Nouv. Tabl. Hist. Nat. p. 173 (1842) ;
Brooke, Proc. Zool. Soc. 1878, p. 922; Lydekker, dn. Mus. La Plata—
Pal. Argent. vol. ii. p. 80 (1893); Ward, Records of Big Game, p. 57
(1896).

(?) Mazama furcata, Gray, List Mamm. Brit. Mus. p. 176 (1843).

Cervus (Elaphus Blastocerus) paludosus, da Schreber’s Sdugethiere,
vol. iv. p. 367 (1844).

Cervus (Blastocerus) paludosus, Sundevall, K. Vet. Ak. Handl. for 1844,
p- 182 (1846).

Blastocerus paludosus, Gray, Proc. Zool. Soc. 1850, p. 237, Cat. Ungulata
Brit. Mus. p. 224 (1852), Cat. Ruminants Brit. Mus. p. 87 (1872), Hand-hst
Ruminants Brit. Mus. p. 157 (1873); Miiller, Zoo/. Garten, vol. xxxvii.
p. 49 (1896).

Blastoceros paludosus, Fitzinger, SB. Ak. Wien, vol. Ixviii. part 1. p. 359
(re73), xxviull. parti -p. 350 (18760).

Blastoceros paludosus fossilis, Ameghino, Mamm. Foss. Répub. Argent. p. 608
(1891); see Lydekker, 4n. Mus. La Plata—Pal. Argent. vol. ii. p. 80
(1893).

(?) Paraceros ensenadensis, Ameghino, /oc. cit. ; see Lydekker, doc. cit.

Cervus (Blastoceros) paludosus, Uhering, Mammiferos de S. Paul, p. 14

(1894).
Cariacus palustris, Lydekker, Royal Nat. Hist. vol: ii: p- 387 (1894).

28.4 American Deer

Characters—Largest of the South American deer, the size being appar-
ently fully equal to that of a red deer, but the build more slender. Antlers
large and rugose, with both prongs of the main fork dividing more than
once, and the upper prong usually somewhat larger than the lower one.
Muzzle bluntly pointed ; ears large, and filled internally with woolly white
hair ; tail bushy. Hair long and coarse, without radiating whorls on the
back and neck. General colour of upper-parts bright rufous chestnut in

Fic. 73.—Side view of Antlers of Marsh-Deer. From a specimen in the British Museum.

summer, in winter brownish red, becoming lighter on the flanks, neck, and
chest ; legs black from the knee and hock downwards, and the tarsal tuft
also black ; abdomen, inside of thighs, throat, chin, and inside and base of
back of ears white or yellowish white ; a whitish line above, or a ring
round the eyes, most marked in the hinds ; a black band on the muzzle and
upper lip, joining a median dark streak on the nose, and black markings on
the under lip; tail yellowish rusty red above and black beneath. In
immature hinds the limbs are less black, showing fawn on the sides

Marsh- Deer 2.8 5

below the hock and knee, in advance of which the extremities are
white, and the black streak on the nose is wanting. The young are
not spotted.

This deer has never been exhibited in the London Zoological Gardens,
although a specimen was living in the menagerie at Berlin in 1896. I
have never seen a fully adult example, and the species is best known to me

Fic. 74.—Young Marsh-Deer. From a photograph by the Duchess of Bedford.

by an immature mounted specimen at Woburn Abbey, and the equally
young examples forming the subject of Fig. 74. It is somewhat curious to
note that the coloration of this species is almost identical with that of the
so-called maned wolf (Canis jubatus) of the same districts.

Although ten is the normal number of their points, the antlers of this
species are liable to “sport” in an extraordinary degree ; specimens as
complex as the one represented in Fig. 75 being by no means uncommon.
The following dimensions of antlers are taken from Mr. Rowland Ward’s

Records of Big Game :—

286 American Deer

On Cee) renee Tip to Tip. Widest inside, pane
242 5 16 18 eG
233 $ 26 25 seeds 1
23% 5 15 165 4-5
234 6 ? 194 12
228 61 20 203 bige.
223 53 ou P 28
21d 5 123 16 5-5

Distribution.—In suitable localities throughout Brazil, and perhaps part
of Guiana, through Paraguay, Entre Rios and Uruguay to the Chaco, or

Fic. 75.—Malformed Antlers of Marsh-Deer. (Rowland Ward, Records of Big Game.)

wooded interior districts of Argentina. According to Admiral Kennedy,
the species is, however, now very rare in Uruguay and Entre Rios. It is
abundant in the neighbourhood of Matto-Grosso, Brazil. Its fossilised
remains occur in the superficial deposits of the Argentine pampas; the
antlers from these formations described under the name of Paraceros appar-
ently belonging either to the present or a closely allied form.

With regard to the adoption of the name dichotoma, instead of the
usually accepted pa/udosa, it may be mentioned that although Iliger gives
no description of the animal he designates by the former name, yet as he
refers to the gouzoupoucou of Azara, which is the present species, his

name is entitled to stand.

Pampas Deer Dey

Habits —According to Dr. Goeldi, the veado galheiro inhabits the
thick jungles on the borders of the great Brazilian rivers, where it goes
about in small parties of from three to five individuals. During the day it
lies hidden in the rank vegetation, issuing forth to feed about sunset, and
remaining abroad till after sunrise ; its food consisting of reeds and various
other aquatic plants. Its sense of smell and hearing being very acute, and
its flight rapid, this deer is difficult to stalk. The hind produces a single
fawn annually, which after four or five days is able to run by the side of
its dam. If taken young it can be easily tamed, but its constitution is very
delicate. Although its flesh is only eaten where other food cannot be
obtained, its skin is largely used for leather by the natives.

2. THE Pampas DEER—-MAZAMA BEZOARTICA

Cervus bezoarticus, Linn. Syst. Nat. ed. 12, vol. i. p. 67 (1766).

Cervus campestris, F. Cuvier, Dict, Scz. Nat. vol. vii. p. 484 (1817) ;
Burmeister, Descript. Phys. Répub. Argent. vol. iii. p. 463 (1879) ; Goeldi,
Mammiferos do Brasil, p. 107 (1893).

Cervus leucogaster, Schreber, Sdugethiere, vol. v. p. 1127 (1817).

Cervus (Mazama) campestris, H. Smith, in Grithth’s Animal Kingdom,
VOMMIVe Dl 3O,.V. Ps 317 (1827).

Cervus azare, Wiegmann, Isis, 1833, p. 954."

Mazama campestris, Jardine, Naturalists Library—Mamm. vol. ii. p. 174.
(1835) ; Gray, Last Mamm. Brit. Mus. p. 176 (1843).

Dorcelaphus campestris, Gloger, Handbuch Naturgeschichte, p. 140
(1841).

Cariacus campestris, Lesson, Nouv. Table Hist. Nat. p. 173 (1842) ;
EOOl<cs lo 70c8 200). Soc. 1878, Pp. 9235 Aplin, zbid. 1894, p. 313; Ward,
Records of Big Game, p. 59 (1896).

Cervus (Elaphus Blastocerus) campestris, Wagner, Schreber’s Sdugethiere,
vol. iv. p. 369 (1844).

Cervus (Elaphus Blastocerus) comosus, Wagner, op. cit. p. 368 (1844).

Furcifer campestris, Gray, Knowsley Menagerie, p. 68 (1850).

Blastocerus campestris, Gray, Proc. Zool. Soc. 1850, p. 237, Cat. Ungulata

1! The original description is stated to be in a work entitled Abbi/dungen merkwurd. Sdaugethiere
(p. 69), very few copies of which appear to have been published.

era

288 American Deer

Brit. Mus. p. 224 (1852), Cat. Ruminants Brit. Mus. p. 87 (1872), Hand-lst
Ruminants Brit. Mus. p. 158 (1873).

Blastoceros campestris, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 359
(1873), Ixxvill. part i. p. 364 (1879).

Blastoceros comosus, Fitzinger, op. cit. \xvili. p. 359 (1873), Ixxvill.
p- 356 (1879).

Blastoceros azarae, Fitzinger, op. cit. Ixxvili. p. 359 (1879).

Cervus comosus, Burmeister, Descript. Phys. Répub. Argent. vol. il. p. 465
(1879).

Oxotoceros campestris, Ameghino, Rev. Arg. Hist. Nat. vol. 1. p. 243
(1891).

Cervus (Blastoceros) campestris, Ihering, Mammiferos de S. Paulo, p. 15

(1894).
Plate XXII

Characters.—Much smaller than the last, and of the approximate size of
the European roe, although more delicately and slenderly built. Antlers of
moderate size, with the lower or front prong of the main fork simple, and
the upper or posterior prong divided, and thus much the more complex,
the usual number of tines being three. Muzzle rather pointed; ears
moderately large and filled with white hair; tail moderately bushy. Hair
short and smooth, that of the middle of the back forming a whorl, and a
second whorl at the base of the neck, so that the fur of the withers is directed
forwards.'. General colour of upper-parts and limbs light reddish brown,
with the individual hairs of the back light gray at the base, then darker
gray, followed by a ring of reddish brown, and the tip black ; face darker,
occasionally a black patch on the crown of the head, extending backwards
as a line to the level of the ears ; tarsal tuft, a patch at the base of the back
of the ears, a ring round the pedicles of antlers, another round each eye, the
lips, throat, chest, under-parts, front and inner side of thighs, of buttocks, and
of upper part of fore-legs, whitish ; flanks, outer side of limbs, and middle
of throat lighter than back ; tail dark blackish brown above and white
below. Upper canines generally present in the male. Very young fawns
lighter coloured, with a row of white spots on each side of the back, and a
second row running from the shoulder to the thigh.

! A similar peculiarity occurs in Cervus albirostris.

Tue Deer oF Att Lanps. Pirate XXII.

oS

4

PAMPAS DEER.

Pablished by Rowland Ward Led . :

Pampas Deer 289

The above description is taken from Paraguayan and Argentine ex-
amples,—the B/astoceros azare of Fitzinger. Inthe true Brazilian campestris,
which the latter writer regards as a distinct species, the colour tends more
to yellowish gray, and upper canines are stated to be always wanting.
These, however, are occasionally not developed in males from Argentina
and Paraguay. Fitzinger states that the Brazilian form lacks the strong
scent of the Argentine race, but this is denied by Dr. Goeldi. In any case,
' the difference between the northern and southern races is not of more than
sub-specific importance. A young specimen from Santa Fe, in the chaco
districts of Argentina, living at Woburn Abbey in 1897, and forming the
subject of Fig. 76, had a very strongly marked black patch and line on the
crown of the head, and the white of the buttocks bordered by a thin black
line ; the base of the upper surface of the tail being also black, and the
general colour brown fawn. Such slight differences can, however, scarcely
be regarded as indicating even a distinct sub-species.

The Brazilian Cervus comosus was regarded by Fitzinger as a smaller and
longer-haired form allied to the marsh-deer ; but Burmeister has shown
that it was founded on an old male of the present species, in which, as is
commonly the case in aged examples of that sex, the hairs on the buttocks,
tail, and under-parts had become greatly elongated.

A fraction over 144 inches is the length of the longest pair of antlers of
this species recorded in Mr. Rowland Ward’s book ; the specimen being
in the British Museum.

Although this species is universally known by the specific name of
campestris, there is no justification for the suppression of the Linnean
bezoartica, which was applied to a South American deer with cylindrical,
three-tined antlers ; such deer being evidently the present species. It. may
be added that although the names campestris and /eucogaster date from the
same year, the former is the earlier, as it is quoted in Schreber’s work, in
which the latter is given for the first time.

Distribution —The open campos of Brazil, Paraguay, and Uruguay to the
pampas of Argentina and Northern Patagonia, and also extending into the
chaco, or wooded country of Argentina in the neighbourhood of Santa Fe.

Habits —The veado campeiro, as this species is called in parts of Brazil,
differs from the veado galheiro by being an inhabitant of dry open plains,
and also by the strong disagreeable odour emitted by bucks after their first

2P

290 American Deer

year. It never enters the forests and thickets of Brazil and Paraguay, but
restricts itself to the open campos which extend here and there among the
forest tracts, and are also the habitat of the Brazilian rhea. On the
Argentine pampas the animal is found in more or less entirely open country.
Formerly, when the tussocks of tall pampas-grass were dotted more or
less thickly over all the plains, it had plenty of covert ; but in the more
settled districts it now has to live almost completely in the open, and has

Fic. 76.—Young Male Pampas Deer. From a photograph by the Duchess of Bedford.

consequently become wary in the extreme. The presence of a buck,
especially in the evening, may, however, often be detected by the character-
istic odour which will be perceptible at the distance of a mile or so.
These deer may be found either in pairs, in parties of three or four, or in
small herds, but the old bucks are often or always solitary for the
greater part of the year. ‘Their times of feeding are much the same as
those of the marsh-deer, the animals remaining concealed under tussocks
of pampas-grass, or in such other shelter as they can find, and issuing forth

Extinct Species 291

to graze in the evening. Their alarm-cry is a low, whistling bark. When
a hind with her fawn is surprised by hunters, she stands stock-still until
her offspring has stolen quietly away and concealed itself, after which she
moves off in the opposite direction, at first slowly, and sometimes even
with a limping gait, in order to draw away the hunters from the hiding-
place of her young one.

In certain parts of Brazil this deer is known as the guazuti, but in
‘Uruguay, where it has now become very scarce, it is termed gama. If
taken sufficiently young, it becomes excessively tame, wandering about an
estancia among the men, horses, cattle, and dogs with the most complete
unconcern. The presence of strange men or dogs at once, however,
causes alarm. In the domesticated state, it will eat almost any kind of
vegetable food, either fresh or cooked ; and, like most of the deer tribe,
displays great partiality for salt. Its speed is considerable, and it requires
a good horse to overtake it ; by the natives it is captured either with the
bolas, or, less commonly, with the lasso.

3. THe Lujan Marsu-DEER—MaAzaMA BRACHYCEROS (Extinct)

Cervus brachyceros, Gervais and Ameghino, Mamm. Foss. Amer. Sud.
p- 126 (1880).

Cervus lujanensis, Ameghino, Diagnosis Mam. Fos. Neuv. p. 12 (1888),
Mam. Fos. Repub. Argent. p. 603 (1889).

Cervus paleoplatensis, Ameghino, op. cit. p. 13 (1888), and 604 (1889).

Cariacus brachyceros, Lydekker, An. Mus. La Plata—Pal. Argent. vol.
i. p. 79 (1893).

Characters—There are several species of extinct deer from the
Plistocene deposits of Buenos Aires more or less nearly allied to the marsh-
deer, but with more complicated antlers, some of which may be noticed
here. In the present species the antlers are very large, with the posterior
prong of the main fork greatly exceeding the anterior one in length and
complexity, and thus to a great extent obscuring the dichotomous forking.
In adult antlers, of which the inner view of a specimen from the left side
is shown in the accompanying figure, the front or lower prong of the
main fork is simply bifurcated, but the hinder prong divides into no less

292 American Deer

than six tines, of which all but one are on the front edge. The whole
antler is much flattened, and has its main fork only a short distance above
the burr. It is a much more complicated type than that of the existing
marsh-deer ; and indicates that the symmetrically dichotomous form of the
latter is a feature of no very great

— —~ ~

structural importance.
Distribution —Argentina during the
Plistocene period.

4. THe Buenos Aires MArsH-DEER—
MaAZzAMA ULTRA (Extinct)

|
Cervus ultra, Ameghino, Diagnosis
Mam. Fos. Neuv. p. 14 (1888).

Antifer ultra, Ameghino, Mam. Fos.
Repub. Argent. p. 610, plate xxxviil,
fig. 2 (1889).

Cariacus ultra, Lydekker, An. Mus.
La Plata—Pal. Argent. vol. i. p. 83
(1893).

Characters. —'This species, which
forms the type of the genus Antifer,
and was founded upon part of a left
antler from the Plistocene formation

Si pe

of Buenos Aires, appears to have been
Fic. 77.—Inner view of Left Antler of Lujan allied to the marsh-deer, although (or
Marsh-Deer. From a specimen in the La ‘ 5 5
considerably larger size, and with the
antlers much flattened. The antler
forks in the usual dichotomous manner ; the hinder tine of the terminal
fork of the posterior branch of the main fork is again divided, but the
front tine of the same is simple. The fragment of an antler on which the
genus Epreuryceros was based appears to be merely the basal portion of an
old antler of the present species; and if this be so, the main bifurcation
took place nearer to the burr than is the case in the living marsh-deer.

Distribution.—Argentina during the Plistocene epoch.

Plata Museum.

Extinct Species 2.93

5. AMEGHINO’s MarsH-DrER—MaAzaMa FRAGILIs (Extinct)

Cervus fragilis, Ameghino, Diagnosis Mam. Fos. Neuv. p. 11 (1888).

Paraceros fragilis, Ameghino, Mam. Fos. Repub. Argent. p. 606, plate

xxxvin, fig: 7 (1889).

Cariacus fragilis, Lydekker, dn. Mus. La Plata—Pal. Argent. vol. i.

p- 82, plate xxx (1893).

Characters.—The type of this well-
marked species is an imperfect left
antler from the superficial deposits of
the pampas of Buenos Aires. The
beam is cylindrical, and the main
bifurcation takes place about two
inches above the burr. ‘The anterior
branch of the main fork is given off
at an obtuse angle to the posterior
one, and is apparently simple. The
much larger posterior branch forks
about five inches above the main
bifurcation, and the hinder tine again
divides dichotomously. The figured
specimen, from which the front prong
of the main fork is broken away, is
one of a pair in the museum at La
Plata. Both antlers show five points,
and apparently indicate a species allied
to the pampas deer, but having antlers
of a more complex type.

Distribution. — Argentina during
the Plistocene epoch.

ae S Wavftyee

Fic. 78.—Front view of Left Antler of Ameghino’s
Marsh-Deer. From a specimen in the La Plata
Museum.

iii. THE GuemMAL Group—Sus-GENus XENELAPHUS

Hippocamelus, Leuckart, De Equo bisulco Molina, p. 23 (1816).

Cervequus, Lesson, Nouv. Tabl. Hist.

Nate pi t7 30 (842).

294 American Deer

Furcifer, Sundevall, K. Svenska Vet. Ak. Handl. for 1844, p. 183
(1846), in part, as a sub-genus ; Gray, Proc. Zool. Soc. 1850, p. 236, Cat.
Ungulata Brit. Mus. p. 226 (1852), Cat. Ruminants Brit. Mus. p. 88
(1872), dun. Mag. Nat. Hist. ser. 4, vol. xilt. p. 331 (1074); Brooke:
Proc. Zool. Soc. 1878, p. 923; Riitimeyer, 6h. schweiz. pal. Ges. vol. viii.
p. 49 (1881) ; mec Fitzinger, 1843.

Xenelaphus, Gray, Proc. Zool. Soc. 1869, p. 498, Cat. Ruminants Brit.
Mus. p. 89 (1872), Ann. Mag. Nat. Hist. ser. 4, vol. xiii. p. 331 (1874).

Anomalocera, Gray, Scientific Opinion, 1869, p. 384; Philippi, Wiegmann's
Archiv, 1870, p. 46; nec Templer, 1837.

Huamela, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. xi. p. 217 (1873) ;
Hand-list Ruminants Brit. Mus. p. 159 (1873).

Creagoceros, Fitzinger, SB. 4k. Wien, vol. \xviii. part i. p. 358 (1873),
Ixxviii. part 1. p. 369 (1879).

Characters —Antlers small and simple, forming a single dichotomous
fork, of which the front prong is the smaller, and curves upwards and
backwards towards the hinder one. Metatarsal gland and tuft absent.
Tail short and rather bushy. Face moderately long ; face-gland large and
exposed, and the gland-pit in the skull deep and triangular, but not very
large. Upper canines present in both sexes, although not projecting
beyond the lips. Fur very coarse and brittle. Size medium. Fawns
uniformly coloured.

This sub-genus is well distinguished by the simply forked antlers, the
coarse brittle hair, and the absence of the metatarsal gland. The absence
of the latter, and the uniform coloration at all seasons and all ages are
evidently specialised features ; and it accordingly seems that the simple
form of the antlers is due to degradation, and cannot therefore be regarded
as indicating the ancestral type from which the more complex forms of
Mazama antlers have been evolved.

The synonymy of the sub-genus and its two existing species is singularly
involved and complex ; the clearing of the web of confusion being mainly
due to Mr. Sclater. As noted below, the early names Hippocamelus and
Cervequus are rejected on account of their inappropriate nature. The name
Furcifer being preoccupied, and also apparently first applied to the pampas
deer, Xenelaphus and Anomalocera come next, the second of these being
likewise barred by an earlier use. Although Xenelaphus was originally

PERUVIAN GUEMAL.

Published by Rowland Ward, Lid

Tue Deer or Aut Lanps. PuatE XXII.

Guemals 2.95

applied to a specimen with abnormal antlers, this affords no reason for its
rejection, and Fitzinger’s creation of Creagoceros is therefore quite
superfluous.

Daistribution.—W estern and southern portions of the Neotropical region.

1. THE PERUVIAN GUEMAL—MAZAMA ANTISIENSIS

Cervus antisiensis, D’Orbigny, Arch. Mus. Parts, vol. ili. p. gt (1834) ;
Sclater, Proc. Zool. Soc. 1875, p. 46; Matschie, SB. Ges. nat. Berlin, 1894,
p02.

Cervus (Elaphus Furcifer) antisiensis, Wagner, Schreber’s Sdugethiere, vol.
iv. p. 384 (1844).

Cervus (Furcifer) antisiensis, Sundevall, K. Vet. Ak. Hand. for 1844,
Pp! 183 (1846).

Anomalocera huamel, Gray, Scientific Opinion, 1869, p. 384.

Xenelaphus huamel, Gray, Proc. Zool. Soc. 1869, p. 497.

Xenelaphus leucotis, Gray, Cat. Ruminants Brit. Mus. p. 89 (1872).

Xenelaphus anomalocera, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. x.
Pp: 445 (1872).

Xenelaphus chilensis, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. xii. p. 61
(1873), Hand-list Ruminants Brit. Mus. p. 159 (1873).

Creagoceros antisiensis, Fitzinger, SB. Ak. Berlin, vol. \xvili. part 1.
Peg 5o(lo7 2). xxvil. part 1. p. 369 (1879).

Furcifer chilensis, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. xill. p. 332
(1874).

Cariacus antisiensis, Brooke, Proc. Zool. Soc. 1878, p. 924; Schaft, Zoo/.
Garten, vol. xxxi. p. 226 (1890).

Furcifer antisensis, Nehring, SB. Ges. nat. Berlin, 1895, p. 9.

Plate XXIII

Characters.—Size somewhat less than that of the Virginian deer.
Build rather stout and heavy ; ears long, narrow, and sharply pointed, well
haired on both sides, and the hair at the base on both surfaces elongated ;
hair short, thick, rough, and brittle, longest between the ears, on the fore-
head) vand’ on. the tail; Pelage of upper-parts and outer side of limbs
coloured a mixture of brown and yellowish white, so as to give a general

296 American Deer

speckled yellowish gray-brown appearance ; the flanks, head, and forehead
speckled brownish white ; the individual hairs are whitish at the root, then
ringed with brownish, followed by yellowish white, and terminating in a
black tip, but on the sides of the head and forehead the tips are white. A
dark median line running down the forehead to the tip of the nose ; under
surface of lower jaw, throat, part of fore-neck, lower neck, hinder portion
of abdomen, and inner sides of buttocks and limbs white ; chest and under-
parts uniform brownish, darker than the back; lower part of limbs
yellowish gray-brown externally ; tail with a narrow brown and white
streak along the middle of the terminal portion of the upper surface, at the
base, sides, lower surface, and tip white; ears grayish- brown externally,
with a grayish-white heart-shaped spot at the base. Tarsal tuft dark
blackish brown. Female smaller and darker than the male. In the
specimen shown in plate xxiii the under surface of the tail is as described
above, but in a second skin in the Museum it is chestnut-brown.

Distribution —Andes of Peru, Ecuador, Bolivia, and Northern Chili,
usually at heights of between 14,000 and 16,000 feet above the sea-level,
but probably ranging down to 11,000 feet. In Ecuador abundant on
Chimborazo, Pechincha, and Cotopaxi.

2. THe CHILIAN GUEMAL—MAZAMA BISULCA

Equus bisulcus, Molina, Saggio Storia Nat. Chil, p. 320 (1782).

Hippocamelus dubius, Leuckart, De Equo bisulco Moline, p. 24 (1816),
ists, 1825, p..362;

Auchenia huamel, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 764
(1827).

Cervequus andicus, Lesson, Nouv. Tabl. Hist. Nat. p. 173 (1842).

Cervus chilensis, Gay and Gervais, Ann. Sci. Nat. 1846, p. 91 ; Sclater,
Proc. Zool. Soc. 1875, p. 45; Burmeister, Descript. Phys. Répub. Argent. vol.
lil. p. 462 (1879).

Capreolus leucotis, Gray, Proc. Zool. Soc. 1849, p. 64.

Furcifer huamel, Gray, Proc. Zool. Soc. 1850, p. 236, Cat. Ungulata
Brit. Mus. p. 227 (1852).

Furcifer antisiensis, Gray, Cat. Ungulata Brit. Mus. p. 226 (1852), Gat.
Ruminants Brit. Mus. p. 88 (1872).

Guemals 207

Huamela teucotis, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. xi. p. 219
(1873), Hand-list Ruminants Brit. Mus. p. 160 (1873).

Creagoceros chilensis, Fitzinger, SB. Ak. Wien, vol. |xviii. part i. p. 358
(Ho72) ai xxvill. parti. p.. 372 (1879).

Cartacus chilensis, Brooke, Proc. Zool. Soc. 1878, p. 923.

Furcifer chilensis, Sclater, List Anim. Zool. Gardens, p. 178 (1883) ;

Nehring, SB. Ges. nat. Berlin, 1895, p. 12.
_ Characters —Nearly allied to the last, but of smaller size, and dis-
tinguished by the following details of form and coloration. Antlers stouter,
with the front prong more decidedly shorter than the hinder one, and the
fork occurring at a considerable distance above the burr, instead of close to
it. Fore part of face relatively deeper, with a distinctly convex profile ;
eyes placed wider apart. No dark line from the forehead to the nose ;
throat and fore-neck yellowish brown like the rest of the neck; only a
small portion of the inner surface of the thigh and upper part of the
fore-leg white, the remainder of the limbs being yellowish brown.

Distribution —The Andes of Southern Chili and the whole of Patagonia ;
the animal being far more abundant in the southern than in the northern
portion of its range.

The history of this species is somewhat remarkable. Ancient travellers
in Chili brought reports of a cloven-hoofed animal known to the natives as
guemal, or huemal. This animal Molina regarded as a horse, to which
he gave the name Equus bisu/cus ; although his description is very vague,
the specific name is adopted. Leuckart renamed it Hippocamelus dubius,
both names being objectionable. Hamilton Smith regarded it as a llama,
under the name of duchenia huamel. Lesson’s generic name of Cerveguus
is nearly as objectionable as Hippocamelus.1

This deer is an inhabitant of the mountain-valleys of the Cordilleras in
situations where dense forests afford it a suitable habitat ; during the winter,
however, it descends to the plains, but never wanders any distance from the
foot of the mountains. In Western Patagonia it is by no means uncommon ;
and it is there hunted by the Indians, who call it guamul, guemul, or
huamel, and bring down the skins to Carmen and Bahia Blanca on the
Atlantic sea-board.

1 If this name is adopted, it antedates Mazama, and therefore will have to stand for all the American
deer, if they are included in one genus.

2 Q

298 American Deer

3. THe PuisrocEnE GuEMAL—MazaMaA SENELITICA (Extinct)

Cervus seneliticus, Ameghino, Diagnosis Mam. Fos. Nuev. p. 14 (1888).

Furcifer seneliticus, Ameghino, Mam. Fos. Repub. Argent. p. 612, plate
XXxix, fig. 4 (1889).

(?) Furcifer sulcatus, Ameghino, op. cit. p. 611, plate xxxix, fig. 3 (1889).

Cartacus seneliticus, Lydekker, dn. Mus. La Plata—Pal. Argent. vol. ii.
p- 53 (1893).

Characters.—Nearly allied to the existing representatives of the group,
but with larger antlers. The species was founded on a skull showing the
simple anterior branch of the antlers, which curves upwards and forwards,
starting at right angles to the posterior branch, which is broken away on
both sides. The left frontlet and antler forming the type of F. sw/catus
does not appear to be specifically distinct.

Remains of one or other of the existing species occur in the superficial
deposits of Ecuador.

Distribution. — Argentina during the Plistocene epoch.

iv. THe TypicaAL, oR BRockKET GRoup—Sus-Genus MAzamMa

Mazama, Rafinesque, Amer. Month. Mag. vol.1. p. 44 (1817) ; Merriam,
Serence, Ser. 2, VOI. A. p.22091(T305)).

Subulo, H. Smith, in Griffith’s Animal Kingdom, vol. v. p. 318 (1827),
as a sub-genus, nec Swbu/a, Schumacher, 1817; Fitzinger, SB. dk. Ween,
vol. luxix part d paulo 79):

Passalites, Gloger, Handbuch Naturgeschichte, p. 140 (1841).

Subula, Lesson, Nouv. Tabl. Hist. Nat. p. 174 (1842), mec Schumacher,
1817.

Coassus, Gray, List Mamm. Brit. Mus. p. 174 (1843), Proc. Zool. Soc.
1850, p. 240, Cat. Ungulata Brit. Mus. p. 238 (1852), Cat. Ruminants Brit.
Mus. p. 91 (1872); Brooke, Proc. Zool. Soc. 1878, p. 9245) Rutimever
Abh. schweiz. pal. Ges. vol. vill. p. 31 (1881).

Homelaphus, Gray, Cat. Ruminants Brit. Mus. p. go (1872).

Nanelaphus, Fitzinger, SB. Ak. Wien, vol. \xviii. part 1. p. 360 (1873),
Ixxix. part 1. p. 31 (1879), in part.

Doryceros, Fitzinger, op. cit. Ixviii. p. 360 (1873), xxix. p. 23 (1879).

Brockets 299

Characters —Antlers in the form of simple unbranched spikes. Meta-
tarsal, and in two instances also the tarsal, gland and tuft absent. Tail very
short. Face elongated ; face-gland small and exposed, and the gland-pit
deep and triangular ; the crown of the head tufted, and the hair of the face
radiating from two whorls, one of which is situated on the crown of the
head and the other below the line of the eyes, so that while the hair of the
forehead, as in other deer, is directed upwards towards the antlers, that of
the nose is directed downwards towards the muzzle. Upper canines occa-
sionally present in old males. Hair fine and smooth. Size small ; build
clumsy, with the back much arched; the profile of the face convex. Fawns
spotted with white. The ears are of medium length ; the upper lip has a
pair of white spots on each side of the nose, and the lower lip is marked by
a larger spot of white in the middle line.

It may be mentioned that the term brocket properly belongs to young
red deer stags, but in semi-scientific literature is now universally applied to
the members of the present group. Fossil remains of brockets occur in the
cavern-deposits of Lagoa Santa, Minas Geraes, Brazil, and likewise in the
Plistocene beds of the Argentine pampas, but their specific determination
stands in need of revision.

In regard to the specific representatives of the group, Sir Victor Brooke,
in 1878, wrote as follows :—It is now many years since I commenced the
study of this difficult group of the Cervide ; but although I have examined
the specimens contained in nearly all the continental museums, and made a
private collection of some importance, I must confess that I am still far
from a satisfactory understanding of the subject. The complete absence of
cornual and cranial characters renders it exceedingly difficult to grasp the
characteristic peculiarities of the different modifications of the form, six or
seven of which are, I think, probably persistent, and worthy of specific
recognition.” Fuller series of specimens are still urgently required before
our knowledge of the group can be regarded as approaching completion.
In the retention of a spotted coat by the fawns the sub-genus is less
specialised than the last; but the loss of the metatarsal gland and the
simple form of the antlers are specialised, or degraded, and not primitive
features. As already mentioned, the circumstance that species of the
Blastoceros group with antlers nearly or quite as highly developed as the
living forms occur in the Monte Hermoso beds of Argentina, which were

300 American Deer

deposited just about the time that deer had first reached South America,
and the fact that brockets are known only from Central and South America,
forms a sufficient proof that the group is a comparatively modern and
degraded one.

Mazama is taken as the name for the sub-genus, and hence for the
genus, on the authority of Dr. Merriam in the passage cited above. Sir
Victor Brooke rejected the earlier Sw4u/o in favour of the later Coassus on
account of its similarity to the still earlier Swbw/a. The genus Doryceros
was established by Fitzinger for the wood-brocket on account of the
absence of the tarsal gland, but since the metatarsal gland is so variable and
inconstant in the sub-genus Dorce/aphus, this can scarcely be regarded as a
character of even sub-generic value.

Distribution.—The hottest portions of the Neotropical region.

1. THe Rep BrockeET—MazAMA RUFA

Cervus rufus, F. Cuvier, Dict. Scz. Nat. vol. vii. p. 127 (1817) ; Bur-
meister, Descript. Phys. Répub. Argent. vol. iii. p. 465 (1879).

Cervus (Subulo) rufus, H. Smith, in Griffith’s Animal Kingdom, vol. iv.
p-. 140, v. p. 318 (1827) ; Goeldi, Mammiferos do Brasil, p. 108 (1893).

Subulo rufus, Jardine, Naturalists Library—Mamm. vol. iii. p. 178
(1835).

(?) Subulo apura, Swainson, Classif. Quadrupeds, p. 295 (1835).

Coassus rufus, Gray, List Mamm. Brit. Mus. p. 174 (1843), Cat. Ungu-
lata Brit. Mus. p. 238 (1852), Cat. Ruminants Brit. Mus. p. 92 (1872), Hand-
list Ruminants Brit. Mus. p. 161 (1873); Quelch, Zoo/ogist, ser. 3, vol. xvii.
p- 19 (1893).

Cervus (Subulo) dolichurus, Wagner, in Schreber’s Sdugethiere, vol. iv.
p- 389 (1844).

Subulo dolichurus, Fitzinger, SB. Ak. Wien, vol. \xviil. part 1. p. 359
(1873), lxxix. part 1. p. 10 (1879).

Subulo rufus, Fitzinger, op. cit. Ixviii. p. 360 (1873), xxix. p. 11
(1379).

Cariacus rufus, Brooke, Proc. Zool. Soc. 1878, p. 925.

Cervus (Coassus) rufus, Ihering, Mammiferos de S. Paulo, p. 15 (1894).

Brockets 301

Characters.—Size relatively large, and build heavy, the height at the
shoulder being about 27 inches. General colour of pelage glistening
brownish red, sometimes with minute dark tips to the hairs ; sides of neck
and flanks reddish gray ; throat, under surface of upper part of neck, and
inner side of thighs whitish gray ; tail brownish red above, white below and
at the tip ; a small whitish streak on the rump, a white spot on the front of
the lower lip, and a smaller spot on each side of the upper lip below the
nose ; antlers yellowish white.

The large size, heavy build, and uniformly brownish red coloration
at all seasons and all ages (except the very young fawns), are the most
distinctive features of this handsome species. The type specimen
was obtained from Dutch Guiana, and is preserved in the museum
aie acs:

Distribution. —Guiana, through Brazil, to Paraguay.

Habits—In Brazil, where it is known by the name of veado pardo, or
veado mateiro, this brocket is found alike in dense forests and on the open
campos, generally singly, and apparently only at certain seasons in pairs.
Like most deer, it is of an inquisitive, but at the same time of a very timid
and cautious disposition. It is nocturnal in its habits, issuing forth to feed
about sunset, and returning to its lair, which is generally under some thick
bush, soon after the sun is above the horizon. Various plants and the leaves
and young shoots of trees form its chief nutriment, but in the neighbour-
hood of cultivated lands it inflicts much damage on the crops, being very
partial to the young shoots of melons, budding maize, cabbages, and especi-
ally beans. The female gives birth, as a rule, only to a single fawn, which
is born in December or January ; the young animal being able to follow its
mother in from three to five days after birth. When danger is at hand, the
doe conceals her fawn in thick covert, and herself takes to flight. Where
the forest is not too thick, these brockets can be run down by good hounds
in half an hour or so ; if taken sufficiently young, the fawns can be readily

tamed.

2. THE STREAK-EYED BrocKET—MAaAZAMA SUPERCILIARIS

Coassus supercitiaris, Gray, Proc. Zool. Soc. 1850, p. 242, plate xxv, Ann.
Mag. Nat. Hist. ser. 2, vol. 1x. p. 432 (1852), Cat. Ungulata Brit. Mus.

302 American Deer

p. 239 (1852), Cat. Ruminants Brit. Mus. p. 92 (1872), Hand-list Ruminants
Brit. Mus. p. 160 (1873).

Cervus (Subulo) supercilaris, Wagner, Schreber’s Sdugethiere, vol. v.
p. 386 (1855).

Subulo superctharis, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 360
(1873), lxxix. part i. p. 18 (1879).

Cartacus superciliaris, Brooke, Proc. Zool. Soc. 1878, p. 926.

Characters.—Very closely allied to the last, of which it is not improb-
ably merely a local variety or colour-phase. General colour of pelage
shining brownish red ; neck, head, hocks, and front of fore-legs whitish
gray ; forehead blackish, with a distinct streak over the eye.

The type specimen is in the British Museum.

Distribution.—Probably Brazil.

3. THe Biack-racepD BrockKET—MazAMA TEMA

Mazama tema, Rafinesque, Amer. Month. Mag. vol. i. p. 44 (1817).

Cervus rufinus, Pucheran, Arch. Mus. Paris, vol. vi. p. 491 (1851).

Coassus rufinus, Gray, Hand-list Ruminants Brit. Mus. p. 162 (1873).

Subulo rufinus, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 360 (1873),
Ixxix. paft 1.\p\.17{ 1874).

Cariacus rufinus, Brooke, Proc. Zool. Soc. 1878, p. 925.

Cariacus tema, Alston, Bil. Centr. Amer.— Mamm. p. 118 (1879) ;
Rhoads, Amer. Naturalist, vol. xxvill. p. 526 (1894).

Characters—Smaller than the red brocket, the height at the shoulder
being about 25} inches. General colour of pelage bright shining brownish
red ; throat, chest, and under-parts blackish red, becoming darker on the
abdomen ; lower part of hind-leg, front and outer side of fore-leg, and lower
part of face shaded with bluish black ; hinder and inner side of fore-leg like
the under-parts, upper part of hind-leg like the back ; tail like back above,
white below; the usual white spots on the lips; antlers whitish horn-
colour, and less rugose than those of the red brocket.

The distinctive features of this species are the size, the dark shading of
the face and limbs, and the blackish red (instead of whitish) throat. The
type specimen is in the museum at Paris.

Distribution —Ecuador, at elevations of 12,000 feet and over.

ECUADOR

Tur DEER or Ati Lanps

PUDU (1) anp CENTRAL AMERICAN BROCKET (2),

Publi ghad by Kowland Ward Lia

. Prare XXD

Brockets 30

iGo)

4. THE CENTRAL AMERICAN BrocKET—MaAzaMA SARTORI

Cervus humboldtu, Wiegmann, Isis, 1833, p. 954,—no description.

Cervus sartoru, de Saussure, Rev. Mag. Zool. ser. 2, vol. xii. p. 252
(1860).

Subulo humboldtn, Fitzinger, SB. Ak. Wien, vol. \xxix. p. 20 (1879).

Cariacus rufinus, Alston, Biol. Centr. Amer.—Mamm. p. 118 (1879), nec

Pucheran.
Plate XXIV, fig. 2

Characters.—Distinguished from the preceding mainly, if not entirely,
by its inferior dimensions, the height at the shoulder being only 203
inches. The throat, neck, and chest are light fawn, and the abdomen
white ; the dark shading occupies the lower part of the face, the front of
the fore-legs, and the outer side of the hind-limbs.

This brocket, of which the type is in the Paris Museum, was long
confounded with the black-faced brocket ; and it should probably be re-
garded as a sub-species rather than a species. The figure in the plate is
taken from a mounted specimen now exhibited in the British Museum.

Distribution —Central America, especially Guatemala.

5. Tue Woop-BrockET—MazaMA NEMORIVAGA

Moschus americanus, Erxleben, Syst. Regn. Animal. p. 324 (1777).

Cervus nemorivagus, F. Cuvier, Dict. Sci. Nat. vol. vii. p. 485 (1817) ;
Ihering, Mammiferos de S. Paulo, p. 14 (1894), as synonym of swmplicicorns.

Cervus simplicicornis, Wied. Naturgeschichte Brasil, vol. 11. p. 596 (1826) ;
Burmeister, Descript. Phys. Répub. Argent. vol. i. p. 466 (1879) ; Goeldi,
Mammiferos do Brasil, p. 108 (1893).

Cervus (Subulo) nemorivagus, H. Smith, in Griffith’s Animal Kingdom,
Voleive P1425 Vie Pp. SI Qa(1O27).

Cervus (Subulo) simpheicornis, H. Smith, op. cit. vol. iv. p. 141, v. p. 381
(1827).

Passalites nemorivagus, Gloger, Handbuch Naturgeschichte, p. 140 (1841).

Coassus nemorivagus, Gray, Cat. Ungulata Brit. Mus. p. 238 (1852), Cat.

304 American Deer

Ruminants Brit. Mus. p. 91 (1872), Hand-list Rummants Brit. Mus. p. 160
(1873) ; Quelch, Zoo/ogist, ser. 3, vol. xvii. p. 19 (1893) ; Rendall, zd.
ser. 4; Vol. h. p. 245 (2897). :

Coassus simplicicornis, Gray, loc. cit.; Quelch, Zoologist, ser. 3, vol. xvii.
p. 19 (1893).

Doryceros nemorivagus, Fitzinger, SB. Ak. Wien, vol. \xviii. part i. p. 360
(1873), lxxix, parti.p. 29 (1079):

Cariacus simplicicornis, Brooke, Proc. Zool. Soc. 1878, p. 925.

Cartacus nemorivagus, Brooke, /oc. cit. (1878).

Cervus (Coassus) simplicicorms, Ihering, Mammiferos de S. Paulo, p. 15
(1894).

Characters.—The type of the genus Doryceros, which, according to
Fitzinger, is distinguished from all the preceding forms by the absence of
the tarsal gland and tuft. Size small, typically 19 inches at the shoulder.
General colour of the typical form varying from pale pepper-and-salt brown
to grayish or whitish, with a distinct streak on the forehead before the
front of the eyes; never any sign of reddish at any season; hairs of the
back ringed with yellowish red below the tip; antlers short, fine, and
dirty white in colour. Build light and slender.

In the form known as C. s¢mplicicornis, which is regarded by Fitzinger,
Goeldi, and Ihering as specifically inseparable from nemorivaga, although
kept distinct by Sir Victor Brooke, the streak on the forehead is wanting.
As both forms are met with in British Guiana, they seem scarcely worthy
even of sub-specific separation.

The type specimens of both forms are preserved in the Paris Museum,
that of memorivaga being from Guiana, and that of semplicicornis from Brazil.
The occurrence of both forms in British Guiana is mentioned by Mr.
Quelch in the passage cited. Five inches and one-eighth is the maximum
recorded length of the antlers.

Distribution.—Guiana, Colombia, Bolivia, Brazil, and Trinidad.

Habits.—In Brazil, where it is known as the vira, or veado catingueiro,
this brocket inhabits the open campos .and thin jungles of the interior,
avoiding the dense forests of the coast region. In Trinidad it abounds on
the borders of the high forests, where it inflicts much damage on young
plantations of cocoa and nutmeg. Its extreme wariness renders it difficult

of approach. The female produces a pair of fawns annually.

Brockets 205

6. THe PeruviAN BrocKET—MazAMA TSCHUDII

Cervus (Subulo) tschudu, Wagner, Schreber’s Saugethiere, vol. v. p. 386
(1855).

Cervus (Subulo) simplicicornis major, Wagner, Joc. cit. (1855).

Doryceros tschudu, Fitzinger, SB. Ak. Wien, vol. Ixviii. part i. p. 360
(1073) pilxxix. Part Lp. 30 (1879).

Coassus whitelyt, Gray, Ann. Mag. Nat. Hist. ser. 4, vol. xii. p. 163
(1873), Hand-list Ruminants Brit. Mus. p. 162, plate xxxii (1873).

Cartacus whitelyt, Brooke, Proc. Zool. Soc. 1878, p. 926.

Characters.—Closely allied to the wood-brocket, from which it is dis-
tinguished by its somewhat inferior size, the nearly smooth antlers, and
certain details of coloration. The pelage of the back is darker than in
M. nemorivaga, and the under-parts and inner side of the limbs pure white
instead of yellowish white; the hairs of the back lack the sub-terminal
yellowish red ring. C. white/yi was founded on an immature skull from
Peru in the British Museum, probably belonging to the present form. The
species is only known to me by description.

Distribution.—Peru, at elevations of 16,000 feet above the sea-level on
the western slopes of the Cordillera, in the same districts inhabited by
M. antisiensis.

7. THE Picmy BrockET—MazaMA NANA

Cariacus nanus, Lesson, Nouv. Tab/. Hist. Nat. p. 173 (1842).

Cervus (Subulo) nanus, Wagner, Schreber’s Sdugethiere, vol. v. p. 386
(1855).

Nanelaphus namby, Fitzinger, SB. Ak. Wien, vol. \xvi. part 1. p. 361
(ia74). ixxix.<part‘i. p. 32 (1879).

Nanelaphus nambi, Goeldi, Mammiferos do Brasil, p. 108 (1893).

Cervus (Coassus) nanus, Ihering, Mammiferos de S. Paulo, p. 16 (1894).

Characters.—Imperfectly known. Size very small, inferior to that of the
wood-brocket, from which this form is said to be distinguished by the
presence of a small tarsal tuft and the larger size of the face-gland. Tail
terminating in a tuft of longish hairs; hair rough and thick ; ears thickly
haired externally, short-haired internally. General colour of upper-parts

2R

306 American Deer

uniformly dark brown with a tinge of reddish ; under-parts lighter ; tail
coloured like the back above, white below and at the tip; a small white
spot beneath each eye; lips white, the upper one becoming dirty white
near the muzzle; ears externally dark brown, internally bluish white ;
tarsal tuft white.

This species is the type of Fitzinger’s genus Nane/aphus, in which the
Chilian pudu was also included. In Europe it appears to be represented
only by a specimen in the museum at Vienna, and further information is
required as to its affinities. Judging from the description, it seems to be
a brocket rather than a pudu; the tail being more developed than in the
latter, and the tarsal gland also present. ‘The white under surface of the
tail, and the white on the lips are also characters of the brockets, among
which group the species is placed by Dr. von Thering.

Distribution —Central Brazil, in the neighbourhood of Mato-Grosso,
where the species is known as noambibororoca, or veado pequeno. It lives
more generally on the borders of the campos than in the forests.

InceRTA SEDIS
1. Mazama inornata

Homelaphus inornatus, Gray, Cat. Ruminants Brit. Mus. p. 90 (1872).

Coassus inornatus, Gray, Hand-list Ruminants Brit. Mus. p. 162 (1873).

Named on the evidence of an immature stuffed male in the British Museum,
measuring rather over 24 inches in height. Gray’s description is as follows :—‘ It is
of a pale brown colour, more dusky on the head, back of the neck, shoulders, and
outside of the legs; a distinct yellowish streak over the upper edge of the orbit; the
inside of the upper part and the front edge of the thighs and the under side of the
tail white. It has no appearance of natural metatarsal glands. The ears are moderate
and nakedish. The nose and middle of the chin are dark blackish, with a large triangular
spot on the front of the upper lip,’ and a narrow white cross-band immediately under

the front of the lower lip.” The antlers are represented by small knobs.

2. Mazama aurita

Coassus auritus, Gray, Proc. Zool. Soc. 1850, p. 242, Ann. Mag. Nat. Fist. ser.
2, vol. ix. p. 432 (1852), Cat. Ungulata Brit. Mus. p. 239 (1852), Cat. Ruminants
Brit. Mus. p. 92 (1872).

' In the original misprinted /g.

Pudus B07

Cervus (Subulo) auritus, Wagner, Schreber’s Saugethiere, vol. v. p. 336 (1855).
Subulo auritus, Kitzinger, SB. Ak. Wien, vol. Ixviu. part 1. p. 360 (1873), Ixxix.

part 1. p. 19 (1879).

Founded on a skin reported to be from Brazil, but now lost ; not improbably the
female of M. rufa.

MI tae Pupus——Genus Pupua

Pudu, Gray, Proc. Zool. Soc. 1850, p. 242, Cat. Ruminants Brit. Mus.
Puo2zn1s72)).

Nanelaphus, Fitzinger, SB. dk. Wien, vol. Ixviil. part i. p. 360 (1873),
exit. pact. p- 3% (1879), in part.

imuadgeKaartod. Proc. Zool. Soc. 1877, p. 18; Brooke, zd. 1878,
p. 296 ; de Winton, sid. 1896, p. 508.

Characters—Skull and metacarpals generally as in Mazama. Size very
small ; hair coarse and brittle ; antlers in the form of short, simple spikes ;
cannon-bones very short; tail very short or wanting; no whorls in the
hair of the face, which is directed regularly backwards from the muzzle to
the crown; ears large and rounded; face-gland moderately large and
exposed, and the gland-pit very deep and oval; naked portion of muzzle
below nostrils deep ; tarsal and metatarsal glands and tufts absent ; upper
canines wanting, and the middle pair of lower incisors not greatly expanded ;
in the ankle-joint (tarsus) the external cuneiform united with the naviculo-
cuboid bone.

In the characters of the skull and several other features the pudus are
very closely connected with the guemals and brockets, from the latter of
which they are distinguished by the uniformly upward direction of the hair
of the face. This, however, is only a character of sub-generic importance
(according to the scheme of classification here employed) ; and the justifica-
tion for ranking the group as a separate genus rests upon the peculiar
structure of the ankle-joint, which is quite different from that of all the
other American deer.

Distribution.—W estern side of the Neotropical region.

ite Cumann LupUu-—euDUA PUDU

Capra pudu, Molina, Saggio Storia Nat. Chili, p. 310 (1782).
Ours pudu, Kerr, Animal Kingdom, p. 35 (1792).

308 Pudus

Cervus humilis, Bennett, Proc. Zool. Soc. 1831, p. 27.

Cervus (Pudu) humihs, Gray, Proc. Zool. Soc. 1850, p. 242.

Pudu humilis, Gray, Cat. Ungulata Brit. Mus. p. 240 (1852), Proc. Zool.
Soc. 1864, p. 105, Cat. Ruminants Brit. Mus. p. 93 (1872), Hand-lst
Ruminants Brit. Mus. p. 163 (1873).

Pudu chilensis, Gray, Cat. Ungulata Brit. Mus. plate xxxvi (1852).

Cervus pudu, Sclater, Proc. Zool. Soc. 1871, p. 238.

Nanelaphus pudu, Fitzinger, SB. Ak. Wien, vol. Ixviti. part 1. p. 361
(1873)2 lexi: part 1. po 34: (1070):

Pudua humilis, Garrod, Proc. Zool. Soc. 1877, p. 18; Brooke, wid. 1878,
p. 927; de Winton, zd. 1896, p. 510.

Coassus humilis, Ritimeyer, Abh. schweiz. pal. Ges. vol. viii. p. 31 (1881).

Characters.—The type species. Size very small, the height at the
shoulder being about 134 inches. Tail distinct. General colour of upper-
parts and outer surface of limbs reddish brown, with a speckling of fawn,
darkest on the head, passing into grayish on the neck, on the fore-neck
more distinctly speckled with yellowish fawn, and on the under-parts and
inner surface of the limbs fawn, yellower in some parts, and redder in
others. Ears relatively long. In the skull the premaxille separated by a
distinct interval from the nasal bones.

Distribution. —Chilian Andes and southwards to the archipelago of
Chiloe, descending in winter to the plains of the southern provinces.
Nothing seems to be recorded with regard to the mode of life of this pretty
little deer, which has several times been represented by living examples in
the London Zoological Gardens.

2. Tue Ecvuapor Pupvu—PupvuA MEPHISTOPHELES

Pudua mephistopheles, de Winton, Proc. Zool. Soc. 1896, p. 508, plate xix.
Plate XXIV, fig. 1

Characters—Known by a single immature female, measuring about 12
inches at the shoulder. Hair of the body long and coarse, its basal portion
peculiarly brittle and pith-like ; the terminal half of each hair black with
a ferruginous tip, producing a rich brown colour. Back darker than the
flanks, owing to the hairs having broader black bands with correspondingly

Musks 309

reduced coloured tips; on the neck the tips paler and longer, the black
being much reduced, which gives a tawny appearance ; towards the head
the black again increases, till the shorter hairs of the ears, face, and chin
are almost black. Ears thickly haired inside and out, the hairs on the inside
being broadly tipped with white. Feet black, most of the hairs being
minutely tipped with buff; inner sides of the legs and the abdomen
clothed with long yellowish hair of a finer and more ordinary character.
Ears very short and partially concealed by the rough hair. Naked
portion of the muzzle from the nostrils downwards deep. ‘Tail entirely
wanting. When full grown, the animal would probably be about 14 or
15 inches high at the shoulder, and thus taller than the Chilian species.

The distinctive features of this species are the absence of the tail, the
blackish-brown ground-colour sprinkled with bright rufous, and the nearly
black face and legs. The ears, too, are relatively short, with long fur,
especially internally, where they are white. In the skull the premaxille
ascend to reach the nasal bones, whereas in P. pudu the two are separated
by a short interval.

Distribution —Paramo of Papallacta, Ecuador, where it appears to be
very scarce.

B. Tue ABERRANT DEER—Sus-Famity MoscH1n&

Characters.—Distinguished from the Cervine by the following structural
features. The liver is provided with a gall-bladder, as in many hollow-
horned ruminants. In the skull the canal situated just within the margin
of the eye and leading into the chamber of the nose opens by a single
aperture, instead of two orifices, as in the true deer. There is no face-
gland or gland-pit below the eye. The hemispheres of the brain are
comparatively smooth, having few convolutions. There are also differences
in certain internal organs of the female. Antlers wanting in both sexes.

This sub-family contains only the musks, or musk-deer, of which there
appear to be two species.

That the musks are widely different from the typical Cervide, must be
freely admitted ; and were it not that the multiplication of groups is to be
deprecated, there is much to be said in favour of referring them to a family

apart (Moschide). ‘They were excluded from the Cervide by Fitzinger,

310 Musks

and Sir Victor Brooke seemed rather inclined to take the same view.
Garrod, also, was strongly against their inclusion in the Cervid@, remarking
that “when we consider the genus Moschus in its relations to the other
ruminants, it seems to me that to call it a deer is altogether against the
tendency of the facts at our disposal.” On the other hand, Ritimeyer
unhesitatingly classed the genus in the deer family, even going so far as to
place it next Hydre/aphus. But Sir William Flower, while also including
it in the Cervide, regarded it as a very early and distinct offshoot from the
other branch of that family. A somewhat similar relationship to the true
deer is presented by the giraffes, and it must be confessed that if the latter
are regarded as a distinct family, and not a sub-family of Cervida, it is a
little illogical not to give the same rank to the musks. Still, I am loth to
interfere with the system of classification generally adopted in England
more than seems necessary.

XIV. THe Musxks—Genvus Moscuus

Moschus, Linn. Syst. Nat. ed. 12, vol. i. p. gt (1766) ; Flower, Proc.
Zool. Soc. 1875, p. 159; Garrod, iid. 1877, p. 287; Riitimeyer, dA.
schweiz. pal. Ges. vol. viii. p. 19 (1881).

Characters.—Build stout and heavy, with the limbs long and thick, the
hinder pair considerably exceeding the front in this respect, and the rump
elevated. Hair coarse, thick, brittle, minutely waved, and in structure
resembling pith. Ears large. Upper canines greatly developed in the
males, and projecting far below the level of the lips, in females much smaller.
No tarsal or metatarsal glands or tufts ; lateral metacarpal bones represented
by their lower extremities ; main hoofs narrow and pointed, lateral hoofs
very large and functional. Tail very short and glandular, in the male
terminating in a tuft, in the female evenly haired throughout. The male
with a globular gland in the skin of the abdomen secreting the substance
known as musk, at least during the breeding-season. Naked portion of
muzzle large and completely surrounding the nostrils.

Among the many structural peculiarities of the genus, not the least
remarkable are those connected with the lateral digits of the limbs. In
the fore-limbs the joints of these digits are supplied with special tendons
from the deep flexor muscle of the leg, so that the lateral hoofs are capable

Himalayan Musk ee 6

of a large amount of movement and lateral expansion. <The special office

>

performed by the lateral phalanges,” writes Sir Victor Brooke, “is the
prevention of the animal slipping when rushing at speed down slopes or
sinking in swampy ground ; and this function is effected without danger of
the displacement of these delicate structures by means of strong ligamentous
bands developed in the sheath surrounding them, which bind them firmly
in their normal position.”

Distribution —Eastern part of the Eastern Holarctic, and some adjacent

districts of the Oriental region.

1. THE Himatavyan Musk—Moscuvus MoscHIFERUS

Moschus moschiferus, Linn. Syst. Nat. ed. 12, vol. 1. p..gt (1766) ; H.
Smith, in Griffith’s Animal Kingdom, vol. v. p. 301 (1827); Gray, Cat.
Ungulata Brit. Mus. p. 244 (1852), Cat. Ruminants Brit. Mus. p. 96 (1872) ;
Milne-Edwards, Ann. Sct. Nat. ser. 5, vol. it. p. 119 (1864), Rech. Mamm.
p- 176 (1868-74) ; Jerdon, Mamm. India, p. 266 (1867) ; Kinloch, Large
Game Shooting, part i. p. 41 (1869) ; Flower, Proc. Zool. Soc. 1875, p. 159 ;
Sterndale, Mamm. India, p. 494 (1884) ; W. L. Sclater, Cat. Mamm. Ind.
Mus. part ul. p. 172 (1891); Blanford, Fauna Brit. Ind—Mamm. p. 552
(1891) ; Biichner, Mélanges biol. Ac. St. Péersbourg, vol. xiii. p. 163
(1890-91).

Moschus sibtricus, Pallas, Spicil. Zool. vol. xiii. p. 29 (1799) ; Gray, Cat.
Ungulata Brit. Mus. p. 243 (1852).

Moschus altaicus, Eschscholtz, Isis, 1830, p. 606.

Moschus moschiferus altaicus, Brandt, Medicin. Zool. vol. 11. p. 347 (1833).

Moschus chrysogaster, Hodgson, ‘fourn. As. Soc. Bengal, vol. viii. p. 203
(1839) ; Gray, Cat. Ungulata Brit. Mus. p. 245 (1852), Cat. Ruminants
Bri. Mas. p. 97 41872).

Moschus leucogaster, Hodgson, ‘fourn. As. Soc. Bengal, vol. vill. p. 203
(1839) ; Gray, Cat. Ungulata Brit. Mus. p. 245 (1852), Cat. Rumuinants
Brit. Mus. p. 96 (1872).

Moschus saturatus, Hodgson, ‘fourn. As. Soc. Bengal, vol. vil. p. 203
(1839).

Moschus moschiferus fasciatus, Milne-Edwards, Ann. Sci. Nat. ser. 5, vol.
lil. p. 119 (1864) ; Gray, Cat. Rumiants Brit. Mus. p. 96 (1872).

312 Musks

Moschus moschiferus maculatus, Milne-Edwards, Ann. Sct. Nat. ser. 5, vol.
ii. p. 120 (1864) ; Gray, Cat. Ruminants Brit. Mus. p. 96 (1872).

Moschus moschiferus concolor, Milne-Edwards, Ann. Sct. Nat. ser. 5, vol.
ii. p. 121 (1864) ; Gray, Cat. Ruminants Brit. Mus. p. 96 (1872).

Characters.—Height at shoulder about 20 inches, at rump two inches
more. General colour of upper-parts some shade of rich dark brown, more
or less mottled and speckled with light gray, the individual hairs being
white for about three-quarters of their length, then with a white band,
followed by a blackish tip; under-parts and inner side of limbs paler ;
chin, inner border of ears, and inside of thighs whitish ; in some instances a
white spot on each side of the throat. Considerable individual or local
variation of colour is observable, and upon such differences a number of
nominal species or varieties have been described, although there is at present
little decisive evidence that any of these are entitled to rank as distinct sub-
species. While some individuals are paler than ordinary, others tend to the
development of a yellowish tint. The animal to which the name M.
chrysogaster was applied is depicted in Hodgson’s sketches as bright sepia-
brown sprinkled with golden red above, and golden red or orange beneath,
but the tints are probably exaggerated. In the form described as M@.
/eucogaster the upper-parts are very dark, with splashes of black on the back
and hips, while the under-parts are white or whitish. Other specimens,
again, are described as yellowish white all over the upper-parts, and white
beneath and on the inner side of the thighs ; and a variety from Kashmir
is stated to show lines of grizzled gray spots on the back. The fawns are
spotted with yellowish or pure white ; those from Kashmir territory being
of a paler tint than the individuals from the Eastern Himalaya.

Distribution.—F orest-clad districts in the Himalaya as far westwards as
Gilgit, at elevations of about 8000 feet or higher in summer, thence extend-
ing into Tibet, and so on to Siberia and Western China, where the species
is found in the southern districts of the province of Kansu. In the Kash-
mir area the range terminates with the northern limits of forest, the musk-
deer being quite unknown in the arid treeless districts of Dras and Ladak,
as it is in Baltistan.

Habits —Although formerly very abundant in the Himalaya, the musk-
deer has been so persecuted for the sake of its valuable secretion, that it is
a marvel it has not been exterminated. ‘Twenty years ago I found it still

Himalayan Musk 273

numerous in the more remote valleys north of Kashmir, such as Tilel and
Wardwan, but even in such districts it now appears to have become very
scarce, although the game laws recently put in force in Kashmir territory
may be doing something towards checking the diminution in its numbers.
Musk-deer are solitary animals, generally going about alone, and it is seldom, if
ever, that more than two adults are seen in company. Although they are
chiefly nocturnal, lying up during the greater part of the day under the

Fic. 79.—Musk Deer. From a photograph by the Duchess of Bedford.

shelter of some protecting bush, I have occasionally seen them in remote
districts feeding in the early afternoon. In Kashmir territory they are
chiefly found near the upper limits of the pine forests, and still more com-
monly in the still higher zone of birch, where the snow lingers till late in
the summer. On the Chinab I have, however, seen them in the dense
mixed forest at much lower levels, and in some districts they are met with
among the rhododendron forests. When disturbed, they start off in a series
of enormous bounds, halting after a few of such leaps to turn and gaze upon
the intruder, when they generally afford an easy shot. Cold seems to be a
DE

314 Musks

matter of absolute indifference to them, the thick coat of pithy hair afford-
ing an insulating protector capable of maintaining the natural heat in almost
any degree of temperature. In its agility and capacity of maintaining its
foot-hold on smooth and slippery surfaces of rocks the musk-deer is the
equal of the African klipspringer ; but the feet of the two animals display a
remarkable difference in their conformation. In the klipspringer the lateral
hoofs are minute and functionless, and the main hoofs short and rounded, so
that the animal rests on their tips alone. In the musk-deer, on the other
hand, both the main and the lateral hoofs are long and pointed, and the
animal appears to use the latter by grasping ridges or points of rock.

With regard to its food, General Kinloch states that this consists chiefly
of leaves and flowers, although another writer mentions grass and lichens ;
but the difference in these statements may be due to locality or season.
When standing at gaze, either before or after the first series of bounds,
musk-deer will occasionally utter a loud hiss, but, except for loud screams
when wounded or captured, no other cry seems to have been noticed.
From specimens kept in captivity many years ago in the residency at
Katmandu, Nepal, it appears that the pairing-season takes place in January,
and that the young are born in the following June. Although occasionally
there are twins, there is usually but a single fawn produced at a birth. To
make up for this slow rate of propagation, the fawns are able to breed
within a year from their birth.

The musk-pod of the males, which, as already stated, is situated on the
abdomen, is taken out entire and dried; the contents usually weighing
about an ounce, and selling for some sixteen or eighteen shillings. The
quantity of musk in the pod varies, however, according to the season and
the age of the animal from which it is taken. It has been stated that the
secretion is only yielded during the pairing-season, but if Hodgson be right
in saying that this occurs in January, the former statement cannot be
correct, as I have frequently shot musk-deer with full pods in May and
June. The flesh has no musky flavour, if the pod be removed immediately
after death, and makes excellent soup.

In addition to the large number shot, many musk-deer are captured
by means of snares in certain parts of the Himalaya. General Macintyre
describes the method as follows :—‘ A low fence is made of boughs, etc.,

along the ridge of a hill, sometimes a mile or more in length. At intervals

Kansu Musk eee is

of 100 or 150 yards are gaps. The musk-deer, crossing the ridge from one
valley to another, come across this fence, and to save themselves the trouble
of jumping over it, walk alongside until, seeing a little gap, they try to go
through it. But in each gap a noose of strong string is placed on the
ground, and tied to a stout sapling, bent downwards. The noose is so
arranged that when the deer tread inside it, the sapling is loosed and flies
back, leaving the noose tied tightly round the animal’s leg. The people
visit these fences every two or three days, and secure the deer thus caught,
and repair the fences and nooses, which are often carried away or destroyed
by larger game.”

The Hindustani name of the musk-deer is kastura. In English parks
these deer flourish well, and there are many at Woburn Abbey, and also at
Leonard’s Lee, Sussex. They show but little timidity, and allow them-
selves to be approached closely before starting off with the characteristic
bounds.

2. THe Kansu Musk—Moscuus sIFANICUS

Moschus sifanicus, Biichner, Mélanges biol. Ac. St. Pétersbourg, vol. xii.
p. 162 (1891, separate copies dated 1890).

Characters.—Size and general coloration similar to the last, but the ears
much longer, and, instead of being similar externally to the back, differently
coloured. Externally they are more or less completely deep black, or black
at the base with a broad yellowish tip, the margin of the upper half having
a blackish or brownish band; internally the margin is covered with
yellowish hair showing a more or less decided rufous tinge. The skull is
more massive, and longer in its anterior half, the nasal bones being
narrower, more elongated, and articulating with only a small portion of the
frontals.

There is no example of this form in the British Museum.

Distribution.—Southern Kansu, China, where the Himalayan species

also occurs.

“tl [Ve a 2
| eee
— - ‘

Ter P BIN DX

BIBLIOGRAPHY OF THE CERVIDZE

Tue following list, which makes no pretence to be complete, includes only works
and papers devoted exclusively, or almost so, to deer :—

Auston, E. R. On Female Deer with Antlers. Proc. Zool. Soc. 1879, pp. 296-299.

Banos, O. The Florida Deer. Proc. Soc. Washington, vol. x. pp. 25-28 (1896).

Barron, B. S. Account of the Cervus wapiti, or Southern Elk of Northern
America. Philad. Med. and Phys. fourn. vol. i. part 1. suppl. pp. 36-55
(1808).

Beavan, R.C. Notes on the Panolia Deer (Cervus eldi). Proc. Zool. Soc. 1867,
Pp: 759-766.

Biranrorp, W. T. On some Stags’ Horns from the Thian-Shan Mountains in
Central Asia. Proc. Zool. Soc. 1875, pp. 637-640.

- Exhibition of, and Remarks upon, two Heads and a Skin of the Yarkand Stag.

Ibid. 1892, pp. 116 and 117.

On a Stag, Cervus thoroldi, from Tibet, and on the Mammals of the Tibetan
Plateau. bid. 1893, pp. 444-449, plate xxxiv.

Bryru, E. A general Review of the Species of True Deer, or Elaphoid form of
Cervus, comprising those more intimately related to the Red Deer of Europe.
Journ, As. Soc. Bengal, vol. x. pp. 736-750 (1841).

Note on the Races of Reindeer. bid. vol. xxix. pp. 376-384 (1861).

—— Notes upon three Asiatic Species of Deer, Rucervus duvaucelli, R. schomburgki,
Panolia eldi. Proc. Zool. Soc. 1867, pp. 835-842.

Brooke, Sir V. On Hydropotes inermis and its cranial characters, as compared with
those of Moschus moschiferus. Proc. Zool. Soc. 1872, pp. §22-§25.

On Sclater’s Muntjac and other Species of the genus Cervulus. Ibid. 1874,
PP: 33-42-
——— On a new Species of Deer from Persia. Tc. pp. 42 and 43.

318 Appendix

Brookg, Sir V. Ona new Species of Deer from Mesopotamia. Proc. Zool. Soc.
1875, pp. 261-266.

——— Supplementary Note on Cervus mesopotamicus. Ibid. 1876, pp. 298-303.

—— On Cervus schomburgki (Blyth). T.c. pp. 304-307.

On the Deer of the Philippine Islands, with the Description of a new Species.
Ibid. 1877, pp. 51-60.

On the Classification of the Cervzde, with a Synopsis of the existing Species.
Ibid. 1878, pp. 883-928, plate lv.

Biicuner, E. Bemerkungen uber die Verbreitung des Edelhirsches im Ostlichen
Russland. dun. Mus. Zool. St. Pétersbourg, 1896, pp. 387-399.

Cameron, A. G. The value of the Antlers in the Classification of the Deer. Fe/d,
1892, April 30, p. 625 5; May 14, p. 7035 21, p. 7413 Jue 11, p.seo:

Deer and Deer-Stalking in the Scottish Highlands. Jéid. 1896, Aug. 8,
P: 222 215, p.267 522 1pi BOs 2954p. Bae

Caton, J.D. The Antelope and Deer of America. New York, 1877, 8vo.

Dawkins, W. B. Contributions to the history of the Deer of the European Pliocene
and. Miocene Strata. Quart. Fourn. Geol. Soc. vol. xxxvii. pp. 402-420 (1877).

—_——— British Pleistocene Mammalia—Part vi. Cervide. Mon. Pal. Soc. 1887.

Depéret,C. Nouvelles études sur les Ruminants pliocenes et quaternaires d’ Auvergne.
Bull. Soc. géol. France, ser. 3, vol. xii. pp. 247-284, plates v-viii (1884).

Evans, G. H. Notes on the Thamin, or Brow-antlered Deer. Yourn. Bombay Nat.
Hist. Soc. vol. ix. pp. 326-333 (1895).

Exiot, D. G. Remarks upon two Species of Deer of the genus Cervus from the
Philippine Archipelago. Pudl. Field Columbian Mus.—Zool. vol. 1. p. 157
(1897).

Firzincer, L. J. Die Gattungen der Familie der Hirsche (Cervz) nach ihrer natur-
lichen Verwandschaft. SB. dk. Berlin, vol. Ixviii. part 1. pp. 332-362 (1873).
— Kritische Untersuchungen tiber die Arten der naturlichen Familie der Hirsche
(Cervi). Ibid. vol. |xix. part 1. pp. 519-604 (1874), Ixx. part 1. pp. 239-333
(1874), Ixxviii. part 1. pp. 301-376 (1879), Ixxix. part 1. pp. 7-71 (1879).

Frower, W. H. On the structure and affinities of the Musk-Deer (Moschus moschi-
ferus, Linn.). Proc. Zool. Soc. 1875, pp. 159-190.

Fores, W. H. Supplemental notes on the Anatomy of the Chinese Water-Deer
(Hydropotes inermis). Proc. Zool. Soc. 1882, pp. 636-638.

Garrop, A. H. On the Chinese Deer named Lophotragus michianus by Mr.
Swinhoe. Proc. Zool. Soc. 1876, pp. 757-765.

Appendix 319

Garrop, A. H. Notes on the Anatomy of the Musk-Deer (Moschus moschiferus).
Ibid. 1877, pp. 287-292.

Notes on the Anatomy of the Chinese Water-Deer. .c. pp. 789-792.
Gray, J. E. Memoir on the genus Moschus, Linn., with descriptions of two new
Species. Proc. Zool. Soc. 1836, pp. 63-65.

Observations upon the Tufts of Hair observable upon the posterior Legs of
the genus Cervus, as a character of that group, and as means of subdividing it
into sections. Tc. pp. 66-68.

On a very large Roe-Deer (C. /eucotis) in the collection of the Earl of Derby.
Ibid. 1849, pp. 64 and 65.

Synopsis of the species of Deer (Cervina), with description of a new Species
in the Gardens of the Society. Jid. 1850, pp. 222-243, and Aun. Mag. Nat.
LADGRCCE 2 VOIX: pps 403-493 (1852 ),

On Rusa japonica, a new Species of Rusa Deer from Japan. Ann. Mag. Nat.
Hist. ser. 3, vol. vi. pp. 218 and 219 (1860).

——— Notice of a Stag from North China sent by Mr. Swinhoe to the Zoological
Society. Proc. Zool. Soc. 1861, pp. 236-238.

—-— On the Long-eared or Mule-Deer of North America. Aun. Mag. Nat.
Hist. ser. 3, vol. xviii. pp. 338 and 339 (1866).

On the Guemal or Roebuck of Southern Peru (Xenelaphus huamel). Proc.
Zool. Soc. 1869, pp. 496-499.

A new Deer from Chili (4nomalocera huamel). Scientific Opinion, 1869,
B99:

The Ahu (Capreolus pygargus). Ann. Mag. Nat. Hist. ser. 4, vol. x. pp.
407 and 408 (1872).

On the Guemal (Huamela leucotis). T.c. pp. 445 and 446 (1872).

Further remarks on the Guemal of Patagonia (Huamela leucotis). Ibid. vol.
Xi. pp. 214-220 (1873).

—— On the Wood-Deer of Brazil, Blastocerus sylvestris. Ibid. vol. xi. pp. 426
and 427 (1874).

——-—— On Xenelaphus, Furcifer, and Coassus peruvianus. Ibid. vol. xii. pp. 331 and
332 (1874).

Grevé, C. Das Elen in Aussterben. Zool. Garten, vol. xxxvi. pp. 267-269
(1895).

GrimB_eE, A. The Deer Forests of Scotland. London, 1896, 4to.

Hartine, J. E. The Deer of Epping Forest. Essex Naturalist, vol. 1. pp. 46-62
Gusis 7).

32.0 Appendix

Hevupe, P. M. Cerfs des Philippines et de L’Indo-Chine. Mém. hist. nat. emp.
Chinots, vol. 11. pp. 1-64 (1888).
—— Catalogue revisé des Cerfs Tachetés (Ska). Tic. pp. 146-168 (1892),
and 111. pp. 98-107 (1897).
Apercu sommaire du genre Hippelaphus. Ibid. vol. i. pp. 47-52 (1896),
and 92-97 (1897).
Hopeson, B. H. Contributions in Natural History, the Musk-Deer and Cervus
jaral. Gleanings in Science, vol. 1. pp. 320-324 (1831).

— Note relative to the account of the Jaral. ourn. As. Soc. Bengal, vol. 1.

pp. 66 and 67 (1832).

On the Ratwa Deer of Nepal (Cervus ratwa). Asiat. Researches, vol. xvii.
Pp- 139-149 (1833).

—__— Specific description of a new Species of Cervus (C. elaphoides). “fourn.
As. Soc. Bengal, vol. iv. pp. 648 and 649 (1835).

———— Notes on the Cervus duvaucelii, Cuvier, or C. elaphoides and bahraiya,
Hodgson. Jdid. vol. v. pp. 240-242, and Proc. Zool. Soc. 1836, pp. 46-47.
—_ On three new Species of Musk (Moschus) inhabiting the Himalayan Districts
(M. chrysogaster, M. leucogaster, M. saturatus). ‘ourn. As. Soc. Bengal,
vol. vill. p. 202 (1839).

——_— Note on the Cervus elaphus (elaphoides ?) of the Saal Forest of Nepal (hodie
C. affinis). Ibid. vol. x. pp. 721-724 (1841).

——— On a new Organ in the genus Moschus. T.c. pp. 795 and 796 (1841).

——— On a new Species of Cervus (C. dimorphe). Ibid. vol. xu. pp. 889-898
(1843).

—— On the Shou or Tibetan Stag (Cervus affinis). Ibid. vol. xix. pp. 388-394

(1851).

Hose, C. H. Description of a new Deer from Mount Dulit, Eastern Sarawak.
Ann. Mag. Nat. Hist. ser. 6, vol. xii. p. 206 (1893).

Huet, J. Liste des Espéces dans les Families des Cervidés, Cervulideés, Tragulidés, et
des Moschidés. Bull. Soc. Acclim. Paris, ser. 4, vol. v. pp. 274-284, 407-508,
5§45-555, and 721-737 (1888).

Keirwack, K. Ueber einen Damhirsch aus dem deutschen Diluvium. 7B. preuss.

geol. Landesanst. 1887, pp. 283-290, plate xi (1888).

Lockuart, J. G. Notes on the Habits of the Moose in the Far North of British
America in 1865. Proc. U. S. Mus. vol. xiii. pp. 305-308 (1891).

Lypekker, R. Onaremarkable Antler from Asia Minor. Proc. Zool. Soc. 1890,
pp- 363-365, plate xxx.

Appendix BoE

Lypekker, R. On an apparently new Deer from North China in the Menagerie of
the Duke of Bedford at Woburn Abbey. J/did. 1896, pp. 930-934, plates xlviii.

and xlix.

On the Deer allied to Cervus sica. Ibid. 1897, pp. 37-46, plate i.

Macpuerson, H. A., and others. The Red Deer. Fur and Feather Series.
London, 1896, 12mo.

Mancotp, C. Cervus alces in Aussterben begriffen. Zool. Garten, vol. xxxvi.

Pp- 157-159 (1895).

Mearns, E. A. Description of a new Deer (Dorcelaphus texanus) from
Texas and Northern Mexico. Proc. Soc. Washington, vol. xii. pp. 23-26
(1898).

Merriam, C. H. Cervus roosevelti, a new Elk from the Olympics. Proc. Soc.
Washington, vol. x1. pp. 271-275 (1897).

Meyer, A. B. Die Hirschgeweih-Sammlung im koniglichen Schlosse zu Moritzburg,
bei Dresden. 2 vols. fol., Dresden, 1883 and 1887.

Mitvats, J. G. British Deer and their Horns. London, 1897, 4to.

Mirne-Epwarps, A. Note sur le Mi-lou, ou Sseu-pou-siang (E/aphurus
davidianus), du Nord de la Chine. dun. Sci. Nat.—Zool. vol. v. pp. 380-382
(1866).

Moe renporrr, O. F. Ueber die Sikahirsche. Zool. Fahrb. 1887, pp. 588-590.

Moyjsisovics, A. Ueber die Geweihbildung des Hochwildes von Bellye. Mr. Ver.
Stetermark, 1888, pp. 53-73.

Miier, J. Ein Sumpfhirsch (Blastocerus paludosus, Gray) im Berliner Zoologischen
Garten. Zool. Garten, vol. xxxvii. pp. 49 and 50 (1896).

Neasrine, A. Ueber Furcifer antisiensis, d’Orb., and Cervus brachyceros, Philippi
SB. Ges. Berlin, 1895, pp. 9-18.

Nirscue, H. Studien uber das Elchwild, Cervus alces, Linn. Zool. Anzeiger, vol.
Xiv. pp. 181-191 (1891).

Bemerkungen tiber zwei aus Spitzbergen stammends Rentierschadel. Fh.

Ver. Nat. Wurit. 1893, pp. 111-127.

Noack, T. Zur Saugethierfauna der Mantschurischen Subregion—Part 1. Cervide.
Humboldt, vol. viii. pp. 1-10 (1889).

Poutic, H. Die Cerviden des thuringischen Diluvial-Travertines, mit Beitragen
uber anders diluviale und tber recente Hirschformen. Paleontographica, vol.
XXXIX. Pp. 215-262, plates xxiv-xxvii (1892).

Powerscourt, Viscount. On the Acclimatisation of the Japanese Deer at Powers-
court. Proc. Zool. Soc. 1884, pp. 207-209.

aly

22, Appendix

PucHEeRAN, J. Monographie des Espéces du genre Cerf. Arch. Mus. Paris,
vol. vi. pp. 264-492 (1852).

Quetcn, J. J. The Deer of British Guiana. Zoologist, ser. 35 vol. xvil. p. 19
(1893).

Rocer, O. Ueber die Hirsche. CB. Ver. Regensburg, 1887, pp. 50-93.

Riimeyer, L. Beitrage zu einer nattirlichen Geschichte der Hirsche. bh. schweiz.
pal. Ges. vol. vil. pp. 3-8 (1880), vill. pp. 9-97 (1881), X. pp. 3-192 (1883).

ScLaTeR, P. L. Note on the Japanese Deer living in the Society’s Menagerie. Proc.
Zool. Soc. 1860, pp. 375-377.

Note on the Deer of Formosa. Jdid. 1862, pp. 150-152, plates xvi and
XVil.

—— On certain species of Deer now or lately living in the Society’s Menagerie.
Trans. Zool. Soc. vol. vii. pp. 333-352, plates xxvili-xxxix (1871).

Remarks on Cervus chilensis and Cervus antisiensis. Ann. Mag. Nat. Hist.
ser. 4, vol. xi. pp. 213 and 214 (1873).
—— Remarks on the Deer of Chili and Western Peru. Proc. Zool. Soc. 1875.
PP: 44-47.
ScLtaTerR, W. L. Description of a Stag’s Head allied to Cervus dybowskii. Fourn.
As. Soc. Bengal, vol. lviii. pp. 186-188, plate xi (1889).
Scorr, W. B. Cervalces americanus, a fossil Moose, or Elk, from the Quaternary
of New Jersey. Proc. Acad. Philadelphia, 1885, pp. 181-202, plate i.
SwinHoe, R. On the Japanese and Formosan Deer. dun. Mag. Nat. Hist. ser. 3,
vol. vill. p. 192 (1861).
——— Letter with Observations on some Chinese Deer. Proc. Zool. Soc. 1864,
pp. 168 and 169.
—— On the Cervine Animals of the Island of Hainan (China). J/éid. 1869,
pp- 652-660.
— Ona new Deer from China, Hydropotes inermis. Ibid. 1870, pp. 89-92.
— On Chinese Deer, with the description of an apparently new species, Cervus
kopschiit. Ibid. 1873, pp. 572-576.
- On a small, tufted, Hornless Deer from the mountains near Ningpo (Lopho-
tragus michianus). Ibid. 1874, pp. 452-454.

Taczanowsk1, L. Description d’un nouveau Cerf tacheté du pays d’Ussuri
méridional (Cervus dybowskii). Proc. Zool. Soc. 1876, pp. 123-125.

Truz, F. W. Description of a new Species of Deer (Cariacus clavatus) from Central
America. Proc. U. S. Mus. vol. xi. pp. 417-424 (1888).

Appendix 32.3
WaLSINGHAM, Viscount.

On the Distribution of the different species of Deer and
other Ruminants in Northern California and Oregon.

Proc. Zool. Soc. 1873,
pp. 561-563.

Wuiraker, J. A Descriptive List of the Deer Parks and Paddocks of England.
London, 1892, 8vo.

Wrecmann, A. F. A. Eine neue Art des Hirsch-geschlectes.
970.

575,913.33) Pp= 9.52=

Fic. 80.—Skull and Antlers of Caspian Red Deer.

From a specimen shot in September 1891 by
Mr. St. George Littledale in the Western Caucasus, at an elevation of about 6000 feet.

ACAPULCENSIs, Cervus, 263

acuticornis, Dama, 196

acuticornis, Panolia, 196

affinis, Cervus, 88, 281

Ahu, 230

ahu, Cervus, 227

albicornis, Cervus, 144

albipes, Cervus, 204

albipes, Prox, 204

albirostris, Cervus, 91

Alce, 18, 49, 125

alce, Alcelaphus, 52

alce, Cervus, 52

Alcelaphus, 49

Alces, 18, 49

alces, Alces, 52

alces, Cervus, 52

alces latifrontis, Cervus, 59

alfredi, Axis, 172

alfredi, Cervus, 172

alfredi, Melanaxis, 172

altaicus, Moschus, 311

Altai wapiti, 164

ambiguus, Cervus, 239

Ameghino’s marsh-deer, 293

American deer, 243

americana couesi, Mazama, 261

americana gymnotis, Mazama, 265

americana macrura, Mazama, 257

americana, Mazama, 249

americana mexicana, Mazama, 261

americana nemoralis, Mazama, 264

americana osceola, Mazama, 259

americana peruviana, Mazama, 267

americana savannarum, Mazama,
266

americana texana, Mazama, 261

americana tolteca, Mazama, 263

americana typica, Mazama, 252

americanus, Alce, 52

americanus, Alces, 52

americanus, Cariacus, 249

americanus, Dorcelaphus, 249

americanus, Moschus, 303

Amphitragulus, 215

andicus, Cervequus, 296

anoceros, Dremotherium, 217

INS DEE OX

anocerus, Cervus, 217
anocerus, Dicrocerus, 217
Anoglochis, 19, 238
Anomalocera, 294
anomalocera, Xenelaphus, 295
Antifer, 282

antisiensis, Cariacus, 295
antisiensis, Cervus, 295
antisiensis, Creagoceros, 295
antisiensis, Furcifer, 295, 296
antisiensis, Mazama, 295
apura, Subulo, 300

arcticus, Rangifer, 47
arcticus, Tarandus, 47

Ardé deer, 239

ardeus, Anoglochis, 239
ardeus, Cervus, 239
argyropus, Hydropotes, 221
aristotelis, Cervus, 144
aristotelis heterocervus, Rusa, 144
aristotelis, Hippelaphus, 144
aristotelis leschenaulti, Rusa, 144
aristotelis nigra, Rusa, 144
aristotelis, Rusa, 144

aureus, Cervulus, 204
aureus, Stylocerus, 204
aurita, Mazama, 306

auritus, Cervus, 269, 307
auritus, Coassus, 306

auritus, Subulo, 307

Axis, I41

axis, Cervus, 179

axis ceylonensis, Cervus, 179
azare, Blastoceros, 288
azare, Cervus, 288

BAHRAINJA, Cervus, 189
barbarus, Cervus, 74
Barren-ground reindeer, 47
Basilan sambar, 158
Bavian deer, 174
bedfordianus, Cervus, 81
belgrandi, Cervus, 139
bezoartica, Mazama, 287
bezoarticus, Cervus, 287
bisulca, Mazama, 296
bisulcus, Equus, 296

Black-faced brocket, 302
Black-tailed deer, 278
Blastomeryx, 216
Blastoceros, 20, 282

Bokhara deer, 108
borbonicus, Cervus, 240
brachyceros, Cariacus, 291
brachyceros, Cervus, 281, 291
Brockets, 298

brookei, Cervus, 151
Brow-antlered deer, 197
browni, Cervus, 131

Brown’s fallow deer, 131
Buenos Aires marsh-deer, 292
Burmese thameng, 200

CaLaMIANES DEER, 173
Californian mule-deer, 276
cambojensis, Cervulus, 193
cambojensis, Rucervus, 193
campestris, Blastoceros, 287
campestris, Cariacus, 287
campestris, Cervus, 287
campestris, Dorcelaphus, 287
campestris, Furcifer, 287
campestris, Mazama, 287
campestris, Ozotoceros, 288
canadensis asiaticus, Cervus, 104
canadensis eustephanus, Cervus,
105
canadensis, Cervus, 94
canadensis, Elaphus, 94
canadensis luehdorfi, Cervus,
canadensis occidentalis, Cervus,
IOI
canadensis, Strongyloceros, 94
canadensis typicus, Cervus, 96
canadensis verus, Cervus, 96
caprea, Capreolus, 224
caprea, Capreolus, 224
Capreolus, 19, 222
capreolus, Cervus, 224
capreolus fossilis, Cervus, 224
capricornis, Cervus, 249
cariacou, Cervus, 263
cariacou, Reduncina, 263
Caribou, 37

326

caribou, Rangifer, 42
Cariacus, 248
carnutorum, Cervus, 140
cashmerensis, Cervus, 83
cashmeriensis, Cervus, 83
cashmirianus, Cervus, 83
cashmirianus yarcandensis, Cervus,
88
cashmirianus typicus, Cervus, 85
Caspian red deer, 75
caspicus, Cervus, 186
Catoglochis, 238
Central American brocket, 303
Central American deer, 264
cephalophus, Elaphodus, 213
Cervalces, 49
Cervequus, 293
Cervine, 17, 32
Cervulus, 19, 202
Cervus, 18, 61, 62
chilensis, Cariacus, 297
chilensis, Cervus, 296
chilensis, Creagoceros, 297
chilensis, Furcifer, 295, 297
chilensis, Pudu, 308
chilensis, Xenelaphus, 295
Chilian guemal, 296
Chilian pudu, 307
Chinese muntjac, 208
Chinese water-deer, 221
Chital, 179
chrysogaster, Moschus, 311
clactonianus, Cervus, 131
cladoceros, Cervus, 239
clavatus, Cariacus, 267
clavatus, Cervus, 249
Coassus, 298
Colombian deer, 266
columbiana, Mazama, 278
columbianus, Cariacus, 278
columbianus, Cervus, 278
columbianus, Dorcelaphus, 278
columbianus, Eucervus, 278
columbicus, Cervus, 265
Common American deer, 249
Common sika, 111
comosus, Blastoceros, 288
comosus, Cervus, 288
concolor, Moschus, 312
coronatus, Cervus, 52
Corsican red deer, 74
corsicanus, Cervus, 74
Cosoryx, 216
couesi, Dorcelaphus, 261
Creagoceros, 294
crinifrons, Cervulus, 210
crookei, Mazama, 268
Crooke’s black-tailed deer, 268
crooki, Dorcelaphus, 268
culionensis, Cervus, 173
curvostylis, Cervulus, 204

Index

| cusanus, Capreolus, 232
| cusanus, Cervus, 232
| cylindroceros, Cervus, 239

DacryLoceros, 125

| Dama, 18, 125

dama americana, Cervus, 249
dama browni, Cervus, 131
dama, Cervus, 127

dama, Dactyloceros, 127
dama, Dama, 127

dama, Palmatus, 127
Damine group, 125
davidianus, Cervus, 236
davidianus, Elaphurus, 236
dawkinsi, Cervus, 140
dejeani, Cervus, 156
Desert mule-deer, 277
dichotoma, Mazama, 283
dichotomus, Cervus, 283
dicranius, Cervus, 240
dicranoceros, Cervulus, 217
dicranoceros, Cervus, 217
Dicroceros, 19, 218
dimorpha, Axis, 190
dimorpha, Rusa, 189
dimorphe, Cervus, 189
dolichurus, Cervus, 300

dolichurus, Subulo, 300
Dorcelaphus, 20, 248
Doryceros, 299
Dremotherium, 19, 215
dubius, Hippocamelus, 296
Duke of Bedford’s deer, 81
duvauceli, Cervus, 189
duvaucellii, Axis, 189
duvaucellii, Rucervus, 189
dybowsku, Cervus, 91, 117

East American wapiti, 96
Ecuador pudu, 308

eedii, Panolia, 196

Elaphine group, 62
Elaphoceros, 110

Elaphodus, 19, 212
elaphoides, Cervus, 189
Elaphurus, 19, 233

Elaphus, 62

elaphus antiqui, Cervus, 75
elaphus barbarus, Cervus, 74.

| elaphus canadensis, Cervus, 94
elaphus, Cervus, 64

elaphus corsicanus, Cervus, 74
elaphus minor, Cervus, 74
elaphus germanicus, Cervus, 65
elaphus maral, Cervus, 75
elaphus typicus, Cervus, 65
elaphus varius, Cervus, 75
elegans, Dicroceros, 218

eldi, Cervus, 195

eldii, Panolia, 196

eldi platyceros, Cervus, 200

eldi typicus, Cervus, 200

Elk, 49, 52

ensenadensis, Paraceros, 283
Epieuryceros, 282

equina pennantii, Rusa, 151
equina, Rusa, 151

equinus, Cervus, 150

equinus, Hippelaphus, 151
etueriarum, Cervus, 122
euceros, Cervus, 189

Eucervus, 248

Eucladoceros, 238

eucladoceros, Cervus, 189
eudpis, Cervus, 112

europeus, Capreolus, 224
European red deer, 65

European roe, 224

euryceros belgrandi, Cervus, 139
euryceros carnutorum, Cervus, 140
euryceros, Cervus, 134
euryceros dawkinsi, Cervus, 140
euryceros germaniz, Cervus, 136
euryceros hiberniw, Cervus, 137
euryceros italiw, Cervus, 139
eustephanus, Cervus, 105

FaLLow DEER, 127
fasciatus, Moschus, 311
fee, Cervulus, 209

fitchii, Cervus, 140
Five-tined deer, 242
Florida deer, 259
Formosan sambar, 154
Formosan sika, 116
fossilis, Capreolus, 224
fragilis, Cariacus, 293
fragilis, Cervus, 293
fragilis, Mazama, 293
fragilis, Paraceros, 293
frontalis, Cervus, 196
frontalis, Panolia, 196
furcata, Mazama, 283
furcatum, Dremotherium, 217
furcatus, Palaomeryx, 217
furcatus, Prox, 217

Furcifer, 294

GIANT FALLOW DEER, 134

gigantea, Alce, 134

giganteus belgrandi, Cervus, 139

giganteus carnutorum, Cervus,
140

giganteus, Cervus, 134

giganteus italia, Cervus, 139

giganteus rufli, Cervus, 136

giganteus typicus, Cervus, 137

goudotii, Cervus, 265

Greenland reindeer, 47

Guazu deer, 282

Guemals, 293

guettardi, Cervus, 37
gunnii, Cervus, 140
Gymnotis, 248
gymnotis, Cariacus, 265
gymnotis, Cervus, 265

Harry-FRONTED MUNTJAC, 210

Hangul, 83

hastalis, Cervus, 42

hastalis, Tarandus, 42

hemionus californica,
276

hemionus, Cervus, 269

hemionus, Dorcelaphus, 270

hemionus eremica, Mazama, 277

hemionus eremicus, Dorcelaphus,

Mazama,

277
hemionus, Mazama, 269
hemionus peninsule, Mazama,
276

hemionus typica, Mazama, 275
Hensel’s muntjac, 217
heterocervus, Rusa, 144
hibernicus, Megaceros, 134
hibernus, Cervus, 134
Himalayan musk, 311
Hippelaphus, 141

hippelaphus, Cervus, 162
hippelaphus moluccensis, Cervus,

166
hippelaphus, Rusa, 162
hippelaphus timoriensis, Cervus,

170
hippelaphus typicus, Cervus, 164
Hippocamelus, 293
Hog-deer, 175
Homelaphus, 298
hortulorum, Cervus, 117
Huamela, 294
huamel, Anomalocera, 295
huamel, Auchenia, 296
huamel, Furcifer, 296
huamel, Xenelaphus, 295
humboldtii, Cervus, 303
humboldtii, Subulo, 303
humilis, Cervus, 308
humilis, Pudua, 308
Hydrelaphus, 19, 219
Hydropotes, 219
Hyelaphus, 142

INDIAN SAMBAR, 146

Indian spotted deer, 179
inermis, Hydrelaphus, 221
inermis, Hydropotes, 221
inornata, Mazama, 306
inornatus, Coassus, 306
inornatus, Homelaphus, 306
Irish elk, 134

islandicus, Cervus, 134
issiodorensis, Cervus, 122

Index

isubra, Cervus, 102
Isubra deer, 103

JAPANESE SIKA, I12
japonica, Rusa, 112
japonicus, Cervus, III
jarai, Cervus, 144
jarya, Rusa, 144

Javan sambar, 162, 164
Johnson’s elk, 59
jubata, Alces, 52

Kawnsu Musk, 315
Kashmir stag, 85
Kaup’s muntjac, 217
kopschi, Cervus, 114
kuhli, Cervus, 174
kuhli, Rusa, 174

LACHRYMans, Cervulus, 208
La Paz mule-deer, 276
latifrons, Alces, 59
latifrons, Cervus, 59
lepida, Rusa, 185

lepidus, Cervus, 185
leptocephala, Mazama, 281
leptocephalus, Cariacus, 281
leschenaulti, Hippelaphus, 144
leschenaultii, Cervus, 144
leucogaster, Cervus, 287
leucogaster, Moschus, 311
leucotis, Capreolus, 296
leucotis, Huamela, 297
leucotis, Xenelaphus, 295
leucurus, Cariacus, 258
leucurus, Cervus, 257
leucurus, Mazama, 258
lewisii, Cervus, 278

lobata, Alces, 52

lobata coronata, Alces, 52
lobatus, Cervus, 52
Lophotragus, 212
luehdorfi, Cervus, 102
lujanensis, Cervus, 291
Lujan marsh-deer, 291
Luzon sambar, 157

lyratus, Cervus, 196

MACHLIs, Alces, 52

Macrotis, 248

macrotis californicus,
276

macrotis, Cariacus, 269

macrotis, Cervus, 269

macrotis columbianus, Cariacus,
278

macrotis columbianus, Cervus, 278

macrotis, Dorcelaphus, 269

macrotis, Eucervus, 269

macrotis, Mazama, 269

macrotis montanus, Cervus, 275

Cariacus,

1

ehaaf

macrotis, Otelaphus, 269
macrourus, Cervus, 257
macrourus, Mazama, 257
macrurus, Dorcelaphus, 258
maculata, Axis, 179
maculata ceylonensis, Axis, 179
maculatus, Hyelaphus, 179
maculatus, Moschus, 312
malaccensis, Cervus, 151
Malayan sambar, 150
malchis, Alces, 52

major, Axis, 179
Manchurian roe, 231
Manchurian sika, 114
Manchurian wapiti, 102
manchuricus, Capreolus, 231
Mandarin sika, 121
mandarinus, Cervus, 121
mantchurica, Pseudaxis, 114
mantchuricus, Cervus, 114
mantchuricus, Elaphoceros, 114
mantschuricus, Axis, 114
mantschuricus major, Cervus, 121
Maral, 79

maral asiatica, Cervus, 104
maral, Cervus, 75

marianna, Rusa, 157
mariannus, Cervus, 157
marianus, Hippelaphus, 157
Marsh-deer, 283

matheroni, Capreolus, 232
mauricus, Cervus, 127
Mazama, 20, 243, 248, 298
mediterraneus, Cervus, 74
Megaceros, 125

megaceros, Cervus, 134
megaceros ruff, Cervus, 136
Melanaxis, 142

melas, Cervus, 204

melas, Prox, 204
mephistopheles, Pudua, 308
Mesopotamian fallow deer, 132
mesopotamica, Dama, 132
mesopotamicus, Cervus, 132
Mexican deer, 261
mexicana, Mazama, 261
mexicana, Reduncina, 262
mexicanus, Cariacus, 261
mexicanus, Cervus, 261
michianus, Elaphodus, 214
michianus, Lophotragus, 214
Michie’s tufted deer, 214
Micromeryx, 215

micrurus, Cervulus, 209
Milou deer, 233

minor, Axis, 179

minor, Cervus, 176
Miocene muntjacs, 215
Moluccan rusa, 166
moluccensis, Cervus, 166

| moluccensis, Hippelaphus, 166

328

moluccensis, Rusa, 166
Moose, 52

moschatus, Cervulus, 203
moschatus, Prox, 204
moschatus, Stylocerus, 204
moschiferus, Moschus, 311
Moschine, 21, 309
Moschus, 21, 310
Mule-deer, 269

muntjac, Cervulus, 203
muntjac, Cervus, 203
muntjac, Prox, 204
Muntjacs, 202

muntjac, Stylocerus, 204
Muntjacus, 202

Musks, 310

muswa, Alces, 52

naMBI, Nanelaphus, 305
nana, Mazama, 305
Nanelaphus, 298, 307
nanus, Cariacus, 305
nanus, Cervus, 305

nanus, Coassus, 305

nanus, Subulo, 305
nariyanus, Cervus, 91
nemoralis, Cariacus, 264
nemoralis, Cervus, 264
nemoralis, Mazama, 264
nemoralis, Reduncina, 264
nemorivaga, Mazama, 303
nemorivagus, Cariacus, 304
nemorivagus, Coassus, 303
nemorivagus, Doryceros, 304
nemorivagus, Cervus, 303
nemorivagus, Passalites, 303
nemorivagus, Subulo, 303
nepalensis, Rusa, 144
neschersensis, Cervus, 232
Newfoundland reindeer, 45
niger, Cervus, 144

niger, Hippelaphus, 144
nigricans, Cervus, 158
nigricans, Ussa, 158

North African red deer, 74
nudipalpebra, Cervus, 179

OCCIDENTALIS, Cervus, 101
occidentalis, Elaphus, 1o1
occidentalis, Strongyloceros, 101
Ollen, 77

oryzus, Axis, 176

osceola, Cariacus, 259

osceola, Dorcelaphus, 259
Otclaphus, 248

Ozotoceros, 282

PALMOMERYX, 215
palaoplatensis, Cervus, 291
Palmatus, 125

palmatus, Alces, 52

Index

| paludosa, Mazama, 283
paludosus, Blastoceros, 283
paludosus, Cariacus, 283
paludosus, Cervus, 283
paludosus, Dorcelaphus, 283
palustris, Cariacus, 283
palustris, Cervus, 283
Pampas deer, 287

Panolia, 188

Para, 175

Paraceros, 282

paradoxa, Rusa, 157
pardinensis, Cervus, 122
Passalites, 298

Pekin sika, 117

pennanti, Axis, 151

Pére David’s deer, 236
peronii, Cervus, 170
peronii, Hippelaphus, 170
peronii, Rusa, 170
perrieri, Cervus, 122

Peruvian brocket, 305
Peruvian deer, 267
Peruvian guemal, 294
peruvianus, Cariacus, 267
peruvianus, Coassus, 267
Philippine spotted deer, 172
philippinus, Cervus, 157
philippinus, Rusa, 157
philippinus, Stylocerus, 157
Pigmy brocket, 305
platycercus, Panolia, 200
Platyceros, 125

platyceros, Cervus, 127
platyceros, Dama, 127, 200
platyceros, Panolia, 200
Pliocene roe, 232

Pliocene sika, 122
Plistocene guemal, 298
Polycladus, 238
polycladus, Cervus, 239
porcinus, Axis, 175
porcinus, Cervus, 175
porcinus, Hyelaphus, 175
primigenius, Cervus, 65
priscus, Cervus, 65
Propaleomeryx, 215

Prox, 202

Pseudaxis, 18, 110
pseudaxis, .\xis, 123
pseudaxis, Cervus, Ug 23
pseudaxis, Sikelaphus, 123
Pseudocervus, 62

Pudu, 307

Pudua, 20, 307

pudu, Capra, 307

pudu, Cervus, 308

pudu, Nanelaphus, 308
pudu, Ovis, 307

pudu, Pudua, 307

pumilio, Axis, 175

pumilio, Cervus, 175

punctulata, Reduncina, 278

punctulatus, Cariacus, 278

punctulatus, Otelaphus, 278

pusilla, Mazama, 281

pusillus, Eucervus, 281

pygargus, Capreolus, 227

pygargus, Cervus, 227

pygargus mantschuricus, Cervus,
231

ramosus, Cervus, 239
Rangifer, 18, 33

rangifer, T'arandus, 37
ratva, Prox, 204

ratwa, Cervus, 204
ratwak, Stylocerus, 204
Recurvus, 188

Red brocket, 300

Red deer, 62, 64
Reduncina, 248

reevesi, Cervulus, 208
reevesii, Cervus, 208
Reindeer, 33, 37
richardsonii, Cervus, 278
richardsonii, Otelaphus, 278
Roes, 222

roosevelti, Cervus, 101
Rucervus, 18, 188

rufa, Mazama, 300

ruff, Cervus, 136

rufinus, Cariacus, 302, 303
rufinus, Cervus, 302
rufinus, Coassus, 302
rufinus, Subulo, 302

rufus, Cariacus, 300

rufus, Cervus, 300

rufus, Coassus, 300

rufus, Subulo, 300

Rusa, 18, 141, 162
Rusine group, 141
rusoides, Cervus, 123
russa, Cervus, 162

russa moluccensis, Cervus, 166
russa timoriensis, Cervus, 170

SAMBAR, 144

sartorli, Cervus, 303

sartoriil, Mazama, 303
saturatus, Moschus, 311
Savanna deer, 266
savannarum, Cariacus, 266
savannarum, Cervus, 266
savannarum, Reduncina, 266
savini, Cervus, 131
Scandinavian reindeer, 38
scheuchzeri, Paleomeryx, 217
schomburgki, Cervus, 193
schomburgkii, Rucervus, 193
Schomburgk’s deer, 193
schottini, Cervus, 37

sclateri, Cervulus, 208
sedgwicki, Anoglochis, 240
sedgwicki, Cervus, 241
sedgwickii, Eucladoceros, 240
Sedgwick’s deer, 240
senelitica, Mazama, 298
seneliticus, Cariacus, 298
seneliticus, Cervus, 298
Shou, 88

Siamese thameng, 200
Siberian roe, 227

sibiricus, Moschus, 311
sica, Cervus, III

sica manchuricus, Cervus, 114
sica typicus, Cervus, 112
sifanicus, Moschus, 315
Sika, 110

sika, Axis, 112

sika, Cervus, III

sika, Elaphoceros, 112

sika, Pseudaxis, 112
Sikelaphus, 110

Sikine group, 110

similis, Cervus, 281

similis, Mazama, 281
similis, Reduncina, 281
simplicicornis, Cariacus, 304
simplicicornis, Cervus, 303
simplicicornis, Coassus, 304
simplicicornis major, Cervus, 305
simplicicornis, Subulo, 303
sivalensis, Cervus, 201
Siwalik swamp-deer, 201
smithi, Cervus, 196
Sonoran deer, 261

spelaus, Strongyloceros, 75
spinosa, Mazama, 280
spinosus, Cariacus, 280
spinosus, Cervus, 280
Spitzbergen reindeer, 41
steerli, Cervus, 159
Streak-eyed brocket, 301
Strongyloceros, 62
strongyloceros, Cervus, 94
Stylocerus, 202

stylocerus, Prox, 204
subcornutus, Cervulus, 204
subcornutus, Stylocerus, 204
Subulo, 298

sulcatus, Furcifer, 298
superciliaris, Cariacus, 302
superciliaris, Cervus, 301
superciliaris, Coassus, 302
superciliaris, Mazama, 302

Index

superciliaris, Subulo, 302
Swamp-deer, 189
swinhoii, Cervus, 154
Szechuan sambar, 156

TaEvaNnus, Cervus, 116
taévanus, Elaphoceros, 116
taiouanus, Cervus, 116
taivanus, Axis, 116

taivanus, Pseudaxis, 116
tamulicus, Cervulus, 204
Tarandus, 33

tarandus arcticus, Cervus, 47
tarandus arcticus, Rangifer, 47
tarandus caribou, Rangifer, 42
tarandus, Cervus, 37

tarandus grcenlandicus,
|

Cervus,

/
tarandus groenlandicus, Rangifer,
7
tarandus, Rangifer, 37
tarandus spetzbergensis, Rangifer,
41
tarandus sylvestris, Cervus, 42
tarandus terre-nove, Rangifer,
45
tarandus typicus, Rangifer, 38
tema, Cariacus, 302
tema, Mazama, 302
Tenasserim muntjac, 209
terre-nove, Rangifer, 45
tetraceros, Anoglochis, 242
tetraceros, Cervus, 242
Texan deer, 261
texanus, Dorcelaphus, 261
Thameng, 195
thoroldi, Cervus, 91
Thorold’s deer, 91
Tibetan muntjac, 208
Tibetan tufted deer, 213
timorensis, Cervus, 170
Timorese rusa, 170
timoriensis, Cervus, 170
timoriensis, Hippelaphus, 170
toltecus, Cariacus, 263
toltecus, Cervus, 263
toltecus, Coassus, 263
trigonoceros, Cervus, 217
True’s deer, 267
truei, Mazama, 267
tschudii, Cervus, 305
tschudii, Doryceros, 305
tschudii, Mazama, 305
tschudii, Subulo, 3c5

THE END

ee

Tufted deer, 212
tunjac, Cervus, 162
Tuskless muntjacs, 218
Typical deer, 61

uLTRa, Antifer, 292

ultra, Cariacus, 292

ultra, Cervus, 292

ultra, Mazama, 292

unicolor, Cervus, 144

unicolor dejeani, Cervus, 156
unicolor equinus, Cervus, 150
unicolor nigricans, Cervus, 158
unicolor philippinus, Cervus, 157
unicolor, Rusa, 144

unicolor swinhoei, Cervus, 154
unicolor typicus, Cervus, 146

Ussa, 142

vaGINaLis, Cervulus, 204
vaginalis, Cervus, 203
vaginalis, Muntjacus, 204
verticornis, Cervus, 140
Virginian deer, 252
virginiana, Mazama, 249
virginiana, Reduncina, 249
virginianus, Cervus, 249
virginianus, Dorcelaphus, 249
virginianus mexicanus, Cariacus,
262
vulgaris, Capreolus, 224
vulgaris, Dama, 127

WALLICHU, Cervus, 83, 88
wallichii, Pseudocervus, 88
Wapiti, 94

wapiti, Cervus, 94
Water-deer, 218

West American wapiti, 101
Western American deer, 257
whitelyi, Cariacus, 305
whitelyi, Coassus, 305
wiegmanni, Gymnotis, 265
Wood-brocket, 303
Woodland reindeer, 42

XANTHOPYGus, Cervus, 81
Xenelaphus, 20, 293

YARCANDENsIs, Cervus, 88
Yarkand stag, 88
Yucatan deer, 263
yucatanensis, Cervus, 263

Printed by R. & R. Crarx, Limrrep, Edinburgh,

.

Poe £7 ay L

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