i«Bi»EBaa^

THE

CAMBRIDGE NATURAL HISTORY

EDITED BY

S. F. HARMER, Sc.D., F.R.S., Fellow of King's College, Cambridge ;
Superintendent of the University Museum of Zoology

A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge;
University Lecturer on the Morphology of Invertebrates

VOLUME X

By FRANK EVERS BEDDARD, M.A. (Oxon.), F.R.S., Vice-
Secretary and Prosector of the Zoological Society of
London.

Pontoon
MACMILLAN AND CO., LIMITED

NEW YORK : THE MACMILLAN COMPANY
I 9O2

All rights reserved

' ' And also as it come vnto my mynde,
Off bestis sawe I mony diuerss kynde.
The lyoun king, and his fere lyonesse,
The pantere, like vnto the smaragdyne,
The lytill sqerell, full of besynesse,
The slawe ass, the druggar-beste of pyne,
The nyce ape, the werely porpapyne,
The percyng lynx, the lufar vnicorne

The fery tigere, full of felonye,

The dromydare, the standar oliphant."

From The Kingis Quhair, JAMES I. (of Scotland).

PREFACE

INASMUCH as Sir AY. H. Flower and Mr. Lydekker could not
profess to treat the Mammalia exhaustively within the limits
of nearly 800 pages, in their Introduction to the Study of
Mammals, it is obvious that the present volume, which appears
ten years later and is of rather less size, can contain but a selec-
tion of the enormous mass of facts at the disposal of the student
of this group. Thus the chief question for myself was what to
select and what to leave aside. It will be observed that I have
reduced the pages of this book to conformity with those of
other volumes of the series by treating some groups more briefly
than others. It has appeared to me to be desirable to treat fully
such groups as the Edentata and the Marsupialia, and permissible
to be more brief in dealing with such huge Orders as those of the
Eodentia and Chiroptera. Lengthy disquisitions upon such
familiar and comparatively uninteresting animals as the Lion and
Leopard have been curtailed, and the space thus saved has been
devoted to shorter and more numerous accounts of other creatures.
As there are nearly six hundred genera of living Mammals known
to science, omission as well as compression became an absolute
necessity. I have given, I hope, adequate treatment from the
standpoint of a necessarily limited treatise to the majority of the
more important genera of Mammals both living and extinct ; but
the length of this part of the book had to be increased by the dis-
coveries, which give me at once an advantage and a disadvantage
as compared with the two authors whose names I have quoted, of
a considerable number of important new types in the last ten years.

vi PREFACE

Such forms as Notoryctes, Romerolagus, Caenolestes, " Neomylodon"
and Ocapia could not possibly have been omitted.

In preparing my accounts of both living and extinct forms
I have nearly invariably consulted the original authorities, and
have often supplemented or verified these accounts by my own
dissections at the Zoological Society's Gardens. My rule has not,
however, been invariable in this matter, inasmuch as there exist
two recent and trustworthy text-books of Mammalian Palaeontology
— Professor Zittel's Handbucli der Palaeontologie, and Dr. A. Smith
Woodward's manual, Outlines of Vertebrate Palaeontology, in the
Cambridge Biological Series. Where the name of a genus only
or its range, or merely one or two facts about it, are mentioned,
I have not thought it necessary to go further than these two
works. But a good deal has been done even since the appear-
ance of these two volumes which it will be found that I have
not ignored.

I have to thank my editors for the trouble which they have
taken in the revision of the proofs and for many suggestions. To
Professor Osborn, of Columbia University, New York, I am indebted
for some kind suggestions. My daughter Iris has assisted me
in various ways. Finally, I desire to express my indebtedness to
Mr. Dixon and to Mr. M. P. Parker for the care which they have
taken in the preparation of the figures which were drawn by
them especially for this work.

FRANK E. BEDDARD.

LONDON, February 28, 1902.

CONTENTS

PACK

PREFACE . . . . •. . . . . . . . . v

SCHEME OF THE CLASSIFICATION ADOPTED IN THIS BOOK ix

CHAPTER I

INTRODUCTORY

CHAPTER II
STRUCTURE AND PRESENT DISTRIBUTION OF THE MAMMALIA

CHAPTER III
THE POSSIBLE FORERUNNERS OF THE MAMMALIA 90

CHAPTER IV
THE DAWN OF MAMMALIAN LIFE 96

CHAPTER V
THE EXISTING ORDERS OF MAMMALS : PROTOTHERIA — MONOTREMATA . 105

CHAPTER VI
INTRODUCTION TO THE SUB-CLASS EUTHERIA 116

CHAPTER VII

EUTHERIA— MARSUPIALI A 122

30447

viii CONTENTS

CHAPTER VIII

. PAGE

EDENTATA— GANODONTA ..... ."*'** ... . .161

CHAPTER IX

UNGULATA — CONDYLARTHRA — AMBLYPODA — ANCYLOPODA — TYPOTHERIA —

TOXODONTIA — PROBOSCIDEA — HYRACOIDKA . . .*• . . . 195

CHAPTER X

UNGULATA (CONTINUED] — PERISSODACTYLA (OUD-TOED UNGULATES) — Lrro-

PTERNA 235

CHAPTER XI

UNGULATA (CONTINUED) — ARTIODACTYLA (EVEN-TOED UNGULATES) — SIRENIA 269

CHAPTER XII
CKTACEA— WHALES AND DOLPHINS . . . . . . .339

CHAPTER XIII
CARNIVORA— FISSIPEDIA 386

CHAPTER XIV
CARNIVORA (CONTINUED)— PINNIPEDIA (SEALS AND WALRUSES)— CREODONTA 446

CHAPTER XV

RODENTIA — TlLLODONTIA 458

CHAPTER XVI
INSECTIVORA — CHIROPTERA 508

CHAPTER XVII
PRIMATES .533

INDEX 591

SCHEME OF THE CLASSIFICATION ADOPTED
IN THIS BOOK

Order.

MONOTRE- |

MATA

(p. 106)
7ALLOTHERIA

(p. 96).

Sub-Class PROTOTHERIA (p. 105).

Sub-Order. Family.

{ECHIDNIDAE (p. 110).
ORNITHORHYNCHIDAE
(p. 112).

Sub-Family.

Sub-Class EUTHERIA (p. 116).

MARSUPI-
ALIA

(p. 122)

EDENTATA

(p. 161)

GANODONTA

(p. 190)

Diproto-

clontia
(p. 128)

-

Polyproto-

cloatia
(p. 149)

1

Xenarthra
(p. 166)

Nomarthra
I (P- 186)

1

MACROPODIDAE (p. 129)

PHALANGERIDAE (p. 138)

„ EPANORTHIDAE (p. 145).
DASYURIDAE (p. 149).

DlDELPHYIDAE (p. 155).

PERAMELIDAE (p. 156).

NOTORYCTIDAE (p. 158).

' MYRMECOPHAGIDAE
(p. 166).

BRADYPODIDAE (p. 170).

DASYPODIDAE (p. 173).

MYLODONTIDAE (p. 179).

MEGALONYCHIDAE (p. 183).

MEGATHERIIDAE (p. 183).

GLYPTODONTIDAE (p. 184).

ORYCTEROPODIDAE(P. 187).
\ MANIDAE (p. 188).
/ STYLINODONTIDAE (p. 191).

\_ CONORYCTIDAE (p. 193).

f Macropodinae (p. 132).
| Potoroinae (p. 137).

\Hypsiprymnodontinae-
. (p. 138).

( Phalangerinae (p. 140).
| Phascolarctinae (p. 142).
"I Phascolomyinae (p. 144).
V. Tarsipedinae (p. 145).

SCHEME OF CLASSIFICATION

Order.

UNGULATA

(p. 195)

Sub-Order.

r Condylarthra

(p. 202).
Amblypoda

(p. 205).
Ancylopoda

(p. 211).
Typotheria

(p. 212i.
Toxodontia

(p. 214).
Proboscidea

(p. 216)
Hyracoidea

(p. 232).

Perisso-
dactyla
(p. 235) .

Litopterna i
(p- 267) \

Family.

Sub-Family.

ELEPHANTIDAE (p. 217).

DlNOTHERIIDAE (p. 231).

Artiodactyla ^
(p. 269)

EQUIDAE (p. 237).

LOPHIODOXTIDAE (p. 247).

PALAEOTHERIIDAE (p. 247).
TAPIHIDAE (p. 250).
RHINOCEROTIDAE (p. 253).

TlTANOTHERIIDAE (p. 264).

MACRAUCHENIIDAE
(p. 267).

HlPPOPOTAMIDAE (p. 273).

SUIDAE (p. 275).

DlCOTYLIDAE (p. 278).

TRAGULIDAE (p. 282).
PROCERATIDAE (p. 284).
CAMELIDAE (p. 285).

CERVIDAE (p. 291)

GlRAFFIDAE (p. 301).

ANTILOCAPRIDAE (p. 306).
BOVIDAE (p. 307).
ANTHRACOTHERIIDAE

(p. 328).
CAENOTHERIIDAE (p. 329).

XlPHODONTIDAE (p. 329).

OREODONTIDAE (p. 330).
ANOPLOTHERIIDAE (p. 332).

/ Cervinae (p. 293).
\ Moschinae (p. 299).

SIRENIA (p. 333).

CETACEA

(p. 339)

Mystacoceti
(p. 353)

Odontoceti

(p. 362)

CARNIVOEA

(p. 386)

Fissipedia
(p. 387)

C BALAEXOPTERIDAE
-^ (p. 355).
I.BALAEXIDAE (p. 358).

I PHYSETERIDAE (p. 362)

-! DELPHIXIDAE (p. 372).

PLATANISTIDAE (p. 380).
1^ SQUALODOXTIDAE (p. 384).

-! ZEUGLODONTIDAE (p. 384).

FELIDAE (p. 390).
MACHAERODOXTIDAE
(p. 401).

VlVEURIDAE (p. 403)

/ Physeterinae (p. 363).
\_ Ziphiinae (p. 367).

Euplerinae (p. 403).
Galidictiinae (p. 404).
Cryptoproctinae (p. 404).
Viverrinae (p. 405).
Herpestinae (p. 409).

SCHEME OF CLASSIFICATION

XI

Order.

CARNIVORA

(Continued)

CREODONTA

(p. 455).

Sub-Order.

Fissipedia

(Continued)

Pinnipedia

(p. 446)

INSECTIVORA

Insectivora
Vera

(p. 509)

Dennoptera

I (p. 520)

.

•

Simpliciden

RODENTIA

(p. 458)

(p. 462)

TILLODONTIA

(p. 506).

Dupliciden-
tata
I (p. 502)

Family.

HYAENIDAE (p. 411).
CANIDAE (p. 413).
PROCYONIDAE (p. 426).

MUSTELIDAE (p. 431)

URSIDAE (p. 442).
OTARIIDAE (p. 450).
TRICHECHIDAE (p. 451).
PHOCIDAE (p. 452).

ANOMALURIDAE (p. 462).
SCIURIDAE (p. 463).
CASTORIDAE (p. 467).
HAPLODONTIDAE (p. 469).
GLIRIDAE (p. 470).

MURIDAE (p. 471)

BATHYERGIDAE (p. 480).
SPALACIDAE (p. 482).
GEOMYIDAE (p. 483).
HETEROMYIDAE (p. 484).
DIPODIDAE (p. 484).
PEDETIDAE (p. 486).

OCTODONTIDAE (p. 487)

CTENODACTYLIDAE (p.490).
CAVIIDAE (p. 491).
DASYPROCTIDAE (p. 493).
DINOMYIDAE (p. 495).
CHINCHILLIDAE (p. 496).
CERCOLABIDAE (p. 497).
HYSTRICIDAE (p. 499).

LEPORIDAE (p. 502).
LAGOMYIDAE (p. 505).

f ERINACEIDAE (p. 509).

TUPAIIDAE (p. 511).

CENTETIDAE (p. 511).

POTAMOGALIDAE (p. 513).
SOLENODONTIDAE (p. 513).

CHRYSOCHLORIDAE (p. 514).
MACROSCELIDAE (p. 515).
TALPIDAE (p. 516).
SORICIDAE (p. 518).

| GALEOPITHECIDAE (p. 620).

Sub-Family.

f Melinae (p. 432).
\ Mustelinae (p. 433).
[ Lutrinae (p. 439).

' Murinae (p. 471).

Phlaeomyinae (p. 473).

Hydromyinae (p. 474).

Rhynchomyinae (p. 474).

Gerbillinae (p. 475).

Otomyinae (p. 475).

Dendromyinae (p. 476).

Lophiomyinae (p. 476).

Microtinae (p. 477).

Sigmodontinae (p. 479).
. Neotominae (p. 480).

( Octodontinae (p. 487).
•{ Loncherinae (p. 488).
_ Capromyinae (p. 489).

Xll

SCHEME OF CLASSIFICATION

Order.

CHIROPTERA

(p. 521)

PRIMATES

(p. 533)

Sub-Order.
Megachiro-
ptera

(P- 524)

Microchiro-
ptera

(p. 526)

Lemuroidea

(p. 534)

Anthro-
poidea

(p. 554)

Family.

Sub-Family.

LEMURIDAE (p. 538)

-! PTEROPODIDAE (p. 524).

RHINOLOPHIDAE (p. 527).
NYCTERIDAE (p. 527).
VESPERTILIONIDAE (p. 528).
EMBALLONURIDAE (p. 530).
PHYLLOSTOMATIDAE
(p. 531).

f Indrisinae (p. 538).
I Lemurinae (p. 540).
j Galagininae (p. 542).
VLorisinae (p. 545).
CHIROMYIDAE (p. 548).
TARSIIDAE (p. 550).
ANAPTOMORPHIDAE (p. 552).
CHRIACIDAE (p. 552).
, MEGALADAPIDIDAE (p. 554).
( HAPALIDAE (p. 556).

CKBIUAE (p. 557).
4 CERCOPITHECIDAE (p. 562).

I SlMIIDAE (p. 570).
\. HOMINIDAE (p. 585)

CHAPTER I

INTRODUCTORY

THE Mammalia form a group of vertebrated aninials which
roughly correspond with what are termed in popular language
" quadrupeds," or with the still more vernacular terms of " beasts "
or " animals." The name " Mammal " is derived from the most
salient characteristic of the group, i.e. the possession of teats ;
but if the term were used in an absolutely strict etymological sense,
it could not include the Monotremes, which, though they have
mammary glands, have not fully-differentiated teats (see p. 16).
There are, however, as will be seen shortly, other characters which
necessitate the inclusion of these egg-laying quadrupeds within
the class Mammalia.

The Mammalia are unquestionably the highest of the Verte-
brata. This statement, however, though generally acceptable,
needs some explanation and justification. " Highest " implies
perfection, or, at any rate, relative perfection. It might be said
with perfect truth that a serpent is in its way an example of
perfection of structure : not incommoded with limbs it can slip
rapidly through the grass, swim like a fish, climb like a monkey,
and dart upon its prey with rapidity and accuracy. It is an
example of an extremely specialised reptile, the loss of the limbs
being the most obvious way in which it is specialised from
more generalised reptilian types. Specialisation in fact is often
synonymous with degradation, and, this being the case, implies
a restricted life. On the other hand, simplification is not always
to be read as degeneration. The lower jaw, for instance, of
mammals has fewer bones in it than that of reptiles, and is more
concisely articulated to the skull ; this implies greater efficiency

VOL. x %< B

2 LARGE SIZE OF MAMMALIA CHAP.

as a biting organ. The term highest, however, includes increased
complexity as well as simplification, the two series of modifica-
tions being interwoven to form a more efficient organism. It
cannot be doubted that the increased complexity of the brain of
mammals raises them in the scale, as does also the complex and
delicately adjusted series of bonelets which form the organ for
the transmission of sound to the internal ear. The separation of
the cavity containing the lungs, and the investment of the parti-
tion so formed with muscular fibres, renders the action of the
lungs more effective; and there are other instances among the
Mammalia of greater complexity of the various parts and organs
of the body when compared with lower forms, which help to
justify the term "highest" generally applied to these creatures.

Complexity and finish of structure are often accompanied by
large size ; and the Mammalia are, on the whole, larger than any
other Vertebrates, and also contain the most colossal species. The
huge Dinosaurs of the Mesozoic epoch, though among the largest
of animals, are exceeded by the Whales ; and the latter group
includes the mightiest creature that exists or has ever existed,
the eighty-five-feet-long Sibbald's Rorqual. Confining ourselves
rigidly to facts, and avoiding all theorising on the possible
relation between complexity and nicety of build and the capacity
for increase in bulk, it is plain from the history of more than
one group of mammals that increase in bulk accompanies specialis-
ation of structure. The huge Dinocerata when compared with
the ancestral Pantolamlda teach us this, as do many similar
examples. Within the mammalian group, as in the case of other
Vertebrates, difference of size has a certain rough correspondence
with difference of habitat. The Whales not only contain the
largest of animals, but their average size is great ; so too with
the equally aquatic Sirenia and very aquatic Pinnipedia. Here
the support offered by the water and the consequent decreased
need for muscular power to neutralise the effects of gravity
permit of an increase in bulk. Purely terrestrial animals come
next ; and finally arboreal, and, still more, " flying " mammals
are of small size, since the maintenance of the position when
moving and feeding needs enormous muscular effort.

The Mammals are more easily to be separated from the
Vertebrates lying lower in the series than any of the latter are
from each other in ascending order. A large number of char-

DEFINITION OF THE CLASS

acters might he used in addition to those which will be made
use of in the following brief catalogue of essential mammalian
features, were it not for the low-placed Monotremata on the one
hand and the highly specialised Whales on the other. Including
those forms, the Mammalia are to be distinguished from all other
Vertebrates by the following series of structural features, which
will be expanded later into a short disquisition upon the general
structure of the Mammalia. The class Mammalia may, in fact, be
thus denned : —

Hair-clad Vertebrates, with cutaneous glands in the female,
secreting milk for the nourishment of the young. Skull without
prefrontal, postfrontal, quadrato-jugal, and some other bones, and
with two occipital condyles formed entirely by the exoccipitals.
Lower jaw composed of dentary bone only, articulating only with
the squamosal. Ear bones a chain of three or four separate bonelets.
< Vrvical vertebrae sharply distinguished from the dorsals, and if
with free ribs, showing no transition between these and the
thoracic ribs. Brain with four optic lobes. Lungs and heart
separated from abdominal cavity 1 >y a muscular diaphragm. Heart
with a single left aortic arch. Red blood-corpuscles non-nucleate.

The following characters are also very nearly universal, and
in any case absolutely distinctive : — Cervical vertebrae, seven ;
vertebrae with epiphyses. Ankle-joint " cruro-tarsal," i.e. be-
tween the leg and the ankle, and not in the middle of the ankle.1
Attachment of the pelvis to the vertebral column pre-acetabular
in position.

The Mammalia since they are hot-blooded creatures are more
independent of temperature than reptiles ; they are thus found
spread over a wider area of the earth's surface. As however, though
hot-blooded, they hare not the powers of locomotion possessed by
birds, they are not quite so widely distributed as are those
animals. The Mammalia range up into the extreme north, but,
excepting only forms mainly aquatic, such as the Sea Lions, are not
known to occur on the Antarctic continent. With the exception
of the flying Bats, indigenous mammals are totally absent from
New Zealand ; and it seems to be doubtful whether those sup-
posed oceanic islands which have a mammalian fauna are really

1 The degeneration of the hind-limh in "Whales and Sirenia forbids the use of
this character as a distinctive one on the principles advocated by the selection of
the above list. But it would be absurd to leave out hair.

4 NUMBER OF SPECIES CHAP, i

oceanic in origin. The continents and oceans are peopled by
rather over three thousand species of Mammalia, a number which
is considerably less than that of either birds or reptiles. It
seems clear that, so far at any rate as concerns the numbers of
families and genera, the mammalian fauna of to-day is less varied
than it was during the Mid - tertiary period, the heyday of
mammalian life. It is rather remarkable to contrast in this way
the mammals and the birds. The two classes of the animal
kingdom seem to have come into being at about the same period ;
but the birds either have reached their culminating point to-day,
or have not yet reached it. The Mammalia, on the other hand,
multiplied to an extraordinary extent during the Eocene and the
Miocene periods, and have since dwindled. The break is most
marked at the close of the Pleistocene, and may be in part due
to the direct influence of man. At present man exercises so
enormous an effect, both directly and indirectly, that the future
history of the Mammalia is probably foreshadowed by the in-
stances of the White Rhinoceros and the Quagga. On the other
hand, the economic usefulness of the Mammalia is greater than
that of any other animals ; and the next most important era
in their history will be probably that of domesticity and " pre-
servation."

CHAPTER II

STRUCTURE AND PRESENT DISTRIBUTION OF THE MAMMALIA

External Form. — It would be quite impossible for any one to
confuse any other quadrupedal animal with a mammal. The
body of a reptile is, as it were, slung between its limbs, like the
body of an eighteenth century chariot between its four wheels ;
in the mammal the body is raised entirely above, and is
supported by, the four limbs. The axes of these limbs too, as a
general rule, are parallel with the vertical axis of the body of
their possessor. There is thus a greater perfection of the
relations of the limbs to the trunk from the point of view of a
terrestrial creature, which has to use those limbs for rapid move-
ment. The same perfection in these relations is to be seen, it
should be observed, in such running forms among the lower
Vertebrata as the Birds and the Dinosaurs, where the actual
angulation ' of the limbs is as in the purely running Mam-
malia. These relations are of course absolutely lost in the
aquatic Cetacea, and not marked in various burrowing creatures.
The way in which the fore- and hind-limbs are angulated is
considerably different in the two cases. In the latter, which
are most used and, as it were, push on the anterior part of
the body, the femur has its lower end directed forwards, the
tibia and the fibula project backwards at the lower end, while
the ankle and foot are again inclined in the same direction as
the femur. With the fore - limbs there is not this regular
alternation. The humerus is directed backwards, the fore-arm
forwards, and the hand still more forwards. This angulation
seems to facilitate movement, inasmuch as it is seen in even the
Amphibia and the lower Eeptiles, in which, however, the differ-
ences between the fore- and hind*-limbs are less marked, indicat-

INTELLECT

ing therefore a less specialised condition of the limbs. It is
an interesting fact that the angulation of the limbs is to some
extent obliterated in very bulky creatures, and almost entirely
so in the elephants (see p. 217), which seem to need strong and
straight pillars for the due support of their huge bodies.

The alertness and general intellectual superiority of mammals
to all animals lying below them in the series (with the exception
of the birds, which are in their way almost on a level with the
Mammalia) are seen by their active and continuous movements.
The lengthy periods of absolute inotionlessness, so familiar to
everybody in such a creature as the Crocodile, are unknown among
the more typical Mammalia except indeed during sleep. This
mental condition is clearly shown by the proportionate develop-
ment of the external parts of all the organs of the higher senses.
The Mammalia as a rule have well-developed, often extremely
large, flaps of skin surrounding the entrance to the organ of
hearing, often called " ears," but better termed " pinnae." These
are provided with special muscles, and can be often moved and
in many directions. The nose is always, or nearly always, very
conspicuous by its naked character ; by the large surface, often
moist, which surrounds the nostrils ; and again by the muscles,
which enable this tract of the integument to be moved at will.
The eyes, perhaps, are less marked in their predominance over the
eyes of lower Vertebrates than are the ears and nose ; but they
are provided as a rule with upper and lower eyelids, as well as
by a nictitating membrane as in lower Vertebrates. The apparent
predominance of the senses of smell and hearing over that of
sight appears to be marked in the Mammalia, and may account
for their diversity of voice as well as of odour, and for the
general sameness of coloration which distinguishes this group
from the brilliantly-coloured birds and reptiles. The head, too,
which bears these organs of special sense, is more obviously
marked out from the neck and body than is the case with the
duller creatures 'occupying the lower branches of the Vertebrate
stem.

The Hair. — The Mammalia are absolutely distinguished from
all other Vertebrates (or, for the matter of that, Invertebrates)
by the possession of hair. To define a mammal as a Vertebrate
with hair would be an entirely exclusive definition ; even in the
smooth Whales a few hairs at least are present, which may be

HAIR

reduced to as few as two bristles on the lips. The term " hair,"
however, is apt to be somewhat loosely applied ; it has been
made use of to describe, for example, the slender processes of the

&fi

Sit1

FIG. 1. — A, Section of human skin. Co,
Dermis ; 7), sebaceous glands ; /', fat
in dermis ; 6, vessels in dermis ; GP,
vascular papillae ; H, hair ; N, nerves
in dermis ; NP, nervous papillae ; Sc,
horny layer of epidermis ; SD, sweat
j;land ; SD\ duct of sweat gland : SM,
Malpighian layer. B, Longitudinal sec-
tion through a hair (diagrammatic). Ajt,
Band of muscular fibres inserted into tlie
hair-follicle ; C'o, corium (dermis) ; F,
external longitudinal ; fl, internal cir-
cular, fibrous layer of follicle ; Ft, fatty
tissue in the dermis ; Gff, hyaline
membrane between the root-sheath and
the follicle ; HBh, sebaceous gland ;
HP, hair-papilla with vessels in its in-
terior ; M, medullary substance (pith)
of the hair ; 0, cuticle of root-sheath :
H. cortical layer ; .Sc, horny layer of
epidermis ; Sch, Hair shaft ; SM, Mal-
pighian layer of epidermis ; ]\'S, H'.S1,
'outer and inner layers of root-sheath.
Anatomy.)

( From Wieuersheim's Comparative

chitinous skin of the Crustacea. It will be necessary, therefore,
to enter into the microscopical structure and development of the
mammalian hair. Hair is found in every mammal. The first
appearance of a hair is a slight thickening of the stratum
Malpighii of the epidermis, the cells taking part in this being

DEVELOPMENT OF HAIR

elongated and converging slightly above and below. Dr. Maurer
has called attention to the remarkable likeness between the
embryonic hair when at this stage and the simple sense-organs
of lower Vertebrates. Later there is formed below this a
denser aggregation of the corium, which 'ultimately becomes

_ the papilla of the hair.

This is the apparent
homologue of the first
formed part of a
feather, which projects
as a papilla before the
epidermis has under-
gone any modification.
Hence there is from
the very first a differ-
ence between feathers
and hairs — a difference
which must be care-
fully borne in mind,
especially when we
consider the strong
superficial resem-
blance between hairs
and the simple barb-

FiG. 2. — Four diagrams of stages in the development \QQQ feathers. Still
of a hair. A, Earliest stage in one of those mammals

in which the dermal papilla appears first ; B, C, D, later the knob ot epl-

three stages in the development of the hair in the (Jgj-mic cells becomes
human embryo, bib, Hair-bulb ; crn, horny layer

of the epidermis; foil, hair - follicle ; grm, hair- depressed into a tubu-

germ; h, hair in D, projecting on the sur- k structure, which is
face ; ?«MC, Malpighian layer of epidermis ; pp,

dermal papilla ; seb, developing sebaceous glands ; lined with cells also

sh.l, sh.2, inner and outer root-sheaths. (After j • j n .-,

Hertwig.) derived from the

stratum Malpighii, but

is filled with a continuation of the more superficial cells of the
epidermis. This is the hair- follicle, and from the epidermic cells
arises the hair by direct metamorphosis of those cells ; there is
no excretion of the hair by the cells, but the cells become the
hair. From the hair-follicle also grows out a pair of sebaceous
glands, which serve to keep the fully-formed hair moist.

Dr. Meijerle1 has lately described in some detail the parti-

1 " ITber die Haare der Saugethiere, " Morph. Jahrb. xxi. 1894, p. 312.

pp

HAIR AND UNDER-FUR

cular arrangement of the individual hairs among mammals :
they are not by any manner of means scattered without order,
but show a definite and regular arrangement, which varies with
the animal. For instance, in an American Monkey {Midas}, the
hairs arise in threes — three hairs of equal size springing from the
epidermis close together ; in the Paca (Coelogenys) there are in
each group three stout hairs alternating with three slender hairs.
In some forms a number of hairs spring from a common point :
in the Jerboa (Dipus) twelve or thirteen arise from a single hole ;
in UTSHS arctos there is the same general plan, but there is one
stout hair and four or five slender ones. There are numerous
other complications and modifications, but the facts, although
interesting, do not appear to throw any light upon the mutual
affinities of the animals. Allied forms may have a very different
arrangement, while in forms which have no near relationship the
plan may be very similar, as is shown by the examples cited from
Dr. Meijerle's paper. The groups of hairs, moreover, have them-
selves a definite placing, which the same anatomist has compared
with the disposition of the bundles of hairs behind and between
the scales of the Armadillo, and which has led him to the view
that the ancestors of mammals were scaly creatures — a view also
supported by Professor Max Weber,1 and not in itself unreason-
able when we consider the numerous points of affinity between
the primitive Mammalia and certain extinct forms of reptiles.2

The hairs are greatly modified in form in different mammals
and in different parts of their bodies. It is very commonly the
case that a soft under-fur can be distinguished from the longer and
coarser hairs, which to some extent hide the latter. Thus the
" sealskin " of commerce is the under-fur of the Otaria ursina of
the North. The coarser hairs may be further differentiated into
bristles ; these again into spines, such as those of the Hedgehog
and of the Porcupine. Again, the flattening and agglutination
of hairs seems to be responsible for the scales of the Man Is

1 " Bemerkungen iiber den Ursprung der Haare," Anat. Anz. 1893, p. 413.

- See for this matter, p. 90. Dr. Bonavia has recently advanced (Stiidics in
Evolution, London, 1895) the somewhat fantastic view that the pigment-patches
of Carnivorous and other mammals are a reminiscence of an earlier scaly condition.
There is no direct evidence that the primitive mammals were scaly, nor are the
Monotremata or Marsupials furnished with any more traces of such a con-
dition than are other mammals : and they are the most lowly organised of existing
Mammalia.

10 CARPAL SENSE ORGAN CHAP.

and for the horns of the Rhinoceros. It is a matter of common
knowledge that upon the head of various animals, e.g. the
Domestic Cat, long and sensitive hairs are developed, which are
connected with the terminations of nerves, and perform a sensory,
probably tactile function. These occur on the snout, above the
eyes, and in the neighbourhood of the ears. It is an interesting
fact that a tuft of quite similar hairs occurs on the hand of
many mammals close to the wrist, which, at least in the
case of Bassaricyon, are connected with a strong branch from the
arm-nerve. These tufts also occur in Lemurs, in the Cat, various
Rodents and Marsupials, and are probably quite general in
mammals who " feel " with their fore-limbs ; — in which, in fact,
the fore -limbs are not exclusively running organs. That the last
remaining hairs of the Cetacea are found upon the muzzle, is
perhaps significant of the importance of these sensory bristles.
The entire absence of hairs is quite common in this order,
although traces of them are sometimes found in the embryo.
The Sirenia, too, are comparatively hairless, as are also many
Ungulates. Whether the presence of blubber in the former case
and the existence of a very thick skin in the latter animals are
facts which have had anything to do with the disappearance of
hair or not, is a matter for further inquiry.

The intimate structure of the hair varies considerably. The
variations concern the form of the hair, which may be round in
transverse section, or so oval as to appear quite Hat when the
hair is examined in its entirety. The substance of the hair is
made up of a central medulla or pith with a peripheral cortex ;
the latter is scaled, and the scales are often imbricated and
with prominent edges. The amount of the two constituents also
differs, and the cortex may be reduced to a series of bands
surrounding only tracts of the enclosed pith. In the hair is
contained the pigment to which the colour of mammals is
chietiy due. Tracts of brightly-coloured skin may exist, as in
the Apes of certain genera ; but such structures are not general.
The pigment of the hair seems to consist of those pigmentary
substances known as melanins. It is remarkable to find such an
uniform cause of coloration, when we consider the great variety
of feather-pigments found in birds. The variations of colour
of the hair of mammals are due to the unequal distribution of
these brown pigments. There are very few mammals which can

POLORATION i i

be called brightly coloured. The Bats of the genus Kerivoula.
have been compared to large butterflies, and some of the Flying
Squirrels have strongly-marked contrasts of reddish brown, white,
and yellow. The same may be said of the spines of certain
Porcupines. But we find in the hair no bright blues, greens, and
reds such as are common among birds.

There are certain general facts about the coloration of
mammals which require some notice here. Xext to the
usually sombre hues of these animals the general absence
of secondary sexual coloration is noteworthy. In but a few
cases among the Lemurs and Bats do we find any marked
divergences in hues between males and females. Secondary
sexual characters in mammals are, it is true, often exhibited
by the great length of certain hair-tracts in the male, such as
the mane of the Lion, the throat- and leg-tufts of the Bar-
bary Sheep, and so forth ; but apart from these, the secondary
sexual characters of mammals are chiefly shown in size, c.y. the
Gorilla, or in the presence of tusks, e.y. various Boars, or of horns, as
in the Deer, etc. The coloration of mammals frequently exhibits
conspicuous patterns of marking. These are in the form of
longitudinal stripes, of cross-stripes, or of spots ; the latter may
be " solid " spots, or broken up, as in the Leopard and Jaguar,
into groups of smaller spots arranged in a rosette-fashion. We
never find in mammals the complicated " eyes " and other mark-
ings which occur in so many birds and in other lower Verte-
brates. It is important to note that in the Mammalia whose
sense of sight is quite keen there should be a practical absence
of secondary sexual colours. As to the relationship of the various
forms of marking that do occur, it seems clear that there has
been a progression from a striped or spotted condition to uniform
coloration. For we find that many Deer have spotted young ;
that the young Tapir of the New World is spotted, while its
parents are uniform blackish brown ; the strongly-marked spot-
ting of the young Puina contrasts with the uniform brown of the
adult ; and the Lion cub, as every one knows, is also spotted, the
adult lioness showing considerable traces of the spots.

The seasonal change in the colours of certain mammals is a
subject upon which much has been written. The extreme of this
is seen in those creatures, such as the Polar Hare and the Arctic
Fox. which become entirely blanched in the winter, recovering

12 CIVET AND MUSK CHAP.

their darker coat in the spring. This is, however, only an
extreme case of a change which is general. Most animals get
a thicker fur in winter and exchange it for a lighter one in
summer. And the hues of the coat change in correspondence.

Glands of the Skin. — The great variety of integumental
glands possessed by the Mammalia distinguishes them from any
group of lower Vertebrates. This variability, however, only con-
cerns the anatomical structure of the glands in question. Histo-
logically they are all of them apparently to be referred to one of
two types, the sudoriparous or sweat gland and the sebaceous
gland. Simple sweat and sebaceous glands are abundant in
mammals, with but a few exceptions. The structures that we are
now concerned with are agglomerations of these glands. The
mammary glands will be treated of in connexion with the mar-
supium ; they are either masses of sweat glands, or of sebaceous
glands whose secretion has been converted into milk.

Many Carnivora possess glands opening to the exterior, near
the anus, by a large orifice. These secrete various odoriferous
substances, of which the well-known " civet " is an example.
Other odoriferous glands are the musk glands of the Musk-deer
and of the Beaver ; the suborbital gland of many Antelopes ; the
dorsal gland of the Peccary, which has given the name of
Dicotyles to the genus on account of its resemblance in form to
a navel. This gland may be seen to secrete a clear watery fluid.
The Elephant has a gland situated on the temple, which is said
to secrete during certain periods only, and to be a warning to
leave the animal alone. Very remarkable are the foot glands of
certain species of Rhinoceros ; they are not universally present
in those animals, and are therefore useful as specific distinctions.
On the back of the root of the tail in many Dogs are similar
glands. The Gentle Lemur (Hapalemur) has a peculiar gland
upon the arm, about the size of an almond, which in the male
underlies a patch of spiny outgrowths. In Lemur varius is a
hard patch of black skin which may be the remnants of such
a gland. It is thought that the callosities on the legs of Horses
and Asses are remnants of glands.

One of the most complex of these structures which has been
examined microscopically exists in the Marsupial Myrmeco^ius}
On the skin of the anterior part of the .chest, just in front of the
1 Proc. Zool. Soc. 1887, p. 527.

GLANDS OF THE SKIN 13

sternum, is a naked patch of skin which is seen to be perforated
by numerous pores. Besides the ordinary sebaceous and sweat
glands there are a series of masses of glands, opening by larger
orifices, which present the appearance of groups of sebaceous
glands, and are of a racemose character ; but the existence of muscu-
lar fibres in their coats seems to show that they should be referred
rather to the sudoriparous series. Beneath the integument is a
large compound tubular gland quite half an inch in diameter.

In Didelpliys dimidiata there is a precisely similar glandular
area and large underlying gland, the correspondence being re-
markable in two Marsupials so distant in geographical position
and affinities. Even among the Diprotodont genera there is
something of the kind ; for in Dorcopsis luctuosa and D. muelleri
is a collection of four unusually large sebaceous follicles upon
the throat, and in the Tree Kangaroo (Dendrolagus bennettii}
there is the same collection of enlarged hair-follicles, though
they are apparently somewhat reduced as compared with those of
Dorcopsis. These are of course a few examples out of many.

It seems to be possible that the functions of these various
glands is at least twofold. In the first place, they may serve,
where predominant in one sex, to attract the sexes together.
In the second place, the glands may be useful to enable a strayed
animal of a gregaxious species to regain the herd. It is perfectly
conceivable too that in other cases the glands may be a protec-
tion, as they most undoubtedly are in the Skunk, from attacks.
In connexion with the first, and more especially the second, of
the possible uses of these glands, it is interesting to note that
in purely terrestrial creatures, such as the Ehinoceros, the glands
are situated on the feet, and would therefore taint the grass and
herbage as the animal passed, and thus leave a track for the
benefit of its mate. The same may be said of the rudimentary
glands of Horses if they are really glands. The secretion of the
" crumen " of Antelopes is sometimes deposited deliberately by
Oreotragus upon surrounding objects, a proceeding which would
attain the same end. One may even perhaps detect " mimicry "
in the similar odours of certain animals. Prey may be lured to
their destruction, or enemies frightened away. The defenceless
Musk-deer may escape its foes by the suggestion of the musky
odour of a crocodile. It is at any rate perfectly conceivable
that the variety of odours among mammals may play a very

14 HOOFS, NAILS AND CLAWS . CHAP.

important part in their life, and it is perhaps worthy of note
that birds with highly-variegated plumage are provided only
with the uropygial gland, while mammals with usually dull and
similar coloration have a great variety of skin glands. Scent
is no doubt a sense of higher importance in mammals than in
birds.* The subject is one which will bear further study.

Nails and Claws. — Except for the Cetacea (where rudi-
ments have been found in the foetus), the extremities of the
fingers and of the toes of mammals are covered by, or encased in,
horny epidermic plates, known as nails, claws, and hoofs.

The variety in the shape and development of these corneous
sheaths to the digits is highly characteristic of mammals as
opposed to lower Vertebrates. If we take extreme cases, such as
the nail of the thumb in Man, the hoof of a Horse, and the claw of
a Cat, it is easy to distinguish the three kinds of phalangeal horny
coverings. But the differences become extinguished as we pass
from these to related types. The nail of the little finger in Man
approaches the claw -like form ; and the hoofs of the Lama are
almost claws in the sharpness of their extremities. On the
whole it may be said that claws and hoofs embrace the bone
which they cover, while nails lie only upon its dorsal surface.
The form of the distal phalanx which bears the nail shows,
however, two kinds of modification which do not support such a
classification. When those phalanges are clad with hoofs or
covered by a nail they end in a rounded and flattened termina-
tion. On the other hand, when they bear a claw they are them-
selves sharpened at the extremity and often grooved above.

The Marsupium. — It " may appear to be unnecessary at this
juncture to speak of the marsupial pouch, which is so usually
believed to be a characteristic of the group Marsupialia. Eudi-
ments of this structure have, however, been recently discovered
in the higher mammals, and, as Dr. Klaatsch l has remarked, all
researches into the " history of the mammals culminate in the
question whether the placental mammals pass through a mar-
supial stage or not." We cannot, therefore, look upon the
marsupial pouch as a matter affecting only the Marsupials,
though it is true that this organ is at present functional only in
them and in the Monotremata.

1 "tJber Marsupialrudirnente bei Placentaliern," Morph. Jahrb. xx. 1893, p.
276.

POUCH

Iii the Marsupials the pouch shelters the young, which are
born in an exceedingly imperfect state, minute, nude, and blind,
with a " larval " mouth fitted only to grasp in a permanent
fashion the teat, upon which they are carefully fixed by the
parent. But even later the pouch is made use of as a temporary
harbour of refuge : from the pouch of female Kangaroos at the
Zoological Gardens may frequently be observed to protrude the tail

g.m.

FJ<;. -j. — Echidna hystrix. A, Lower surface of brooding female ; B, dissection showing
a dorsal view of the pouch and mammary glands ; tt, the two tufts of hair in the
lateral folds of the mammary pouch from which tlie secretion flow s. b.m, Pouch ;
d, cloaca ; g.m, groups of mammary glands. (From Wiedersheim's Compared ice
Anatomy, alter W. Haacke.)

and hind-legs of a young Kangaroo as big as a Cat, and perfectly
well able to take care of itself.

In the Moiiotremata (in Echidna} there is a deep fold of
the skin which lodges the unhatched egg, and into which the
mammary glands -open, one on either side. This structure" is only
periodically developed, and arises from two rudiments, one corre-
sponding to each mammary area ; but in the female with eggs or
young there is but a single deep depression, which occupies the
same region of the body as the marsupial pouch of the Mar-

1 6 MARSUPIUM AND MAMMARY POUCH CHAP.

supials.1 It is usually held that this structure is not of pre-
cisely the same morphological value as the pouch of the
Marsupial ; and the difference is expressed by terming the one
(that of Echidna} the mammary pouch, and the other the
marsupium. At first sight it may appear to be an unnecessary
refinement to separate two structures which have so many and
such obvious likenesses. It is not quite certain, however, that the
difference is not even more profound than later opinions seem
to indicate. The Monotremata not only have no teats, as has
already been pointed out, but the mammary glands themselves
are of a perfectly different nature to those of the higher mammals,
including the Marsupials. There is therefore no a priori
objection to the view that the accessory parts developed in con-
nexion with the mammary glands should also be different. The
teat of the higher Mammalia grows up round the area upon
which the ducts of the mammary glands open ; it is a fold of
skin which eventually assumes the cylindrical form of the adult
teat, and which includes the ducts of the milk glands. It has
been suggested that the two folds of skin which form the
mammary pouch of Echidna are to be looked upon as the equi-
valent of the commencing teat of the higher mammal.2 In this
case it is clear that the marsupial folds of the Marsupial cannot
correspond accurately with the apparently similar folds of
Echidna, because there are teats as well. It is the teats which
correspond to the marsupial folds of Echidna. This view is in
apparent contradiction to an interesting discovery in a specimen
of a Phalanger by Dr. Klaatsch.3 This Marsupial, ' like most
others, has a well - developed marsupial pouch, in which the
young are lodged at birth ; but round two of the teats is
another distinct fold on either side, the outer wall of which
forms the general wall of the pouch. Dr. Klaatsch thinks
that these smaller and included pouches are the equivalents of
the mammary pouches of Echidna. . They contain teats, but this
comparison does not do away with the validity of Gegenbaur's
suggestion already referred to, because the teats are (see above)

1 See Haacke, "On the Marsupial Ovum, the Mammary Pouch, etc., of the
Echidna," Proc. Roy. Soc. 1885, p. 72 ; and " Uberdie Entstehungder Saugetiere,"
Biol. Centralbl. viii. 1889, p. 8.

2 See Gegenbaur's Elements of Comp. Anat. Transl. by Bell, 1878, p. 421.

3 " tiber die Beziehungen zwischen Mammartasche u. Marsupium," Morph.
Jahrb. xvii. 1891, p. 483.

POUCH OF MARSUPIALS

secondary. If this fact be fairly to be interpreted in the sense
which Dr. Klaatsch attaches to it, we have an interesting case
of the growth of a new organ out of and partly replacing an
old organ. In the Monotremes there is a pouch which facilitates
or performs both nutritive and protective functions ; in the
Phalanger these two functions are carried on in separate
pouches ; finally, in other Marsupials, there is a return to the
^differentiated state of affairs found in the Monotremata, but
with the help of a new
organ not found in them.

Though so character-
istic, of Marsupials, the
marsupial pouch is not
always developed in them.
It is present in all the
Kangaroos, Wallabies, and
Wombats, in fact in the
Diprotodonts. It is also
present in a number of
the carnivorous Polypro-
todont Marsupials ; but in
Pliascologale it is only pre-
sent in rudiment, and in
Myrmecobius it is entirely
obsolete. In the American
Opossums the state of the
pouch is variable. " Gener-
ally absent, sometimes
merely composed of two
lateral folds of skin separ-
ate at each end, rarely complete," is Mr. Thomas' summary in his
definition of the family Didelphyidae.1 Another curious feature
of the pouch in the Marsupials is the variability in the position
of the mouth of the pouch : in all the Diprotodonts it looks
forward ; but in many Polyprotodonts it looks backward. This,
however, has some connexion with the habitual attitude of the
possessor : in the Kangaroo, leaping along on its hind-legs, it is
requisite that the pouch should open forwards; but in the
dog-like Thylacine, going on all fours, the fact that the pouch

1 Catalogue of Marsupials in British Museum, 1886.
VOL. X C

FIG. 4. — Diagram of the development of the nipple
(in vertical sectioii). A, Indifferent stage, gland-
ular area flat ; B, elevation of the glandular area
with the nipple ; C, elevation of the periphery
of the glandular area into the false teat, a,
Periphery of the glandular area ; b, glandular
area ; glt glands. (From Gegenbaur.)

1 8 VESTIGIAL POUCHES CHAP.

opens backwards is less disadvantageous to the contained
young.

The male Thylacine has a pouch which is quite or very
nearly as well formed as in the female. There are also rudi-
ments of a pouch in the male foetuses of many Marsupials,
especially of those belonging to the Polyprotodont section of the
order, though these rudiments are by no means confined to that
subdivision. Up to so late a period as the age of four months
(length 19 '8 cm.) the male Dasyurus ur sinus has a pouch.

We have now to consider the interesting series of facts
relative to the permanence — in a rudimentary condition it is
true — of the mammary pouch in the higher Mammalia, facts
which seem to be an additional proof that they have been
derived from an ancestor in which the pouch was an organ of
functional importance. The first definite proof of the occurrence
of a pouch in any mammal not a Marsupial or a Monotreme was
made by Malkmus, who found this structure in a Sheep. It
seems, however, that the structures found in the higher mammals
are not always comparable to the marsupiurn of the Marsupials,
but sometimes to the mammary pouch of the Monotreme. That
the Marsupials are a side line, and not involved in the ancestry
of the Eutheria, is an opinion which is at present widely held. At
the same time it is reasonable to suppose that the original stock
lying between the Prototheria and the Metatheria, whence the
latter and the Eutheria have arisen, preserved both the mammary
pouch of the lower mammal and the marsupiurn of the further-
developed stage, as docs Phalangista occasionally at the present
day. Hence to find remnants of both structures in existing
mammals would not be incredible. This is what Dr. Klaatsch
believes to be the case. In certain Ungulates, including two
species of Antelope, Dr. Klaatsch found very considerable rudi-
ments of folds provided with unstriated muscular fibre ; there
were in the adult Cervicapra isabellina a pair of pouches, one on
each side, and a rudiment of a second on either side ; possibly
this multiplication of the pouches has relation to the number of
young. That there is more than one pouch makes a comparison
with the mammary pouch rather than with the marsupium
probable. The Ungulate teat, it must be remembered (see p. 16),
is a secondary teat ; hence there is no difficulty in the com-
parison from this point of view. A pouch containing a primary

ii LEMURS AND THEIR YOUNG 19

teat would of course be absolutely incomparable with a mammary
pouch, because in that case the wall of the teat itself would be
the pouch.

Mammals belonging to quite different Orders show traces
more or less marked of a marsupiuni. In young Dogs the teats
are borne upon an area where the skin is thinner, the covering
of hair less dense than elsewhere — all points of resemblance to
the inside of the pouch of a Marsupial ; in addition to this
there are traces of the sphincter marsupii muscle. In other
Carnivora there are similar vestiges. In Lemur catta a more
complete rudiment of a marsupial pouch is to be met with. In
this Lemur the teats are both inguinal and pectoral; the skin
in these regions is thin and but slightly hairy, and extends
forwards as two bands of the same thinness and smoothness on
each side of the densely hairy skin covering the sternum. This
area is sharply separated from the rest of the integument by a
fold which runs parallel to the longitudinal axis of the body,
and can be comparable with nothing save the rudiment of the
marsupial fold.

One is tempted to wonder how far the habit which certain
Lemurs have of carrying their young across the abdomen with
the tail wrapped round the body of the mother is a reminiscence
of a marsupial pouch.

Skeleton.

The skeleton of the Mammalia consists almost solely of the
endoskeleton. It is only among the Edentata that an exo-
skeleton of bony plates in the skin is met with. As in other
Vertebrates, the skeleton is divisible into an axial portion,
the skull and vertebral column, and an appendicular skeleton,
that of the limbs. The bones of mammals are well ossified,
and in the adult there are but few and small tracts of cartilage
left.

Vertebral Column. — The vertebral column of the mammals,
like that of the higher Yertebrata, consists of a number of
separate and fully-ossified vertebrae.

The constitution of a vertebra upon which all the usual
processes are marked is as follows : — There is first of all the
body or centrum of the vertebra, a massive piece of bone shaped
like a disc or a cylinder. The centra of contiguous vertebrae

20

VERTEBRAE

are separated by a certain amount of fibrous tissue forming
the intervertebral disc, and the apposed surfaces of the centra
are as a rule nearly flat. In this last feature, and in the
important fact that the centra are ossified from three distinct
centres, the anterior and posterior pieces (" epiphyses ") remaining
distinct for a time, even for a long time (as in the Whales),
the centra in the mammals differ from those of reptiles and
birds. The epiphyses are not found throughout the vertebral
column of the lowly-organised Monotremata, and they do not
appear to exist in the Sirenia.

Fio. 5. — Anterior surface of
Human thoracic vertebra
(fourth), x §. az, Anterior
zygapophysis ; c, body or
centrum ; I, lamina, and
p, pedicle, of the neural
arch ; nc, neural canal ; t,
transverse process. (From
Flower's Osteology of the
Mammalia.)

FIG. 6. — Side view of first
lumbar vertebra of Dog
(Canis famttiaris). x $.
a, Anapophysis ; az, an-
terior zygapophysis ; w,
metapophysis ; pz, pos-
terior zygapophysis ; *•,
spinous process ; t, trans-
verse process. (From
Flower's Osteology.)

From each side of the centrum on the dorsal side arises a
process of bone which meets its fellow in the middle line above,
and is from there often prolonged into a spine. A canal is thus
formed which lodges the spinal cord. This arch of bone is
known as the neural arch, and the dorsal process of the same as
the spinous process. The sides of the neural arch bear oval
facets, by which successive vertebrae articulate with one another :
those situated anteriorly are the anterior zygapophyses, while
those on the posterior aspect of the arch are the posterior
zygapophyses ; these articular facets do not exist in the tail-
region of many mammals, e.g. Whales.

In addition to the dorsal median spinous process of the

21

vertebra there may be a ventral median process, arising of course
from the centrum, termed the hypapophysis.

From the sides of the neural arch, or from the centrum
itself, there is commonly a longer or shorter process on each
side, known as the transverse process. This is sometimes
formed of two distinct processes, one above the other ; in
such cases the upper part is called a diapophysis, the lower
.a parapophysis.

The neural arch may also bear other lateral processes, of
which one directed forwards is the metapophysis, the other
directed backwards the anapophysis.

The series of bones which constitute the vertebral column
can be divided into regions. It is possible to recognise cervical,
dorsal, lumbar, sacral, and caudal vertebrae. In the case of
animals with only rudimentary hind-limbs, such as the Whales,
there is no recognisable sacral region. The neck or cervical
vertebrae are nearly always seven in number. The well-known
exceptions are the Manatee, where there are six, and certain
Sloths, where there are six, eight, or nine. These rare exceptions
only accentuate the very remarkable constancy in number, which
is very distinctive of the mammals as compared with lower
Yertebrata. There are of course abnormalities, the last cervical,
and sometimes the last two, assuming the characters of the
ensuing dorsals, by developing a more or less complete rib.
There are also recorded examples of Bradypus, in which the
number of cervicals is increased to ten. The characteristics,
then, of the cervical vertebrae are, in the first place, that they
do not normally bear free ribs, and that there is a break as a rule
between the last cervical and the first dorsal on this account. In
birds, for example, the cervicals, differing in number in different
families and genera, gradually approach the dorsals by the
gradually lengthening ribs. The transverse processes of the
vertebrae are commonly perforated by a canal for the vertebral
artery, and are bifid at their extremities. In some Ungulates
these vertebrae, moreover, approximate to the vertebrae of lower
Vertebrata in the fact that there are ball and socket joints
between the centra, instead of only the fibrous discs of the
remaining vertebrae. •

The first two vertebrae of the series are always very
different from those which follow. The first is termed the

22

ATLAS

atlas, and articulates with the skull. The most remarkable
fact about this bone (shared, however, by lower Vertebrates)
is that its centrum is detached from it and attached to the
next vertebra, in connexion with which it will be referred

FIG. 7. — Hum.in atlas (young), showing de-
velopment, x f . as, Articular surface for
occiput ; g, groove for first spinal nerve
and vertebral artery ; i «, inferior arch ;
t, transverse process. (From Flower's
Osteology.)

FIG. 8. — Inferior surface of atlas of
Dog. x ^. sn, Foramen for first
spinal nerve ; r, vertebrarterial
canal. (From Flower's Osteology.)

FIG. 9. — Atlas of Kangaroo.

Parker and Haswell's Zoology.)

to immediately. The whole bone thus gets a ring-like form,
and the salient processes of other vertebrae are but little de-
veloped, with the exception of
the transverse processes, which
are wide and wing - like. In
many Marsupials, such as the
Wombat and Kangaroo, the arch
of the atlas is open below, there
(From being no centre of ossification.
In others, such as Thylacinus,
there is a distinct nodule of bone in this situation not con-
crescent with the rest of the arch.

The second vertebra, which is known as the axis or epi-
stropheus, is a compound structure, the anterior " odontoid process,"
which fits into the ring of the atlas, being in reality the
detached centrum of that vertebra.1 It is a curious fact about
that process that it has independently become spoon-shaped in
two divisions of Ungulates ; that it has become so seems to be
shown by the fact that in the earlier types of both it has the
simple peg-like form, which is the prevailing form. The cervical

1 Its independence from the epistropheus is emphasised in Monotremes and
some Marsupials by its late fusion with that vertebra.

AXIS

vertebrae are occasionally wholly (Eight "Whales) or partially
(many Whales, Jerboa, certain Edentates) welded into a com-

FIG. 10. — Side view of axis of Dog.
x |. o, Odontoid process ; pz,
posterior zygapophysis ; s, spin-
ous process ; t, transverse pro-
cess ; r, vertebrarterial canal.
(From Flower's Osteology.)

FIG. 11. — Anterior surface of axis
of Red Deer, x f . o, Odon-
toid process ; pz, posterior
zygapophysis ; sn, foramen for
second spinal nerve. (From
Flower's Osteology.)

bined mass. Indications of this have even been recorded in the
human subject.

The dorsal vertebrae vary greatly in number : nine (Hyper-
oodon) seems to be the lowest number existing normally ; while
there may be as many as nineteen, as in Centetes, or twenty-two,
as in Hyrax. These vertebrae are to be defined by the fact that
they carry ribs, and the first one or two lumbars are often
" converted into " dorsals by the appearance of a small super-
numerary rib. The spinous processes of these vertebrae are
commonly long, and sometimes very long. It is only among the
Glyptodons that any of these vertebrae are fused together into a
mass.

The lumbar vertebrae, which follow the dorsal, vary greatly
in number. There are as few as two in the whale Neobcdaena.,
as many as seventeen in Tursiops ; this group, the Cetacea,
contains the extremes. Nine lumbars are found in the Lemurs
Indris and Loris. As a rule the number of lumbars is to some
extent dependent upon that of the dorsals. It often happens that
the number of thoraco-lumbar vertebrae is constant for a given
group. Thus the Artiodactyles have nineteen of these vertebrae,
and the Perissodactyles as a rule twenty -three. A greater
number of dorsals implies a smaller number of lumbars, and of
course vice versa. The existence of a sacral region formed of a

SACRUM AND CAUDAL SERIES

number of vertebrae fused together and supported by the pelvic
girdle is characteristic of the mammals, but is not found in the
Cetacea and the Sirenia, where functional hind-limbs are wanting.
Strictly speaking, the sacrum is limited to the two or three
vertebrae whose expanded transverse processes meet the ilia.
But to these are or may be added a variable number of vertebrae
withdrawn from both the lumbar and the caudal series, which

pub

FIG. 12. — Lepus cuniculus. Innomin-
ate bones and sacrum, ventral aspect.
acet, Acetabulum ; il, ilium ; isch,
ischium ; obt, obturator foramen ;
pub, pubis ; sacr, sacrum ; sy, sym-
physis. (From Parker and Haswell's
Zoology.)

FIG. 13. — Anterior surface of fourth
caudal vertebra of Porpoise (Pho-
coena communis). x \. h, Chevron
bone ; m, metapophysis ; s, spin-
ous process ; t, transverse process.
(From Flower's Osteology.)

unite with each other to form the massive piece of bone which
constitutes the sacrum of the adult.

The caudal vertebrae complete the series. They begin in as
fully developed a condition as the lumbars, with well-marked
transverse processes, etc. ; but they end as no more than
centra, from which sometimes tiny outgrowths represent in
a rudimentary way the neural arches, etc. Very often the
caudal vertebrae are furnished with ventral, generally V-shaped,
appendages, the chevron bones or intercentra.1 These are

1 Intercentra are but rarely met with anterior to the caudal series. Mr. Parsons
has, however, recorded their occurrence in the lumbar vertebrae of Atherura.

SKULL OF MAMMALS AND REPTILES

particularly conspicuous in the Whales and in the Edentates. In
the former group the occurrence of the first interceiitrum serves
to mark the separation of the caudal from the lumbar series.
The number of caudals varies from three in Man — and those
quite rudimentary — to nearly fifty in Manis macrura and Micro-
gale longicaudata.

The Skull. — The skull in the Mammalia differs from that
of the lower Vertebrata in a number of important features, which
will be enumerated in the following brief sketch of its structure.

F

Jni

—Sq.ccc.

4-C.OCC.

Fia. 14. — Lateral view of skull of a Dog. O.occ, Occipital condyle ; F, frontal ;
F.inf, infra-orbital foramen ; Jg, jugal ; Jin, premaxilla ; L, lachrymal ; M,
maxilla ; Maud, external auditory meatus ; Md, mandible ; N, nasal ; P,
parietal ; Pal, palatine ; Pjt, process of squamosal ; Pt, pterygoid ; Sph, ali-
sphenoid ; Sq, squamosal ; Sq.ccc, supraoccipital ; T, tympanic. (From Wieder-
sheim's Comparative Anatomy.)

In the first place, the skull is a more consolidated whole than in
reptiles ; the number of elements entering into its formation is
less, and they are on the whole more firmly welded together
than in Vertebrates standing below the Mammalia in the series.
Thus in the cranial region the post- and pre-frontals, the post-
orbitals and the supra-orbitals have disappeared, though now and
again we are reminded of their occurrence in the ancestors of
the Mammalia by a separate ossification corresponding to some
of the bones. Nowhere is this consolidation seen with greater
clearness than in the lower jaw. That bone, or rather each
half of it, is in mammals formed of one bone, the dentary (to
which occasionally, as it appears, a separate mento-Meckelian

V

26 LOWER JAW CHAP.

ossification may be added). The angular, splenial, and all the
other elements of the reptilian jaw have vanished, though the
numerous points from which the mammalian dentary ossifies
is a reminiscence of a former state of affairs ; and here again
an occasional continuance of the separation is preserved, as the
case observed by Professor Albrecht of a separate supra-angular
bone in a Eorqual attests. Among other reptilian bones that are
not to be found in the mammalian skull are the basipterygoids,
quadrato-jugal, and supra temporal. A few of these bones,
however, though no longer traceable in the adult skull save in
cases of what we term abnormalities, do find their representatives
in the foetal skull. Professor Parker, for example, has described
a supra-orbital in the embryo Hedgehog ; a supratemporal also
appears to be occasionally independent.

In the mode of the articulation of the lower jaw to the skull
the Mammalia apparently, perhaps really, differ from other
Vertebrates. In the Amphibia and Eeptilia, with which groups
alone any comparisons are profitable, the lower jaw articulates
by means of a quadrate bone, which may be movably or firmly
attached to the skull. In the mammals the articulation of the
lower jaw is with the squamosal. The nature of this articulation
is one of the most debated points in comparative anatomy.
Seeing that Professor Kingsley l in the most recent contribution
to the subject quotes no less than fifty-two different views, many
of which are more or less convergent, it will be obvious that in a
work like the present the matter cannot be treated exhaustively.
As, however, Professor Kingsley justly says that " no single bone
occupies a more important position in the discussion of the
origin of the Mammalia than does the quadrate," and with equal
justice adds that " upon the answer given as to its fate in this
group depends, in large measure, the broader problem of the
phylogeny of the Mammalia," it becomes, or indeed has long been,
a matter which cannot be ignored in any work dealing with the
mammals. A simple view, due to the late Dr. Baur and to
Professor Dollo, commends itself at first sight as meeting the
case. The last-named author holds, or held, that in all the
higher Vertebrates it is at least on a priori grounds likely
that two such characteristically vertebrate features as the
lower jaw and the chain of bones bringing the outer world
1 Tufts College Studies, No. 6, 1900.

OSSICULA AUDITUS

into communication with the internal organ of hearing would
be homologous throughout the series. He believed, therefore,
that the entire chain of ossicula auditus in the mammal is
equal to the columella of the reptile, since their relations
are the same to the tympanum on the one hand and to

.in i.

FIG. 15. — Head of a Human embryo of the fourth mouth. Dissected to show the
auditory ossicles, tympanic ring, and Meckel's cartilage, with the by old and thyroid
apparatus. All these parts are delineated on a larger scale than the rest of the
skull, an, Tympanic riug ; b.hy, basihyal element ; hy, so-called hyoid bone ; in,
incus ; md, bony mandible ; ml, malleus ; st, stapes ; tp, tympanum ; tr, trachea ;
1. (ink), first skeletal (mandibular) arch (Meckel's cartilage) ; II. second skeletal
(hyoid) arch ; III. third (first branchial) arch ; IV. V. fourth and fifth arches
(thyroid cartilage). (From Wiedersheim's Structure of Man.)

the foramen ovale on the other ; and that the lower jaw
articulates in the same way in both. It follows, therefore, that
the glenoid part of the squamosal must be the quadrate which
has become ankylosed with it after the fashion of concentration
in the mammalian skull that has already been referred to. The
fact that occasionally the glenoid part of the squamosal is a
separate bone1 appeared to confirm this way of looking at the

1 Cf. the Armadillo Peltephilus, p. 186.

28 QUADRATE AND INCUS CHAP.

matter. But the hall-mark of truth is not always simplicity ;
indeed the converse appears to be frequently the case. And
on the whole this view does not commend itself to zoologists
at present. For it must be borne in mind that the lower jaw of
the mammal is not the precise equivalent of that of the reptiles.
Apart from the membrane bones, which may be collectively the
equivalents of the dentary of the mammal, there is the cartilaginous
articular bone to be considered, which forms the connexion
between the rest of the jaw and the quadrate in reptiles. Even
in the Anomodontia, whose relations to the. Mammalia are con-
sidered elsewhere, there is this bone. But in these reptiles the
articular bone articulates not only with the quadrate, but also to
a large extent with the squamosal, the quadrate shrinking in
size and developing processes which give to it very much the
look of either the incus or the malleus of the mammalian ear.
In fact it seems on the whole to fit in with the views of the
majority, as well as with a fair interpretation of the facts of
embryology, to consider that the chain of ear bones in the
mammal is not the equivalent of the columella of the reptile,
but that the stapes of the mammal is the columella, and that
the articulare is represented by the malleus and the quadrate
by the incus. It is very interesting to note this entire change
of function in the bones in question. Bones which in the reptile
serve as a means of attachment of the lower jaw to the skull are
used in the mammal to convey the waves of sound from the
tympanum of the ear to the internal organ of hearing.

Another important and diagnostic feature in the mammalian
skull is that the first vertebra of the vertebral column always
articulates with two separate occipital condyles, which are borne
by the exoccipital bones and formed mainly though not entirely
by them. Certain Anomodontia form the nearest approach to
the mammals in this particular. The two condyles of Amphibia
are purely exoccipital in origin.

In the Mammalia, unlike what is found in lower Vertebrates
(but here again the Anomodontia form at least a partial exception),
the jugal arch does not connect the face with the quadrate, for,
as already said, that bone does not exist, in the Sauropsidan
form, in mammals. This arch passes from the squamosal to the
maxillary, and has but one separate bone in addition to those
two, viz. the jugal or malar.

QUADRATE AND TYMPANIC RING

In connexion with the elaboration of the chain of auditory
ossicles' it is very usual for
mammals to possess a thin
inflated bone, sometimes
partly or entirely formed
out of the tympanic bone,
and known as the tympanic
bulla. Whether this struc-
ture is thin and inflated
or thick and depressed in
form it is characteristic of Ma
the mammals, arid does not
occur below them in the
series. But it is not pre-
sent in all mammals. It
is absent, for example, in
the Monotremes. When
it is present it is some-
times formed from other
bones, as, for instance, from
the alisphenoids. The tym-
panic ring has been held to
be the equivalent of the
quadrate. It is more prob- T

FIG. 16. — Under surface ot the cranium ot a Dog.
ably the quadrato-jugal.1 x ^. apf, Anterior palatine foramen ; as, pos-

Ribs. — All mammals
are furnished with ribs, of
which the number of pairs
differs considerably from
group to group, or it may
be even from species to
species. The ribs are
attached as a rule by two
heads, of which one, the
capitulum, arises as a rule
between two centra of
successive vertebrae. The
other, the tuberculum,
springs from the transverse process. Only in the Monotremes

1 Gegenbaur, Vergl. Anat. Wirbelth. Leipzig, 1898, p. 404.

terior opening of alisphenoid canal ; AS,
alisphenoid ; £0, basioccipital ; £8, basi-
sphenoid ; cf, coudylar foramen ; earn, ex-
ternal auditory meatus ; Ex.0, exoccipital ;
fim, foramen lacerum medium ; flp, foramen
lacerum posterius ; fm, foramen magnum ;
fo, foramen ovale ; fr, foramen rotund ura ;
Fr, frontal ; gf, ^rlenoid fossa ; yp, post-
gleuoid process ; Ma, malar ; MX, maxilla ;
oc, occipital condyle ; op, optic foramen ;
Per, mastoid portion of periotic ; pgf, post-
glenoid fossa ; PI, palatine ; PMx, pre-
maxilla ; pp, paroccipital process ; ppf, pos-
terior palatine foramen ; PS, presphenoid ;
Pt, pterygoid ; sf, sphenoidal fissure or fora-
men lacerum anterius ; sm, stylomastoid fora-
men ; SO, supraoceipital ; Sq, zygomatic
process of squamosal ; Ty, tympanic bulla;
Vo, vomer. (From Flower's Osteology.)

RIBS

are there ribs with but one, the capitular, head. In the
posterior part of the series the two heads often gradually
coalesce, so that there comes to be but one, the capitular,
head. The Whales also, at least the Whalebone Whales, are
exceptional in possessing but one head to the ribs, which
is the capitular. The first rib joins the sternum below, and
a variable number after this have the same attachment. There
are always a number of ribs, sometimes called floating ribs,
which have no sternal attachment. In the Whalebone Whales

1

FIG. 17. — A, First thoracic skeletal segment for comparison with B, fifth cervical vertebra
(Man), b.v, Body of vertebra ; c, first thoracic rib ; c', cervical rib (which lias be-
come united with the transverse process, tr), the two enclosing the costo-transverse
foramen (f.c.t) ; st, sternum ; zy, articular process of the arch (zygapophysis).
(From Wiedersheim's Structure of Man.)

it is the first rib alone which is so attached. As a rule,
to which the Whales mentioned are again an exception, the
rib is divided into at least two regions — the vertebral portion
which is always ossified, and the sternal moiety which is usually
cartilaginous. This is, however, often very short in the first rib.
They are, however, ossified in the Armadillos and in some other
animals. Between the vertebral and sternal portions an inter-
mediate tract is separated off and ossified in the Monotremata.
The ribs of existing mammals belong only to the dorsal region
of the vertebral column, but there are traces of lumbar ribs and
also of cervical ribs. In the Monotremata, indeed, these latter

RIBS AND STERNUM

are persistently free for a very long period, and in some cases
never become ankylosed with their vertebrae. But it should be
noted that in this group there is no approximation to the state
of affairs which exists in many lower Vertebrates, where there is
a gradual transition between tha ribs of the cervical and those of
the dorsal region of the vertebral column ; for that of the seventh
ribs in Monotremes is smaller than those which precede it.

The Sternum. — All the Mammalia so far as is known possess
a sternum. This is the bone,
or series of bones (sternebrae),
which lies upon the ventral
surface of the chest, and to
which the ribs are attached
below. The development of
the sternum has been shown
to take place from the fusion of
the ribs below into two lateral
bands, one on each side ; the
approximation of these bands
forms the single and unpaired
.sternum of most mammals.
Very considerable traces, how-
ever, of the paired state of the
sternal bones often exist ; thus
in the Sperm Whale the first
piece of the sternum is divided
into two by a longitudinal FlG- 18— Ste™u1m/a"d; sternal ribs.of *J,e

* ° Common Mole (Tatpa europaea), with

division, and the second piece the clavicles (cl) and humeri (//) ; M,

is longitudinally grooved. The manubrium sterni Nat. size. (From

• ° Flower s Osteology.)

development of the sternum

out of the fused ends of ribs is shown in a more com-
plete condition in some specis of Manis than in many other
mammals. Thus in M. tricuspis the last ribs of those which are
attached to the sternum are completely fused together into a
single piece on each side.1 As a general rule the last ribs
which come into relation with the sternum do so only in an
imperfect way, being simply firmly attached at their sides to,
but not fused with, the last ribs which are definitely articulated
with the sternum. Contrary to what is found in lower Verte-

1 Killer's Zool. Miscellen. i. 1894.

COMPOSITION OF STERNUM

brates, the sternum of the Mammalia consists of a series of pieces,
as many as eight or nine or even sixteen in Choloepus, of which
the first is called the manubrium sterni, and the last the ensifonn
cartilage, xiphisternum, or xiphoid process. The latter often
remains largely cartilaginous throughout life ; in fact this is
generally but not universally the case with that part of the
breastbone. The most extraordinary modification of the xiphoid
process is seen in the African species of the genus Manis, where
it diverges into two long cartilages, which run
back to the pelvis and then, curving round, run
forwards and fuse together in the middle line
anteriorly. These processes serve for the attach-
ment of certain tongue-muscles. They were looked
upon by Professor Parker as the equivalents of
the " abdominal ribs " of reptiles elsewhere non-
existent among mammals. This view is not,
however, usually held. The manubrium sterni
is often keeled in the middle line below ; this
is so with the Bats, which thus approach the
birds, and probably for the same reason, i.e. the
need of an enlarged origin for the pectoral muscle,
which is concerned in the movements of flight.
In many forms this part of the sternum is much
FIG. 1 9.— sternum broader than the pieces which follow ; this is so

of the Pig (Sus . . T _ _.

scrofa). x £. ms, with the Viscacha. In the rig the precise reverse
Mesostemum; js seen the manubrium being narrower than the

ps, presternum ; °

o», xiphisternum. rest of the sternal bonelets. It will be noticed,
(From Flower s however. that in this and similar cases there are

Osteology.) '

no clavicles. Eibs are attached between the
successive pieces of the sternum. When the sternum is
reduced, as it is in the Cetacea and in the Sirenia, it is the
intermediate part of the series of bones which becomes abbrevi-
ated or vanishes. The Sperm Whale has only a manubrium
sterni and a following piece belonging to the mesosternum.
It is fair to say that the xiphoid process and the rest of
the sternum have disappeared, since among the Toothed Whales
a progressive shortening of the sternum can be seen. In
the Whalebone Whales the sternum is still further reduced ;
the manubrium is alone left, and to it are attached but
a single pair of ribs. In Balaena, however, a rudimentary

INTERCLAVICLE

33

piece, apparently comparable to a xiphoid process, has been
detected.

FIG. 20.— Steruum of Rudolphi's Whale (Bal-
aenoptera borealis), showing its relation to
the inferior extremities of the first pair of
ribs. x Jy. (From Flower's Osteology. )

FIG. 21. — Sternum of a young
Dugong (Halicore indicus).
x J. From a specimen in
the Leydeu Museum, ps,
Presternum ; xs, xiphister-
niim. (From Flower's Oste-
ology).

From the instances which have been described, as well as
from the mode of development of the sternum and from the
number of free ribs, i.e. ribs which are not attached to it, it
would seem that the sternum has undergone a considerable reduc-
tion in its size. This reduction may be possibly accounted for
by the need for respiratory activity, which is clearly increased by
a less-marked fixity of the walls of the thoracic cavity. In the
case of the Whales one can hardly help coming to that conclusion.
The arrangement in the Monotrernata does not, however, point in
the same direction ; for these animals are precisely like the
higher Mammalia in the reduction of the sternum and of the
number of ribs which reach it.

The Episternum. — The Mammalia are as a rule to be dis-
tinguished from lower Vertebrates by the absence of an episternum,
or interclavicle as it is also called. In the Monotremata, however,
there is a large T-shaped bone which does not overlie the ster-
num as in reptiles, but is anterior to it. The relations of this
bone to the clavicles seem to leave no doubt that it is the
equivalent of the Lacertilian interclavicle or episternum. The
Monotremata are not, however, the only mammals in which this
structure is to be seen. The Mole in the embryonic condition is

VOL. x D

34

VESTIGIAL EPISTERNA

CHAP.

provided with pieces of bone which overlie the manubrium sterni

Fia. 22. — Shoulder girdle of Ornithorhynchus. c1, c'-, c3, First, second, third ribs ; d,
clavicle ; e.c, epicoracoid ; es'andes", interclavicle (episternum) ; m.c, metacoracoid ;
m.s, mauubrium sterrii ; sc, scapula ; st, sternebra. (From Wiedersheim's Structure
of Man.)

and are attached to the clavicles, and are no doubt to be regarded

as the same structure. Probably in many
mammals the manubrium will be found
to be partly made up of corresponding
rudiments. In any case, vestiges of an
episternum in the shape of two minute
ossicles have been discovered in Man,
lying in front of the manubrium. They
have been termed ossa suprasternalia.
In Man and in the Mole the paired
..r.c. nature of the episternum is clearly
apparent. It has been suggested that
this structure in its entirety belongs to

FIG. 23.— Episternum of an em- the clavicles, just as the sternum belongs
?c°iS; fy%*Sg to the ribs ; *.* that it formed out of the
tion of the episternum ; es", approximated and fused ends of the

lateral portion of the same; i • i T-VH/T-J.!^ j i

r.c, costal ribs ; st, sternum, clavicles. Dr. Mivart figured a good
(The figure was constructed many years since a pair of ossicles in

from two consecutive hori- ,.- , .

zontai sections.) (From Mycetes, lying in one case between the
Wiedersheim's structure of en(js of the clavicles and the manubrium

Man.) . .

sterni, and in another example anterior

to the ventral ends of the clavicles. Gegenbaur has figured a
1 Proc. Zool. Soc. 1865, p. 567.

EFISTERXUM OF MAN

35

pair of similar bones in the Hamster.1 It is possible that these
are to be referred to the same category. It has also been

FIG. 24.— Episternal
vestiges in Man.
cl, Clavicle, sawn
through ; es, "epi-
sternum " (sterno-
clavicular cartilage) ;
l\ interclavicular
ligament ; /", costo-
clavicular ligament ;
m.s, mannbrium
sterni ; o.s, ossa
suprastemalia ; r.c,
first rib ; st, ster-
num. (From Wie-
dersheim's Structure
of Man.)

suggested that these supposed episternal rudiments are the
vestiges of a pair of cervical ribs.

The Pectoral Girdle. — The skeleton by which the fore-limb
is connected with the trunk is known as the Pectoral Girdle.
The main part of this girdle is formed by the large scapula, or
blade-bone as it is often termed. The coracoidal elements will be
dealt with later. The scapula is not firmly connected with the
backbone ; it is attached merely by muscles, thus presenting a
great difference from the corresponding pelvic girdle. The reason
for this difference is not easy to understand. On the one hand
it may be pointed out that in all running animals at any rate
there is a greater need for the fixation in a particularly firm way
of the hind-limbs ; but, again, in the climbing creatures both
limbs would, one might suppose, be bettered by a firm fixation. It
must be remembered, however, that in the latter case the same
result is at least partly brought about by a well-developed clavicle,
which fixes the girdle to the sternum and so to the vertebral
column by means of the ribs.

Broadly speaking, too, the fore-limbs require a greater freedom
and variety of movement than the hind-limbs, which are supports
1 Veryl. Anat. der WirbeWi. Leipzig, 1898, p. 497.

SHOULDER BLADE

for or serve to push along the rapidly-moving body. Stronger
fixation is therefore a greater necessity posteriorly than anteriorly.
In any case, whatever the explanation, this important difference
exists.

The shoulder-blade of mammals is as a rule a much-flattened
bone with a ridge on the outer surface known as the spine ;

FIG. 25. — Right scapula of
Dog (Cams familiaris).
x £. a, Acromion ; «/,
prescapular fossa ; c,
coracoid ; d>, coracoid or
anterior border ; ess, in-
dicates the position of
the coraco-scapular su-
ture, obliterated in adult
animals by the complete
aiikylosis of the two
bones ; gb, glenoid or
posterior border ; gc,
gleuoid cavity ; pf, post-
scapular fossa ; s, spine ;
ss, suprascapiilar border.
(From Flower's Oste-
ology.)

FIG. 26. — Right scapula of
Red Deer (Ccruus elaphns).
x J. a, Acromion ; af, an-
terior or prescapular fossa ;
c, coracoid ; gc, glenoid
cavity ; pf, postscapular
fossa ; ss, partially ossi-
fied suprascapular border.
(From Flower's Osteology.)

this ridge ends in a freely -projecting process, the acromion,
from which a branch often arises known as the metacromion.
This gives a bifurcate appearance to the end of the ridge. The
spine is less developed and the scapula is narrower in such
animals as the Dog and the Deer which simply run, and whose
fore-limbs therefore are not endowed with the complexity of
movement seen, for instance, in the Apes.

It has been pointed out that the area which lies in front of

SHOULDER BLADE AND LOCOMOTION

37

the spine, the prescapular lamina, is most extensively developed in
such animals as per-
form complex move-
ments with the
Sea

scapula of Dolphin (Tursiops tursio).
x £. a, Acromion ; of, prescapular fossa ; c, cora-
coid ; gc, glenoid cavity ; pf, postscapular fossa.
(From Flower's Osteology.)

fore-
limbs. The Sea Lion
and the Great Anteater
are cited by Professor
G. B. Howes as ex-
amples of this pre-
ponderance of the an-
terior portion of the
scapula over that which
lies behind the spine.
The general shape of
the scapula varies con-
siderably among the
different orders of mammals ; but it always presents the characters

mentioned, which are nowhere seen
among the Sauropsida except among
certain Anomodonts, which will be
duly referred to (see p. 90). The
most conspicuous divergences from
the normal are to be found in the
Cetacea and the Monotremata. In
the former the acromion is approxi-
mated so nearly to the anterior
border of the blade-bone that the
prescapular fossa is reduced to a
very small area ; and in Platanista
the acromion actually coincides
. with the anterior border, so that

FIG. 28. — Side view of right half of

shoulder girdle of a young Echidna that fossa actually disappears. In

the Whales, too, the scapula is as
rule very broad, especially above ;
has frequently a fan-like contour.

•pf

(Echidna hystrix). x f . a, Acro-
mion ; c, coracoid ; cb, coracoid

border ; d, clavicle ; ess, coraco- a rule very broad, especially above ;

scapular suture ; ec, epicoracoid ;
gb, glenoid border ; gc,

In the Monotremata the acromion

glenoid border ; gc, glenoid
cavity ; ic, interclavicle ; pf, post-
scapular fossa ; ps, presternum ; s,
spine ; ss, suprascapular epiphysis ;
ssf, subscapular fossa. (From
Flower's Osteology.}

also coincides with the anterior
border of the scapula ; but the
sameness of appearance which it
thus presents (in this feature) to the Cetacean scapula is appar-

CLAVICLE

ently not due to real resemblance. What has happened in the
Monotremata is, that the prescapular fossa is so enormously
expanded that it occupies the whole of the inner side of the
blade-bone, while the subscapular fossa which, so to speak, should
occupy that situation, has been thus pushed round to the front,
where it is divided from the postscapular fossa by a slight
ridge only.

The clavicle is a bone which varies much in mammals. It is
sometimes indeed, as in the Ungulata, entirely absent ; in other
forms it shows varying degrees of retrocession in importance ; it is
only in climbing, burrowing, digging, and flying mammals that
it is really well developed.

In the higher Mammalia the coracoid1 is present, but does
not reach the sternum as in the Monotremata. It is known to

human anatomists as

-^y^**""^ cl ^^^^^^. "

^3BW^ ° the coracoid process

of the scapula. It
has been found, how-
ever, by Professor
Howes2 and others,
that this process
really consists of
two separate centres
of ossification, form-
ing two separate
bonelets, which in
the adult become
firmly ankylosed to
each other and to
the scapula. These
two separate bones have been met with in the embryo of Lepus,
Sciurus, and the young of various other mammals belonging to very
diverse orders, such as Edentates and Primates. The separation even
occasionally persists in the adult. The question is, What is the
relation of these bonelets to the coracoid of the Monotremata and
to the corresponding regions of reptiles ? Professor Howes terms the
lower patch of bone the nietacoracoid and the upper the epicoracoid ;

1 To this category are perhaps to be referred cartilaginous pieces occurring in
the Rabbit, Mus and Sorcx (see Fig. 29 above).

2 "On the Coracoid of the Terrestrial Vertebrates," P.Z.S. 1893, p. 585.

FIG. 29. — Shoulder girdle, with upper end of sternum (inner
surface) of Shrew (Sorex), after Parker, x 7. a, Acro-
mion ; c, coracoid ; cl, clavicle ; ec, partially ossified
" epicoracoid " of Parker, or rudiment of the sternal
extremity of the coracoid ; ma, metacromial process ;
mss, ossified " mesoscapular segment " ; ost, omosteruum ;
pc, rudiment of precoracoid (Parker) ; ps, presternum ;
sr1, first sternal rib ; sr2, second sternal rib. (From
Flower's Osteology.)

EPICONDYLAR FORAMINA

39

the former is alone concerned with the glenoid cavity. It must
therefore, one would suppose, correspond to the " coracoid " of the
Monotreinata, while the upper piece of bone is the epicoracoid
process of that mammal. The Mammalia, therefore, higher as
well as lower, differ from the reptiles in that the coracoid is
formed of two bones, the exceptions being, among some other
extinct forms, certain of the Anomodontia, a group which it will
be recollected is the nearest of all reptiles to the mammals.

The Fore-limb. — The humerus is of varying length among
mammals. A feature which it sometimes shares with the humerus

FIG. 30.— Distal extrem-
ity of the humerus to
show Epicondylar Fora-
mina. A, In Hatteria ;
B, in a Lizard (Lacerla
ocellata) ; C, in the
Domestic Cat ; D, in
Man. c.e, External con-
dyle ; c.i, internal con-
dyle. In A the two
' foramina are developed
(at i, the entepicondy-
lar ; at ii, the ectepi-
condylar). The only
canal (f) present in the
Lizard (B) is on the
external ulnar side, in
the cartilaginous distal
extremity. In Man (D)
an entepicondylar pro-
cess (pr) is sometimes
developed and con-
tinued as a fibrous band.
(From Wiedersheim's
Anatomy of Man. )

of lower forms is the presence of an entepicondylar foramen, a
defect of ossification situated above the inner condyle of that bone
which transmits a nerve. The same foramen and an additional
ectepicondylar foramen are found in the ancient reptilian type
Hatteria (Sphenodon) ; it occurs also in the Anomodont reptiles.
It is as a rule only the lower forms among mammals which show
this foramen ; thus it is present in the Mole and absent in the

DIGGING FOOT OF MOLE

CHAP.

Horse. The fact that it is occasionally met with in Man is an
additional proof of the, in many respects, ancient structure of the
highest type of Primate.

The radius and the ulna, which together constitute the fore-
arm, are both present in a large number of mammals, btit the
ulna tends to vanish in the purely walking and digitigrade
Ungulates, being present, however, in the
more ancient forms of these Ungulates.
In Man and in many other mammals
the radius can be moved from its normal
position and crossed over the ulna ; this
movement of pronation has been per-
manently fixed in the Elephant, where
the bones are crossed but cannot be altered
in position by the contractions of any
muscles. Other types agree with the
Elephant in this fixation of the two bones.
The bones of the wrist show great
variation among mammals. The greatest
number present are to be seen in such a
type as the Mole. % Here we have a
proximal row, consisting of the scaphoid,
lunar, cuneiform, and pisiform, which are
FIG. 31.— Bones of fore-arm arranged in their proper order, beginning
X-r°x 2M01:,£S with that on the radial side of the limb,

form ; ce, centrale ; I, lunar ; that side which bears the first digit. A
in, magnum; », pisiform; -. ,. -, , •• n -,i

R, ralius ; «, radial sesa- second row articulates proximally with
moid (falciform) ; s, sea- these bonelets and distally with the meta-

phoid ; td, trapezoid ; tm, , t. , ...

trapezium ; u, ulna ; u, carpals ; the bones composing it are, men-
uncitbrm; i-v, the digits, tioning them in the same order, trapezium,

(From Flowers Osteology.) .

trapezoid, centrale, magnum, unciiorm.

The centrale does not, however, really belong to the distal
carpal row, and is as a rule situated in the middle of the carpus
away from articulation with the metacarpals. It is a bone which
is not commonly present in the mammalian hand, but is present
in various lower forms, such as the Beaver and Hyrax. It also
occurs in such high types as the majority of Monkeys ; it is to
be found in the Human foetal carpus. Many extinct forms
possessed a separate centrale. Its importance in the formation of
the interlocking condition of the Ungulate foot is referred to later,

FINS OF WHALES 4 I

on p. 196. The only mammal which appears to have the proper
five bones in the distal row of the carpus corresponding to the
five metacarpals is Hyperoodon, where this state of affairs at least
occasionally occurs. The final bone of that series, the unciform,
seems to represent two bones fused. Very often the carpus
is reduced by the fusion of certain of the carpal bones ; thus
among the Carnivora it is usual for the scaphoid and the lunar
to be fused. It is interestingly significant that these bones retain
their distinctness in the ancestral Creodonts. In many Ungulates
the trapezium vanishes. The reduction of the toes in fact implies
a reduction of the separate elements of the carpus.

As to the digits of the mammalian hand, the greatest number
is five, the various supplementary bonelets known as prepollex
and postminimus being, it is now generally held, merely supple-
mentary ossifications not representing the rudiments of pre-existing
fingers. They may, however, bear claws.1 The number of
phalanges which follow upon the metacarpals is almost constantly
three in the mammals, excepting for the thumb, which has only two.
This is highly characteristic of the group as opposed to reptiles
and birds, and the increase in the number of these bones in the
Whales and to a very faint degree in the Sirenia is a special re-
duplication, which will be mentioned when those animals are
treated of.

The Pelvic Girdle. — The pelvic girdle or hip girdle is the
combined set of bones which are attached on the one hand to
the sacrum and on the other articulate with the hind -limb.
Four distinct elements are to be recognised in each " os inno-
minatum," the name given to the conjoined bones of each half of
the entire pelvis. These are : — the ilium, which articulates with
the sacrum ; the ischium, which is posterior ; the pubis, which is
anterior ; and finally, a small element, the cotyloid, which lies
within the acetabular cavity where the femur articulates. The
epipubes of the Monotreme and the Marsupial are dealt with
elsewhere (see p. 116) as they are peculiar to those groups.

Professor Huxley pointed out many years since that while
the Eutherian Mammalia differ from the reptiles in the fact that
the axis of the ilium lies at a less angle with that of the sacrum,

1 Horny matter is apt to be formed upon extremities ; instances which are well
known are the "claws" upon the tail of the Lion and Leopard and the Kangaroo
Omjchogale. For an account of the first see Proc. Zool. Soc. 1832, p. 146.

42 HIP GIRDLE

Ornithorliynclms comes nearest to the reptile in the fact that
this axis is nearly at right angles to that of the sacrum. It is
particularly interesting to find that this peculiarity of Orni-
thorhynchus is only acquired later in life, and that the pelvis of
the foetus conforms in these angles -to the adults of other
mammalian groups. In any case, the backward rotation of the
pelvis is a mammalian characteristic, and it is most nearly
approached among reptiles by the extinct Anomodontia, whose
affinities to mammals will be dealt with on a later page (p. 90).
Another peculiarity of the mammalian pelvis appears to be the
cotyloid bone already referred to. In the Eabbit this bone
completely shuts out the pubis from any share in the acetabular
cavity ; later it ankyloses with that bone. In Ornithorhynchus
the cotyloid or os acetabuli is a larger element of the girdle
than is the pubis. In other mammals, therefore, it seems to be
a rudimentary structure. But it seems to be a bone peculiar to
and thus distinctive of the mammals as compared with other
vertebrates. The acetabular cavity is perforated in Echidna as
in birds ; but in certain Eodents the same region is very thin
and only closed by membrane, as in Circoldbes villosus.

The number and the arrangement of the bones in the hind-limb
correspond exactly to those of the fore-limb. The femur, which
corresponds to the humerus, shows some diversities of form. The
neck, which follows upon the almost globular head, the surface
of articulation to the acetabular cavity of the pelvis, has two
roughened areas or tuberosities for the insertions of muscles. A
third such area, known as the third trochanter, is present or
absent as the case may be, and its presence or absence is of
systematic import. As a general rule the thigh-bones of the
ancient types of mammals are smoother and less roughened by
the presence of these three trochanters than in their modern
representatives. The radius and the ulna are represented in the
hind -leg by the tibia and the fibula. These bones are not
crossed, and do not allow of rotation as is the case with the
radius and the ulna. In Ungulate animals there is the same
tendency to the shortening and rudimentary character of the
fibula that occurs in the case of the ulna, but it is more marked.
It has been shown in tracing the history of fossil Ungulates that
the hind -limbs in their degree of degeneration are as a rule
ahead of the fore-limbs. This is natural when we reflect that

NUMBER OF TOES

43

the hind -limbs must have preceded the fore -limbs in their

thorough adaptation to the cursorial mode of progression. In

the Mammalia the ankle- join li is always

what is termed cruro-tarsal, i.e. between

the ends of the limb-bones and the proximal

row of tarsals ; not in the middle of the

tarsus as in some Sauropsida (reptiles and

birds). The bones of the ankle are much

like those of the hand ; but there are

never more than two bones in the proxi-

mal row, which are the astragalus and the

calcaneum. The former is perhaps to be

looked upon as the equivalent of the

cuneiform and lunar together. But the

views as to the homologies of the tarsal

bones differ widely. Below these is the

navicular, regarded as a centrale. The

distal row of the tarsus has four bones,

three cuneiforms and a cuboid. Eeduction FlQ> 32._Anterior aspect of

is effected by the soldering together of right femur of Rhinoceros

• n • j_i TT i .LI_ (Rhinoceros indicus). x i.

two cuneiforms as in the Horse, by the

#,Head; t, great trochan-
ter ; t', third troehanter.
(From Flower's Osteology.)

fusion of the navicular and cuboid as in

the Deer. No mammal has more than five

toes, and the number tends to become reduced in cursorial animals

(Eodents, Ungulates, Kangaroos).

Teeth. — The teeth of the Mammalia l differ from those of
other vertebrated animals in a number of important points.
These, however, entirely concern the form of the adult teeth,
their position in the mouth, and the succession of the series of
teeth. Developmentally and histologically there are no funda-
mental divergences from the teeth of vertebrates lower in the
scale.

In mammals, as for example in the Dog, the teeth consist of
three kinds of tissue — the enamel, the dentine, and the cement.
The enamel is derived from the epidermis of the mouth cavity,
and the two remaining constituents from the underlying dermis.
The teeth originate quite independently of the jaws, with which
they are later so intimately connected ; the independence of
origin being one of the facts upon which the current theory
1 Cf. Tomes, A Manual of Dental Anatomy, 5th ed. London, 1898.

44

STRUCTURE OF TEETH

of the nature of teeth is founded.

HI

FIG. 33. — Diagrammatic sections of various forms
of teeth. /, Incisor or tusk of Elephant,
with pulp cavity persistently open at base ;
//, Human incisor during development, with
root imperfectly formed, and pulp cavity
widely open at base ; ///, completely formed
Human incisor, with pulp cavity opening by
a contracted aperture at base of root ; IV,
Human molar with broad crown and two
roots ; V, molar of the Ox, with the enamel
covering the crown deeply folded, and the
depressions filled up with cement ; the sur-

It has been pointed out
that the scales of the Elas-
mobranch fishes consist of
a cap of enamel upon a
base of dentine, the former
being derived from the epi-
dermis and modelled upon
a papilla of the dermis
whose cells secrete the
dentine. The fact that
similar structures arise
within the mouth (i.e. the
teeth) is explicable when
it is remembered that the
mouth itself is a late in-
vagination from the out-
side of the body, and that
therefore the .retention by
its tissues of the capacity
to produce such structures
is not remarkable.

The relations of the
three constituents of the
tooth in its simplest form
is shown in the accom-
panying diagram, where
the intimate structure of
the enamel, dentine, and
cement (or crusta petrosa
as it is sometimes called)
is not indicated. The latter
has the closest resemblance
to bone. The dentine is
traversed by fine canals

face is worn by use, otherwise the enamel

coating would be continuous at the top of the which run parallel to each

ridges In all the tigures the enamel is black, other ftnd aiiagtomose here

the pulp white ; the dentine represented by

horizontal lines, and the cement by dots, and there. The enamel IS

formed of long prismatic

fibres, and is excessively hard in structure, containing less animal
matter than the other tooth tissues. To this fact is frequently

VESTIGIAL TEETH 45

due the complicated patterns upon the grinding teeth of Ungulates,
which are produced by the wearing away of the dentine and the
cement, and the resistance of the enamel.

The centre of the tooth papilla remains soft and forms the
pulp of the tooth, which is continuous with the underlying
tissues of the gum by a fine canal or a wide cavity as the case
may be. In teeth which persistently grow throughout the life-
time of the animal, as for example the incisors of the Eodents,
there is a wide intercommunication between the cavity of the
tooth and the tissues of the gum ; only a narrow canal exists in,
for instance, the teeth of Man, and in fact in the vast majority
of cases. The three constituents of the typical teeth are not,
however, found in all mammals ; the layer which is sometimes
wanting is the enamel. This is the case with most Edentates :
but the interesting discovery has been made (by Tomes) that in
the Armadillo there is a downgrowth of the epidermis similar to
that which forms the enamel in other mammals, a rudimentary
" enamel organ."

Teeth are present in nearly all the Mammalia ; and where
they are absent there is frequently some evidence to show that
the loss is a recent one. The Whalebone Whales, the Mono-
tremata, Manis, and the American Anteaters among the Edentata
are devoid of teeth in the adult state. In several of these
instances, however, more or less rudimentary teeth have been
found, which either never cut the gums or else become lost
early in life. The latter is the case with Ornitlwrhynchiis,
where there are teeth up to maturity (see p. 113). Klikenthal
has found germs of teeth in Whales, and Rose in the Oriental
Manis. The loss of the teeth in these cases seems to have
some relation to the nature of the food. In ant -eating
mammals, as in the Anteaters and Echidna, the ants are
licked up by the long and viscid tongue, and require no
mastication. Yet it must be remembered that Orycteropus is
also an anteater, like the Marsupial Myrmecolius, both of which
genera have teeth.

The first of the essential peculiarities of the mammalian
teeth as compared with those of other vertebrates concerns the
position of the teeth in the mouth. There is no undoubted
mammal extinct or living in which the teeth are attached to
any bones other than the dentary, the maxilla, and the pre-

46 TEETH AND CLASSIFICATION CHAP.

maxilla. There are no vomeriue, palatine, or pterygoid teeth,
such as are met with in Amphibia and Reptilia.

The other peculiarities of the mammalian teeth, though true
of the great majority of cases, are none of them absolutely
universal.

But it is necessary to go into the subject at some length on
account of the great importance which has been laid upon the
teeth in deciding questions of relationship ; moreover, largely no
doubt on account of their hardness and imperishability, our
knowledge of certain extinct forms of Mammalia is entirely based
upon a few scattered teeth ; while of some others, notably of
the Triassic and Jurassic genera, there is not a great deal of
evidence except that which is furnished by the teeth. Indeed
the important place which odontography holds in comparative
anatomy is from many points of view to be regretted, though
inevitable. " In hardly any other system of organs of verte-
brated animals," remarks Dr. Leche, " is there so much danger of
confounding the results of convergence of development with true
homologies, for scarcely any other set of organs is less con-
servative and more completely subservient to the lightest
impulse from without." Affinities as indicated by the teeth are
sometimes in direct contradiction to those afforded by other
organs ; or, as in the case of the simple Toothed Whales, no
evidence of any kind is forthcoming. Dr. Leche has pointed out
that, judged merely from its teeth, Arctictis would be referred to
the Raccoons, though it is really a Viverrid ; while Bassariscus,
which Sir W. Flower showed to be a Raccoon, is in its teeth a
Viverrid. Mr. Batesoii has been obliged to hamper the subject
with another difficulty.

In dealing with the variations of teeth,1 Mr. Bateson has
brought together an immense number of facts, which tend to
prove that the variability of these structures is much greater than
had been previously recognised ; that this variability is often
symmetrical ; and that in some animals, as in " Canis cancrivorus,
a South American fox, the majority showed some abnormality."
When we learn from Mr. Bateson that " of Felis fontanieri, an
aberrant leopard, two skulls only are known, both showing dental
abnormalities," it seems dangerous to rear too lofty a super-
structure upon a single fossil jaw. It must be noted too that,

1 Materials for the Study of Variation, London, 1894.

HETERODONTY AND HOMODONTY

47

contrary to the prevailing superstition, it is not domestic animals
which show the greatest amount of tooth variation. As to special
homologies between tooth and tooth, with which we shall deal
on a later page, Mr. Bateson has urged almost insuperable
difficulties.

The teeth of the Mammalia are almost without exception
" heterodont," i.e. they show differences of structure in different

p.masc

s.oc

oc.co,

par.oc

FIG. 34. — Skull of Dasyurus (lateral view), al.sph, Alisphenoid ; ang, angular process
of mandible ; fr, frontal ; ju, jugal ; Icr, lachrymal ; max, maxilla ; nas, nasal ;
oc.cond, occipital condyle ; par, parietal ; par.oc, paroccipital process ; p.max,
premaxilla ; s.oc, supraoccipital ; sq, sqnamosal ; sq', z\ gomatic process of squa-
mosal. (From Parker a:id Haswell's Zoology.)

parts of the mouth. As a general rule, teeth can be grouped
into cutting incisors, sharp conical canines, and molars, with a

FIG. 35. — Upper and lower teeth of one side of the mouth of a Dolphin (Lagenorhyn-
chus), illustrating the homodont type of dentition in a mammal. (After Flower
and Lydekker. )

surface which is in the majority of cases suited for grinding. In
this they contrast with the majority of the lower vertebrates,
where the teeth are " homodont " (or, better, homoeodonf), i.e. all
more or less similar and not fitted by change of form to perform
different duties. But there are exceptions on both sides. In

48 INCISORS OF RODENTS CHAP.

the Toothed Whales the teeth are homodont, as they are in
the frog and in most reptiles ; on the other hand, some of the
remarkable reptiles belonging to Professor Huxley's order of the
Anomodontia have distinct canines, and show other differentiations
in their teeth.

A second characteristic of the mammalian dentition is the
limited number of the teeth, which rarely exceeds fifty-four.
Here again the Toothed Whales are an exception, the number of
their teeth being as great as in many reptiles. In the Mammalia
the number of the teeth is fixed (excepting of course for ab-
normalities), while in reptiles there is frequently no precise
normal. Two regions may be distinguished in every tooth—
the crown and the root ; the latter, as its name denotes, is
imbedded in the gum, while the crown is the freely-projecting
summit of the tooth. The varying proportions of these two
regions of the tooth enables us to divide teeth into two series —
the brachyodont and the hypselodont ; in the latter the crown is
developed at the ;expense of the root, which is small ; the
hypselodont tooth is one that grows from a persistent pulp or, at
any rate, one that is long open. Brachyodont teeth on the
contrary have narrow canals running into the dentine. The
primitive form of the tooth seems undoubtedly to be a conical
single-rooted tooth, such as is now preserved in the Toothed
Whales and in the canine teeth of nearly all animals. The de-
velopment of the teeth, that is, the simple bell-shaped form of the
enamel organ, seems to go some way towards proving this ; but it is
quite another question whether we can fairly regard the Whales
as having retained this early form of tooth. In their case the
simplification, as is so often the case where organs are simplified,
seems to be rather degeneration than retention of primitive
characters. But this is a matter which must be deferred for the
present.

The incisor teeth are generally of simple structure and nearly
always single rooted. In the Eodents, in the extinct Tillo-
doutia and in Diprotodont Marsupials, they have grown large, and,
as has been already stated, they increase in size continuously
from the growing pulp. These teeth have a layer of enamel
only on the anterior face, which keeps a sharp chisel-like edge
upon them by reason of the fact that the harder enamel is worn
away more slowly than the comparatively soft dentine. The

81

»

:

n CANINES AND PREMOLARS 49

" liorn " of the Narwhal is another modification of an incisor, as
are the tusks of Elephants. Among the Lemurs the incisors are
denticulate, and serve to clean the fur in a comb-like fashion.
This is markedly the case in Guleo-pilhecus. The incisors arc
sometimes totally absent, as in the Sloths, sometimes partial!}
absent, as in many Artiodactyles, where the lower incisors bite
against a callous pad in the upper jaw, in which no trace of
incisors has been found.

Canine teeth are present in the majority of mammals, but
are absent without a single exception from the jaws of the
Eodentia. The canine tooth of the upper jaw is that tooth
which comes immediately after the suture dividing the pre-
maxillary from the maxillary bone. The canines are as a rule
simple conical teeth, with but a single root; indeed they
resemble what we may presume to have been the first kind of
tooth developed in mammals. In this they resemble also as a
;eneral rule the foregoing incisors. But instances are known
where the canines are implanted by two roots. This is to be
seen in Triconodon, in the pig Hyotherium, in the Mole and
some other Insectivores, and in Galeopitliecus, where the incisors
also may be thus implanted in the jaw. Furthermore, the
simple condition of the crown of the tooth may be departed from.
This is the case with a Fruit Bat belonging to the genus Ptera-
lopex. In the more primitive Mammalia it is common to find
no great difference between the canines and incisors ; such is the
case with the early Ungulate types of Eocene times, such as
Xiphodon. In modern mammals, however, especially among the
Carnivora, the canines tend to become larger and stronger than
the incisors, and in some of the Cats and in the Walrus these teeth
are represented by enormous offensive tusks. It is not rare for
the canines of male animals to be larger than those of their
mates. There are also cases such as the Musk-deer and the
Kanchil where the male alone possesses these teeth, but only in
the upper jaw. The teeth which follow the canines are known
as the grinders or cheek teeth, or more technically as premolars
and molars. These two latter terms separate teeth which arise
at different periods, and their use will be explained later. In the
meantime it may be pointed out that the cheek teeth are the teeth
which show the greatest amount of variation in their structure ;
this is shown by the number and variety of the cusps in which
VOL. x E

50 ' CHEEK TEETH

the biting surface ends. The grinding teeth vary from simple
one - cusped teeth, precisely like canines, to teeth with an
enormous number of separate tubercles. In the former case it is
hard to distinguish between incisors, canines, and cheek teeth
in the lower jaw, where no suture separates the bone. More-
over it is quite common for the first cheek tooth in the lower
jaw to have the characters of a canine, while the true canine
approximates in its form to the antecedent incisors. This is
so, for instance, with the Lemurs, where the first premolar is
caniniform, and the canine shares in the curious procumbent
attitude which distinguishes the lower incisors of many of those
animals.

A variable number of the anterior cheek teeth may be little
more than simple conical teeth ; but the rest of the set are
commonly more complicated. No definite laws can be laid down
as to the complication of the posterior as compared with the
anterior set. Broadly speaking, it is purely herbivorous creatures
in which the least difference can be detected at the two ex-
tremities, and which are at the same time the most elaborately
decorated with tubercles and ridges. The converse is true that
in purely carnivorous animals, including insect- and fish-eating
forms, there is the greatest difference between the anterior set
of grinding teeth and those which follow. In these two respects
such animals as a Lemur and a Rhinoceros occupy the extremes.
Furthermore, it may be said that omnivorous creatures lie, as
their diet would suggest, in an intermediate position. Generally
speaking, when there is a marked difference between the first
premolar and molars at the end of the series, there is a gradual
approximation in structure of a progressive kind. The tubercles
become more numerous in successive teeth ; but the corollary
which is apparently deducible from this, i.e. that the last molar
is the most elaborate of the series, is by no means always true.
The last cheek tooth indeed is often degenerate. On the other
hand, it is very markedly the largest of the series in such diverse
types as the Elephant, the hog Phacochoerus, and the Rodent
Hydrochoerus. It is a rule that the cheek teeth of the upper jaw
are more complicated than the corresponding teeth of the lower
jaw.

The structure of the cheek teeth is very diverse among the
Mammalia. Broadly, two types are to be recognised. There are

MILK DENTITION

teeth in which the grinding surface is raised into a series of two,
to many, tubercles sharper or blunter as the case may be ; — sharper
and fewer at the same time in carnivorous and especially in
insectivorous types, more abundant in omnivorous animals. To
this form of tooth the term " bunodont " is applied. There is
no doubt that this is the earliest type of tooth ; but whether the
fewer or the more cusped condition is the primitive one is a
question that is reserved for consideration at the end of the
present chapter. The other type of grinding tooth is known as
" lophodont." This is exemplified by such types as the Perisso-
dactyla and Ungulates generally, and by the Kodents. The tooth
is traversed by ridges which have generally a transverse direction
to the long axis of the jaw in which the tooth lies. The ridges

p.4

p.3

FIG. 36. — Molar teeth of A cemtherium platycephalum. x %. m.\ -?».3, Molars : mh, meta-
loph ; p.l-pA, premolars ; jph, 'protoloph ; ps.f, parastyle fossa ; te, tetartocone.
(After Osborn. )

may be regarded as having been developed between tubercles
which they connect and whose distinctness as tubercles is
thereby destroyed. Lophodont teeth are only found in vegetable-
feeding animals.

The special characteristics of the teeth of various groups of
animals will be considered further under the accounts of the
several orders of recent and fossil Mammalia.

A very general feature of the teeth of the Mammalia is what
is usually termed the diphyodont dentition. In the majority of
cases there are two sets of teeth developed, of which the first
lasts for a comparatively short time, and is termed on account of
its usual time of appearance the " milk dentition " ; this is
replaced later by the permanent dentition. In lower vertebrates
the teeth are replaced as worn away. There is not, however,
so great an antithesis in this matter between the Mammalia

P.I

DEVELOPMENT OF TEETH

and other vertebrates as was at one time assumed. But in
order to explain this very important part of the subject it will
be necessary to give some account of the development of the
teeth. The type selected is the Hedgehog, which has been
recently and carefully described by Dr. Leche. of Stockholm,

en.

^-fiajji

FIG. 37. — Two stages in the development of the teeth of a Mammal (diagrammatic sec-
tioiis). alv, Bone of alveolus ; dent, dentine ; dent.s, dental sac ; en, enamel ;
en.m, enamel membrane ; en.m2, enamel membrane of permanent tooth ; en.plp,
enamel pulp ; gr, dental groove ; lam, dental lamina ; lam', part of dental lamina
which grows downwards below the tooth germ ; n, neck connecting germs of milk
and permanent tooth ; pap, dental papilla ; pap-, dental papilla of permanent tooth.
(After 0. Hertwig.)

which type has furthermore the advantage of being a " central "
type of mammal. The first step in the formation of the teeth
is a continuous invagination of the epithelium covering the jaw
to form a deepish wall of tissue running in the thickness of the
jaw ; this is perfectly continuous from end to end of the lower
jaw. From this " common enamel germ " (Schmelzleiste of the
Germans1) " special enamel germs " (Schmelzorgane, enamel organs)
are developed here and there as thickenings in the form of buds

1 Morpli. Jahrb. xix. 1892, p. 502.

ii MONOPHYODONT DENTITION 53

which arise on the outer side of the fold of epithelium and some
way above its lower termination. These ultimately acquire a
bell-like form, and are as it were moulded on to a thickened con-
centration of the dermis beneath ; they then become separate
from the downgrowth of the epithelium whence they have arisen.
Finally, each of the eight germs becomes one of the milk teeth of
the animal The lower end of the sheet of invaginated epi-
thelium, the common enamel germ, is the seat of the formation
of the second set of teeth, of which, however, in the animal under
consideration, there are only two in each jaw. But corresponding
to each of the enamel germs of the milk dentition, with the
exception of the first two molars, there is a slight thickening of
the end of the common enamel germ, which at a certain stage is
indistinguishable from the thickening which will become one of
the permanent teeth. We have thus the diphyodont arrange-
ment. But this does not exhaust the series of rudimentary teeth,
though no more come to maturity than those whose development
has already been touched upon. In the upper jaw a small out-
growth of the common enamel germ arises above and to the
outer side of the enamel germ of the third milk incisor ; this does
not develop any further, but its resemblance to the commencing
germ of a tooth seems to indicate that it is the remnant of a
tooth series antecedent to the milk series. Furthermore, there
are indications in the fourth premolar of a fourth series of teeth
posterior in appearance to the permanent dentition. We arrive
therefore at the important conclusion that although here as
elsewhere there are only two sets of calcified teeth ever developed,
there are feeble though unmistakable remains of two other series,
one antecedent to and the other posterior to the diphyodont
dentition. The gap therefore which separates the mammalian
dentition from that of reptiles is less than has hitherto appeared.
Dr. Leche also carefully studied the tooth development of Iguana ;
he found that in this lizard there are four series of teeth which
come to maturity, and a rudimentary series antecedent to these
which never produces fully formed teeth.

In a few mammals there is a kind of dentition known as the
monophyodont, in which only one series of teeth reaches maturity ;
where in fact there is no replacement of a milk series by a per-
manent dentition. Of the monophyodont dentition Whales form
an example. The Marsupials are very nearly an instance of the

54 TEETH OF CAPE ANTEATER CHAP.

same phenomenon ; for Sir W. Flower showed, and Mr. Thomas
confirmed his discovery, that only one tooth, according to Mr.
Thomas the fourth premolar, is replaced in that group. But
even the purely monophyodont dentition of the Toothed Whales
is a more apparent than real contrast to the diphyodont dentition
elsewhere prevalent. An investigation of the embryos of various
Toothed Whales by Dr. Kiikenthal and by Dr. Leche has brought
to light the highly important fact that two dentitions are present,
but that one only comes to maturity ; from this fact obviously
follows the interesting question : — To which of the two dentitions
of more normal Mammalia does the monophyodont dentition of the
Whales and Marsupials belong ? To this question a clear answer
is fortunately possible. As has been pointed out in the fore-
going sketch of tooth development, and has been illustrated in
the figures, the milk teeth develop as lateral outgrowths of the
common enamel germ, while the permanent teeth arise from the
end of the same band of tissue. This fact enables it to be
stated apparently beyond a doubt that in the Whales and in the
Marsupials it is the milk dentition which is the only one to
arrive at maturity. Thus the earlier theoretical conclusion that
the Marsupial dentition " is a secondary dentition with only one
tooth of the primary set left," is proved on embryological grounds
to be untrue. But there are other monophyodont animals than
those already mentioned.1 Orycteropus, the Cape Anteater, is an
example. Mr. Thomas has lately discovered that in this Eden-
tate there is a set of minute though calcined milk teeth which
probably never cut the gum ; here we have a different sort of
monophyodontism, in which the teeth belong to the second and
not to the first set. Between the latter condition and the
diphyodont state are intermediate stages. Thus in the Sea Lions
the milk teeth are developed but disappear early, probably before
the animal is born.

In the typical diphyodont dentition, such as is exhibited for
example in Man and the vast majority of mammals, the milk teeth
eventually completely disappear and are entirely replaced by the
permanent set of teeth, with the exception, of course, of the molars,
which though they are developed late belong to the milk series.

1 It would be of the greatest interest 'in relation to this and many other
problems to ascertain the precise meaning of the monophyodont dentition of
Ornithorhynchus.

IT DENTITION OF SPARASSODONTS 55

Their correspondence with the milk series is shown in an
interesting way by the close resemblance which the last milk
premolar often bears to the first molar. These two extremes of
dentition, i.e. purely monophyodont and, excepting for the molars,
purely diphyodont, are however connected by an intermediate state
of affairs, which is represented by more than one stage. In
Borhyacna (probably a Sparassodont) the incisors and the canines
and two out of the four premolars belong to the permanent
dentition, while the two remaining premolars and of course the
three molars are of the milk series. Prothylacinus, a genus
belonging to the same group, has a dentition which is a step or
two further advanced in the direction of the recent Marsupials.
We lind, according to Ameghino,1 whose conclusions are accepted
by Mr. Lydekker, that the incisors, canines, and two premolars
belong to the milk series, while the permanent series is repre-
sented only by the two remaining premolars. We can tabulate
this series as follows : —

(1) Purely monophyodont, with teeth only of the first set —
Toothed Whales.

(2) Incompletely monophyodont, as in the Marsupials, where
there is a milk dentition with only one tooth replaced.2

(3) Incompletely diphyodont, with the dentition made up
partly of milk, partly of permanent teeth, as in Borliyaena.

(4) Diphyodont, where all the teeth except the molars are
of the second set ; this characterises nearly all the mammals.

As we pass from older forms to their more recent representa-
tives there is as a rule a progressive development of the form of
the teeth. This is especially marked among the Ungulata. The
extremely complicated type of tooth found in such a form as the
existing Horse can be traced back through a series of stages to a
tootli in which the crown is marked by a few separated tubercles
or cusps. Arrived at this point, the differences between the teeth
of ancestral Horses and ancestral Ehinoceroses and Tapirs are hard
to distinguish with accuracy ; and the same difficulty is experi-
enced in attempting to give a definition of other large orders by
the characters of the teeth, such as will apply to the Eocene or

1 Proc. Zool. Soc. 1899, p. 922.

2 Mr. M. Woodward, however (P.Z.S. 1893, p. 467), is disposed to think that in
some Macropodidae at any rate the supposed tooth of the second set really belongs
to the milk dentition, arising late between Pra3 and Pm4.

56 CUSP NOMENCLATURE CHAP.

even earlier representatives of these families. Fig. 36 (p. 51)
illustrating a series of mammalian teeth will illustrate the above
remarks. That there is such a convergence in tooth structure
shows that it is, theoretically at least, possible to determine the
ancestral form of the mammalian tooth. Practically, however,
the difficulties which beset such theorising are great ; that there
are such divergent and such strongly-held antithetical views is

sufficient proof of this. Two
main views hold the field :
one, which has found most
favour in America, and is due
chiefly to the labours and per-
suasiveness of Professors Cope,

FIG. 38.— Molar teeth of A, Plienacodus, and Scott, Osboril, and Others, IS
B, the Creodont Palaeonictis. Knd, endo- ,

conid ; hid, hypoconulid ; hyd, hypo- K11OW11 as " trituberculy.
conid ; med, metacomd ; • prd, proto- The alternative vjew as urgetl
oomd. (After Osborn and Wortman.)

by Forsyth Major, "Woodward,

and Goodrich, attempts to show that the dentition of the
original mammal included grinding teeth which were multi-
cuspidate or "multi tubercular." There is much to be said for
both views, and something to be said against both.

This question is, however, wrapped up in a wider one. Its
solution depends upon the ancestry of mammals. If the Mam-
malia are to be derived from reptiles with simple conical teeth,
then the first stage in the development of trituberculy is proved.
On the other hand, however, the evidence is gradually growing
that the Theromorpha represent more nearly than any non-
mammalian group with which we are acquainted the probable
ancestral form of the mammals. These animals offer some
support to both the leading views. Cynoynathus had triconodont
teeth which, as will be pointed out later, are a theoretically
intermediate stage in the evolution of tritubercular teeth ; on
the other hand, the teeth of Diademodon and some others are
multituberculate, and have been very properly compared to the
multitubercular teeth of such primitive mammalia as the Ornitho-
rhynchus. Professor Osborn is no doubt correct in italicising a
remark of an anonymous writer in Science to the effect that in
Diademodon the teeth, though multitubercular, show the pre-
valence of three cusps arranged in the tritubercular fashion.

1 See for a summary, Osborn, American Nat. Dec. 1897, p. 993.

TRITUBERCULY 57

But this may be only a proof that the multitubercular ante-
dates the tritubercular. It may be, indeed, that the mammalian
tooth was already differentiated among the mammal-like Saurians,
and that from such a form as Cynognatlius the Eutheria and other
forms in which a tritubercular arrangement can be detected were
evolved, and from such form as Tritylodon the Monotre-
matous branch of the mammals. This way of looking at the
matter harmonises a much -disputed question, but involves
a diphyletic origin of the mammals — an origin which for other
reasons is not without its supporters.

We shall now attempt to give a general idea of the facts and
arguments which support or tend to support " trituberculy." As
a matter of fact the name is inaccurate ; for the holders of this
view do not derive the mammalian molar from a trituberculate
condition, but in the first place from a simple cone such as that
of a crocodile !

To this main and at first only cusp came as a reinforcement
an additional cusp at each side, or rather at each end, having
regard to their position with reference to the long axis of the
jaw. This stage is the " triconodont " stage, and teeth exist among
living as well as extinct mammals which show this early form of
tooth. We have, indeed, the genus Triconodon, so named 011 that
very account. Among living mammals the Seals and the Thyla-
cine all show some triconodont teeth. A Toothed Whale, it may
be remarked, is a living example of a mammal with monocono-
dont teeth. The three primary cusps, as the supporters of Cope's
theory of trituberculism denominate them, are termed respectively
the protocone, paracone, and metacone, or, if they are in the teeth of
the lower jaw, protoconid, paraconid, and metaconid. At a slightly
later stage, or coincidently, a rim partly surrounded the crown of
the tooth ; the rim is known as the cingulum, and from a pro-
minent elevation of this rim a fourth cusp, the hypocone, was
developed. The three main cones then moved, or rather two of them
moved, so as to form a triangle ; this is the tritubercular stage.
Teeth of this pattern are common, and occur in such ancient
forms as Insectivora and Lemurs, besides numerous extinct groups.
An amendment has been suggested, and that is to term the teeth
with the simple primitive triangle " trigonodont," and to reserve
the term tritubercular for those teeth in which the hypocoiie has
appeared. The platform bearing the hypocone widened into the

^VJHU^|

^ 'o° ' *"'
I JT '^•'^^

•LIBRAF

•58

ORIGIN OF TOOTH PATTERNS

" talon " ; and this ledge became produced into two additional
cusps, the hypoconule or hypoconulid, and the ectocone or ecto-
conid. Thus the typical sextuberculate tooth of the primitive
Ungulate, and indeed of many primitive Eutherians, is arrived at.

.. .frotocmid

frolixoMiJ.s^

I \nutaf°nid

fianuontd • i x//, w

Ayfotantd

FIG. 39. — Epitome of the evolution of a cusped tooth. 1, Reptile; 2, Dromathermm ;
3, Micrnconodon ; 4, Spalacotherium : me, metaconid ; pa, paraconid ; pr, proto-
conid ; 5, Amphitherium. (After Osborn. )

From this the still further complicated teeth of modern Ungulates
can be derived by further additions or fusions, etc.1 On the other
hand, the development of the Primate molar stops short at the
stage of four cusps.

That such a series can be traced is an undoubted fact. Every
stage exists, or has existed. But whether the stages can be con-
nected or not is quite another question. It is by three main
lines of argument that the view here sketched out in brief is
supported. In the first place, the tracing of the pedigrees of
many groups of mammals has met with very considerable success ;
and it is clear that as we pass from the living Horse and
Rhinoceros, with their complicated molars, to their forerunners,
we find that both can be referred to a primitive Ungulate molar
with but six cusps. Going still further back to the lowest
Eocene and ancestral type as it appears, Euprotoyonia, we still
find in the molar tooth the sextubercular plan of structure. We
can hardly get further back in the evolution of the Perissodactyles
with any probability of security. On the other hand, many facts
point to a fundamental relationship between the primitive Ungu-
lates and the early Creodonts. The latter frequently show plainly
tritubercular molars. Such Ungulates as Ewprotogonia and Proto-
f/onodon, though sex- or quinque-tubercular as to their molars,
have a distinctly prevailing trituberculism, when the size and
importance of three of the cusps is taken into account. But this

1 e.g. the " protoloph, " "metaloph," etc. (see Fig. 36, p. 51), of the modern
Ungulate form of tooth.

ii TRITUBERCULY AND SEXTUBERCULY 59

lacks finality as a convincing proof of the tritubercular tooth as
a primitive Ungulate tooth.

Professor Osborn has ingeniously utilised certain deviations
from the normal type of tooth structure (for the group) in favour
of his strongly -urged opinions. If the stages of development
have been as he suggests, a retrogression would naturally be in
the inverse order ; thus the " apparently ' triconodont ' lower
molar of Thylatinua " may be interpreted as a retrogression from
a tritubercular tooth. In the same way may be explained the
triconodont teeth of Seals and of the Cetacean Zeuglodon. Finally,
the modern Toothed Whales have retrograded into " haplodonty."

Embryological evidence has also been called in, and with
some success, to contribute towards the proof of the tritubercular
theory of teeth. Taeker has shown that in the Horse and the
Pig, and some other Ungulates, there is first of all a single
hillock or cusp, and that later the additional cones arise separately.
An apparent stumbling-block raised by these investigations is
that it is not always the protocone or its equivalent in the upper
jaw which arises first, as it obviously ought to do phylogenetically.
This, however, is not a final argument in either direction. We
know from plenty of examples that ontogenetic processes some-
times do not correspond in their order with phylogenetic changes.
Thus in the mammalian heart the ventricle divides before the
auricle ; and of course, phylogenetically, the reverse ought to
occur, since a divided auricle precedes a divided ventricle. This
method of development has, moreover, been interpreted otherwise.
It has been held to signify that the complex teeth of mammals
are indeed derived from simple cones but by the fusion of a
number of those cones.

On the other hand there are the claims of the multituber-
cular theory of the origin of mammalian teeth to be considered.
The palaeontological evidence has been already, to some extent,
utilised. The occurrence of such teeth among the possible fore-
runners of mammals, and in some of the most primitive types of
Mammalia, has been referred to. Senor Ameghino dwells upon
the sextubercular condition of many primitive mammals even
belonging to the Eutheria. In a recent communication1 he attempts
to identify six tubercles in the molars of types belonging to a

1 " On the Primitive Type of the Plexodont Molars of Mammals," Proc. Zool.
Soc. 1899, p. 555.

60 MULTITUBERCULAR TEETH CHAP.

variety of Orders. The same condition, as has been noted, char-
acterises that ancient Ungulate form Euprotoyonia. Even where
the teetli seem at first sight to be tritubercular a detailed study
shows traces of otherwise vanished cusps.

It must be remembered in basing arguments upon the early
Jurassic and Cretaceous mammals, that our knowledge of them
mainly depends upon lower jaws, the teeth of which are usually
simpler in pattern than those of the upper jaws. Moreover,
another fact, not always insisted upon, must not be lost sight of.
In many of those creatures the jaws were of small size, and yet
accommodated a large series of molar teeth. AmyliitlieriuTin, for
example, had six molar teeth, and five is a number frequently
met with. As the teeth are so numerous and the jaws so small
it seems reasonable to connect the simplicity of the structure of
the teeth with the need for crowding a number together. The
same argument may partly account for the superabundant teeth
of many Toothed Whales. It is true that the Manatee has very
numerous grinders which are yet complex ; but then in this
animal there is a succession, and the jaw does not hold at a
given time the entire series, with which it is provided in relays.
On the other hand, where there are few molars they are often
of the multitubercular type, or at least approach it ; of this
the Multituberculate Polyrnastodon is a good example ; so, too, the
molars of Hydroclioerus, and of many other Bodents.

It is well known that the fourth deciduous molar of the
upper jaw, which is replaced by a permanent premolar in the
fully adult animal, is of a more complex structure than its
successor. This may indeed be extended to premolars earlier in
the series. In the Dog " the second and first milk molars closely
resemble the third and second premolars " ; now the milk premolars
belong evidently to the same dentition as the permanent molars,
and they are earlier teeth than the later-developed replacing
teeth. It is therefore significant that these earlier teeth should
be more cuspidate than the later teeth. It tells distinctly in
favour of the simplification as opposed to the complication of
teeth in time, in the groups concerned.

These facts may possibly be applied in explanation of the
simple teeth of some of the Jurassic and Cretaceous mammals.
It has been mentioned that absolute trituberculy is exceedingly
rare among those ancient creatures ; more generally there are to

MOUTH AND TONGUE

6r

be found at least traces of more cusps. Now in some of them
we may be dealing with instances of a complete tooth change ;
the suppression, save for one tooth, which is found in Marsupials,
was probably not developed in at least some of these early
mammals. The simplicity may therefore have been preceded
t>y complexity, and may have been merely an adaptation to an
insectivorous diet.

Alimentary Canal. — The mouth, of the Mammalia is remark-
able for the fact that with a few exceptions, such as the Whales,
there are thick and fleshy lips. The
office of these is to seize the food.
The roof of the mouth is formed by
the " hard palate " in front, which
covers over the maxillary and pala-
tine regions. This region is often
covered with raised ridges, which
have a symmetrical disposition, and
are particularly strong in Ruminant
animals. They are much reduced in
the Eodents, where the anterior part
of the palate is ill-defined owing to
the way in which its sides fade into
the lateral surface of the face. It
has been shown that these ridges,
in the Cat at least, develop as
separate papilliforrn outgrowths, and
it has been suggested that these
papillae, which later become united
to form the ridges, are the last remnant of palatine teeth such
as occur in lower vertebrates.

The tongue is a well -developed organ, usually playing a
double part. It acts as an organ of prehension, especially in
such animals as the Giraffe and the Anteater, where it is
long and protrusible beyond the mouth for a considerable dis-
tance. It also carries gustatory organs, which serve for the
discrimination of the nature of the food. Beneath the tongue
there may be a hardish plate, known as the sublingua. This is
especially prominent in the Lemurs, where it projects as a horny
structure below the tongue, and has an independent and free tip.
It is supported in some of these animals by a cartilaginous

FIG. 40. — Palatal folds of the Raccoon
(Procyon lotor). p.p, Papilla pala-
tina ; r.p, palatal folds. (From
Wiedersheim's Structure of Max.)

62 INTESTINAL MEASUREMENTS CHAP.

structure. It is held by Gegenbaur that this organ is the
equivalent of the reptilian tongue, and that in the skeletal
vestiges which it contains are to be found the equivalents of the
hyoid skeletal cartilages which support the tongue in lizards.
In this case the tongue of mammals is a subsequently added
structure.

The oesophagus leads from the mouth cavity to the stomt //•//.
The latter organ has commonly a distinctive shape in mammals.
This is well shown in Man. The orifices of the oesophagus and
intestine are somewhat approximated ; and this causes a bulging
of the lower border of the organ, usually spoken of as the greater
curvature. A stomach of this typical form is found in many
orders of mammals, and is unlike the stomach in any of the
groups of lower vertebrates in shape. Sometimes the shape of
the organ is greatly altered : it may be drawn out, sacculated,
or divided, as in the Ruminants and Whales, into a series of
differentiated chambers, each of which plays some special part in
the phenomena of digestion.

The intestine of mammals is always long and much coiled,
though the length and consequent degree of coiling naturally
varies. On the whole it is perhaps safe to say that it is shorter
in carnivorous than in vegetable-feeding beasts. Thus the Paca
has an intestine of 39 inches total length, while the Cat, an
animal of about the same size, has an intestine which is only
36 inches long. A fish diet, however, to judge from the Seals,
is associated with a long intestinal tract. The intestine is
divisible in the vast majority of mammals into a small and a
large intestine. The two are separated by a valvular constriction
save in certain Carnivores ; and in the majority of cases the
distinction is also emphasised by the presence at the junction of
a blindly-ending diverticulum, the caecum. This latter organ
varies greatly in length, being very short in the Cat -tribe and
exceedingly long in Rodents. Its size is, to some extent, de-
pendent upon the flesh -eating or grass -eating propensities of
the animal in which it occurs. One of the longest caeca is
possessed by the Vulpine Phalanger, in which the organ is one-
fifth of the length of the small intestine ; while the opposite
extremity is reached by Felis macroscelis, which has a small
intestine one hundred times the length of the caecum.

An interesting point in connexion with the gut of mammals

SMALL AND LARGE INTESTINE

is the varying proportion of the small to the large intestine.
As a general rule the former is very considerably longer than

De

Dn

FIG. 41. — Different forms of the stomach in Mammals. A, Dog ; B, Mus decumanus ;
C, Mus musculus ; D, Weasel; E, scheme of the Ruminant stomach, the arrow
with the dotted line showing the course taken by the food ; F, Human stomach,
a, Minor curvature ; b, major curvature ; c, cardiac end. G, Camel ; H, Ediidna
uculeata. Oma, Major curvature ; Cmi, minor curvature. I, Bradypus tridactylus.
Du, Duodenum ; MB, coecal diverticulum ; **, outgrowths of duodenum ; f, re-
ticulum ; tf> rumen. A (in E and G), Abomasum ; (7a, cardiac division ; O,
psalterium ; OK, oesophagus ; P, pylorus : R (to the right in E and to the left in G),
rumen ; R (to the leQ; in E and to the right in G), reticulum ; Sc, cardiac
division; Sp, pyloric division; 1\'Z, water-cells. (From Wiedersheim's Own'-
parative Anatomy.)

the latter ; in Paradoxnrus, for instance, the small intestine
may be fifteen times the length of the large. The excess of
length of one section over .the other is not generally so marked

64

PLAN OF MAMMALIAN LIVER

as this. In Plialanger maculatus the two sections of the gut are
as nearly as possible equal in length, while in Phascolarctos the
large intestine is considerably longer than the small, the lengths
being respectively 160 inches and 111 inches. It is common
among the Marsupials and also among the Rodents for these
proportions to exist, i.e. for the large intestine to be as long as,
or longer than, the small. But there are so many exceptions
that no general statements can be extracted from the facts.

Some few details will be found in the systematic part of this
book. Mr. Chalmers Mitchell has brought forward some reasons
for associating a great length of large intestine with an archaic
systematic position, in the birds at any rate. The facts here
briefly touched upon are not at variance with the extension of
such a view to the mammals.

Appended to the alimentary tract are three glands or sets of
glands. Opening into the mouth cavity are the salivary glands,

which are of enormous
size in Anteaters, and
small or absent in
Whales. In their num-
ber and position these
glands are character-
istic of mammals. Into
the intestine open the
ducts of the pancreas
and liver, two glands
> which the mammals

u share with lower verte-

brates. The form of

FIG. 42. — Diagrammatic plan of the liver of a Mammal . ,

(posterior surface). c, Caudate lobe; cf, cystic the Mer IS, however,
fissure; dv, ductus venosus ; g, gall-bladder; Ic, generally characteristic
left central lobe ; If, left lateral lobe ; llf, left » *
lateral fissure ; p, portal vein entering transverse OI mammals. It IS
fissure ; re, right central lobe ; rl, right lateral lobe ; (Jjyj(Je(J as y rule into
rlfj right lateral fissure ; s, Spigelian lobe ; u, um-
bilical vein ; vc, post-caval vein. (After Flower a right and a left half,
andLydekker.) the Hne of division

being marked by the insertion of the umbilical ligament, a
vestige of the primitive ventral mesentery. Each half is again
commonly subdivided into central and lateral lobes. In addition
to these, two other divisions are often to be seen — the Spigelian
and the caudate lobe. The liver is less divided in Cetacea and

ii THE HEART AND ITS VALVES 65

some others, very much- subdivided in Eodents and other groups.
The degree of subdivision and the proportions of the several lobes
frequently offer valuable systematic characters. The gall-bladder
may be present or absent; it is always a diverticulum of the
hepatic duct. The two are never separate, as in birds, for
instance.

Organs of Circulation. — The heart of all mammals is a com-
pletely four-chambered organ. In the adult heart there is no
communication between the right and left halves. The auricles
are comparatively thin- walled, the ventricles thick -walled, in
relation to the amount of work that they have severally to per-
form. The right ventricle, moreover, which has only to drive
the blood into the lungs, is much thinner-walled than the left
ventricle, which is concerned with the entire systemic circulation.
The exits of the arteries and the auriculo-ventricular orifices are
guarded by valves, which are so arranged as only to permit the
blood to flow in the proper direction. But these valves have a
morphological as well as a physiological interest. At the origin
of each artery, the aorta and the pulmonary, there is a row of
three watch-pocket valves, as they have been generally termed on
account of their form. These three valves meet accurately in
the middle of the lumen of the arterial tube when liquid is
poured into them from above, and thus completely occlude the
orifice. The auriculo-ventricular valves differ in structure in the
two ventricles. That of the left ventricle has only two flaps,
and is therefore often spoken of as the bicuspid or mitral valve.
Both these flaps are membranous, and together they completely
surround the exit from the auricle into the ventricle. The edges
of the valve are bound down to the parietes of the heart by
numerous branching tendinous threads, the chordae tendineae,
which often take their origin from pillar- like muscles arising
from the walls of the heart, the so-called musculi papillares. The
valve of the right ventricle is composed of three flaps, and is
therefore often spoken of as the tricuspid valve ; it is in the same
way membranous, and has chordae tendineae and musculi
papillares connected with it. The disposition of the musculi
papillares and their number differ in different mammals, but no
exhaustive study has as yet been made of the arrangements in
different groups ; the amount of individual variation even is not
known, though it is certainly considerable in some cases, for in-

VOL. x F

66 HEART OF MONOTREMATA CHAP.

stance in the heart of the Rabbit. The heart of the Monotre-
mata presents differences of some importance from those of other
Mammalia ; the modern knowledge of the Monotrematous heart
is mainly due to Gegenbaur l and Lankester,2 in whose memoirs
references to the older literature will be found. The principal
features of interest in which the heart of the Monotremata differs
from that of the higher Mammalia are these. When the two
ventricles are cut across transversely, the cavity of the right is
seen to be wrapped round that of the left in a fashion precisely
like that of the bird's heart ; on the other hand in the higher
mammal the two cavities lie side by side. The main difference
between Monotremes and other Mammals concerns the right
auriculo- ventricular valve. The differences which it presents
from the corresponding structure of the rest of the Mammalia
are two : in the first place, the valve itself does not com-
pletely surround the ostium ; it is only developed on one side ;
the septal half (i.e. that turned towards the interventricular
septum) is either entirely absent or more generally represented
by a small bit of membrane ; nevertheless I found 3 recently in
an Ornithorliynchus heart a complete septal half to the right
auriculo -ventricular valve. The second point of interest in
connexion with this valve is, that the musculi papillares instead
of ending in chordae tendineae attached to the free edge of the
valve are directly attached to the valve, and in some cases pass
through its membranous flap, to be attached to its origin at the
boundary of the auricle and of the ventricle. The invading of
the valve-flap by muscle in this way is highly interesting, as it
recalls the heart of the bird and of the crocodile. The im-
perfect condition of the valve (from which, as has already been
stated, the septal half is as a rule nearly absent) is a point of
resemblance to the heart of the bird ; the corresponding valve
of the crocodile's heart being complete.

There are also features in the system of arteries and veins
which are eminently distinctive of mammals. In the first place,
the aorta leaving the heart and conveying blood to the body
is only a half arch, and bends to the left side as seen in
Fig. 43. The right and left halves are present in reptiles,
and meet behind the heart. In the bird the right half alone

1 Jen. Zeitschr. ii. 1866, p. 365.
2 Proc. Zool. Soc. 1883, p. 8. 3 Proc. Zool. Soc. 1894, p. 715.

CAVITIES OF HEART

has remained. This fact, therefore, shows that the mammal
cannot have been derived from a bird-like ancestor, but that

FIG. 43. — Lepus cunicnlus. Ven-
tral view of the vascular sys-
tem. The heart is somewhat
displaced towards the left of
the subject ; the arteries of
the right and the veins of the
left side are in great measure
removed, a.epg, internal mam-
mary artery ; «/, anterior facial
vein ; a.m, anterior mesenteric
artery ; a.ph, anterior phrenic
vein ; az.v, azygos vein ; br,
l>rachial artery ; c.il.a, common
iliac artery ; c.il.v, common
iliac vein ; cce, coeliac artery ;
d.ao, dorsal aorta : e.c, external
carotid artery ; e.il,a, external
iliac artery ; e.ti.v, external
iliac vein ; e.ju, external jugu-
lar vein ; fm.a, femoral artery ;
fm.r, femoral vein ; h.v, hepatic
veins ; i.c, internal carotid
artery ; i.cs, intercostal vessels ;
i.il.a, internal iliac artery ; i.il.v,
internal iliac vein ; i.ju, internal
jugular vein ; i.l, iliolumbar
artery and vein ; in, innomin-
ate artery ; l.au, left auricle ;
l.e.c, left common carotid ar-
tery ; l.pr.c, left pre-caval vein ;
/.?', left ventricle ; m.sc, median
sacral artery ; p.a, pulmonary
artery; p.epg, epigastric a rtery
and vein ; p.f, posterior facial
vein ; p.m, posterior meseu-
teric artery ; p.ph, posterior
phrenic veins ; pt.c, jiost-caval
vein ; p.v, pulmonary vein ; r,
renal artery and vein ; r.av,
right auricle ; r.c.c, right com-
mon carotid artery ; r.prc, right
pre-caval vein ; r.v, right ven-
tricle ; s.cl.a, right sulxilavian
artery ; s.rt.i; subclavian vein ;
s]>m, spermatic artery : s.vs,
vesical artery ; ut, uterine artery
and vein ; vr, vertebral artery.
(From Parker's Zootomy.)

both must have independently come from an ancestor with
both halves of the aortic arch present, of which one half
has disappeared in one group, and the other half in the
other. It is an interesting fact, too, to notice that the four

68 AZYGOS VEIN

cavities of the mammal's heart, which fourfold division it shares
with birds alone, do not exactly correspond compartment for
compartment with those of the bird's heart, at least in so far as
concerns the ventricles. For the reptilian heart is provided with
only one ventricle, and therefore the division of that cavity
must have been independently accomplished in mammals and
in birds.

There are two features in the venous system which distinguish
all the Mammalia (with the exception of Echidna in one of
these points) from vertebrates standing lower in the series. The
hepatic portal system is limited to a vein which conveys to the
liver blood derived from the alimentary tract ; in no mammal
except in Echidna is there any representative of the anterior
abdominal vein of lower vertebrates. In that animal there is
such a vein, which apparently arises from a capillary network
upon the bladder and passes up, supported by a membrane, along
the ventral wall of the abdomen to the liver, thus emptying
blood into that organ exactly as does the anterior abdominal
vein of the frog. In no mammal is there any trace of a renal
portal system. The kidneys derive their blood from the renal
arteries only.

Many mammals have two superior venae cavae ; this is the
case, for instance, in the Elephant and the Kodents and other
types lying comparatively far down in the series. In most if
not in all mammals there are considerable remains of one of the
posterior cardinals, in the form of the azygos vein, which opens
into the vena cava superior or pre-caval vein, i.e. the superior
cardinal just before the latter debouches into the heart. This
one posterior cardinal is usually on the right side ; but it may
be on the left side, for instance in Trichosurus vulpecula. In
Halmaturus bennettii there are two azygos veins, one left and one
right, of which the left is rather the larger.1

Urinary Organs. — The kidneys in the Mammalia have a
compact form, which contrasts with the somewhat diffuse and
vaguely -outlined kidneys of the Sauropsida. In mammals the
organ is as a rule of that peculiar shape which is called " kidney-
shaped "; a depression termed the hilum, which receives the
ducts of the glands, indenting the border of an otherwise oval-
shaped gland. In some few mammals the kidney is broken up
1 Beddard, Proc. Zool. Soc. 1895, p. 136.

n BODY CAVITY AND DIAPHRAGM 69

into lobules ; this is the case with the "Whales, the Bears, the
Oxen, and a few other forms. A curious fact about the kidneys
of the Mammalia is their very general asymmetry of position.
One of them usually lies in a more advanced position than the
other. The ureters lead from the kidneys to the urinary bladder,
which in its form and relations is quite distinctive of the
Mammalia. The bladder is formed out of the remains of the
allantois, and is therefore not the exact homologue of the bladder
of the frog, which is the equivalent of the entire sac which grows
out of the cloaca in the mammal, and is the foetal allantois.
The ureters open into the bladder in the higher Mammalia, but
lower down in the urino-genital passage in the more primitive
mammals.

The Body Cavity. — The Mammalia differ from all other living
vertebrates by the arrangement of the body cavity in which lie
the viscera. That cavity is divided into two by a partly muscular
and partly tendinous partition, the diaphragm. N"o other verte-
brate has this precise disposition of the coelom. The diaphragm
lies usually transversely to the longitudinal axis of the body,
but gets a much more oblique arrangement in the Cetacea and
the Sirenia, whose needs demand a more expanded chamber for
the lungs. For hi front of the diaphragm lie the lungs and
heart ; behind it the stomach, liver, intestines, and the organs
of reproduction and excretion. The diaphragm is used in re-
spiration ; when its muscles contract, the surface directed toward
the pleural cavity becomes less convex, and the cavity of the lungs
is thus increased, allowing them to expand under the pressure of
the entering air.

The Lungs. — The lungs of the Mammalia differ from those
of animals lying lower in the series by the fact, just referred to,
that they occupy a pleural cavity completely shut off from the
abdomen by the diaphragm. As a rule the lungs of the Mammalia
are to be distinguished by their more or less extensive lobation.
In the Whales, however, and in the Sirenia, they are not much
divided, but present the appearance of the simple sac-like lungs
of the reptiles. In some mammals there is a median and
posterior unpaired lobe of the lung, which lies in the post-
pericardial cavity behind the pericardium. This is not uni-
versally present. The lungs are very frequently not symmetrical
in their lobation, the number of separate lobes on the right side

OVA OF MAMMALS

and on the left being different. The lungs of mammals agree
with those of the lower reptiles in being freely suspended within
their coelomic cavity, and in not being, as in birds, crocodiles,
and the Varanidae among lizards, tied down to the dorsal surface
of that cavity by a sheet of peritoneum covering them.

The Gonads (Ovaries and Testes).- — The ovary in the
Mammalia is always paired ; there is never a partial or com-
plete abortion of one gonad as in birds — except of course in
pathological cases. The ovaries are small, and lie in the

fir.ov

bl.-u

FIG. 44. — Part of a sagittal section of an ovary of a child just born, bl.v, Blood-vessels ;
foil, strings and groups of cells derived from the germinal epithelium becoming
developed into follicles ; g.ep, germinal epithelium ; in, ingrowing cord of cells
from the germinal epithelium ; pr.ov, primitive ova. (From Hertwig, after
Waldeyer.)

abdominal cavity behind the kidneys. In the immense majority
of the Mammalia the ova which are produced within the
ovaries are of minute size ; those of even the colossal Rorqual
are, so far as we know, not markedly larger than the ova of a
Mouse. The smallness of size of these reproductive elements
implies necessarily an absence of much nutritive yolk ; and as a
consequence the developing embryo, since it is not hatched in an
early stage as a free living larva, has to be nourished by the
mother, to whose tissues it is attached through the intermediary
of the placenta, a structure partly composed of foetal structures
derived from the embryo, and partly of portions of the lining
membranes of the uterus of the mother. The ova of the

DEVELOPMENT OF OVA

Eutherian mammals, including the Marsupials, are very small as
compared with those of any other vertebrates, excepting only
Amphioxus, where the young are hatched early as free swimming
larvae. They also differ in a highly characteristic way in the
mode of their development within the ovary. These processes
are to some extent illustrated in Fig. 44. The main frame-
work of the ovary is formed of the so-called " stroma," which is
a mass of tissue formed of more or less connective-tissue-like
cells. Within this are numerous cavities, the Graafian follicles.
The very young follicles consist of but a single layer of follicular
cells surrounding the ovum, which lies centrally. The follicular

disc

•mernti

FIG. 45. — Two stages in the development of the Graafian follicle. A, With the folli-
cular fluid beginning to appear ; B, after the space has largely increased, caps,
Capsule ; disc, cumulus proligerus ; mernb, membrana granulosa ; ov, ovum ; sp,
space containing fluid. (After Hertwig.)

cells gradually increase in number until the ovum lies in the
midst of several layers of cells. At this period a vacuity is
formed between some of these cells, and grows into a large
cell-free cavity ; the ovum does not lie loosely in this space, but
is connected at one side with the follicular cells, which still line
the interior of the Graafian follicle by the so-called discus
or cumulus proligerus. The egg or ovum has, moreover, a layer
of cells immediately surrounding itself. All these facts can be
gathered by an inspection of Fig. 45. It has been shown
that, as in lower vertebrates, the cells immediately surrounding
the ovum are connected with it directly by delicate processes
which penetrate the actual membrane of the egg.

The only ova which depart at all in structure from that above
described are those of the Monotremata. The credit of this

OVA OF MONOTREMES

discovery rests with Owen and with Professor Poulton, who
pointed out in 1884,1 that the ovum of Ornithorliynchus is very-
large as compared with those of other Mammalia (6 mm. as
against '2 mm.), that it is filled with yolk, and that it completely
fills the follicle, being surrounded by two layers of follicular
cells only. This latter fact was proved by Caldwell. Subse-
quently Gyldberg 2 and 1 3 described the ovarian ovum of Echidna,

FIG. 46. — Ovarian egg of Echidna, b, Basilar membrane ; fe, follicular epithelium ;
o, oil globules ; vm, vitelline membrane ; yl, y2, yolk. (Partly after Caldwell.)

showing it to be identical with that of Ornithorhynclius. Later
still a more elaborate and beautifully illustrated paper was pub-
lished by Caldwell4 upon the early stages of development in the
Monotremata and Marsupials, in which the ovum of the former
was accurately described (see Fig. 46). In the particulars
mentioned above, the ovum of the Monotremata is practically
identical with that of the large-yolked ova of the Sauropsida.

1 Quart. Journ. Micr. Sei. xxiv. 1884, p. 9.

- S.B. Jen. Gesells. 1885, p. 1.

3 Proc. Roy. Phy's. Soc. Edin. viii. 1885, p. 354.

4 Phil. Trans, clxxviii. 1887. p. 463.

OVIDUCT

73

It is the general rule among vertebrate animals that the
ovaries are completely independent of the ducts which convey
their products to the exterior. In certain fishes, however, there
is an absolute continuity between the two structures, which is
believed to be due to a simple concrescence between the originally
distinct ovary and oviduct. The latter has grown round the
former, an obvious advantage in preventing the eggs from
wandering into the abdominal cavity and becoming lost. In
the Mammalia we find discontinuity as a general rule. But in
quite a number of
forms folds of the
lining membrane of
the abdominal cavity
are developed, which
practically ensure
the passage of the
ova into the ovi-
duct when they are
extruded from the
ovaries. The ovi-
duct, moreover, has
a large and fimbri-

lut'

ated mouth, called •* ' rut

in Tinman anptrmTn- FIG. 47. — Lepus cuniculus. The anterior end of the vagina,

111 llUllldll dildtOIlly ... ., f . , . i- n • *_ i_ j /vr,,i

J with the right uterus, Fallopian tube, and ovary. (A at.

— Which is provided size.) Part of the ventral wall of the vagina is removed,

«7^>i rr>V><«. r>f antl tne proximal end of the left uterus is shown in

longitudinal section, fl.t, Fallopian tube ; fl.t', its peri-

fanciful names • toneal aperture ; Lut, left uterus ; l.ut', left os uteri ;

,1 j. -i |. ov, ovary ; r.ut, right uterus ; r.ut', right os uteri ; s,

tne morSUS aiaDOll. vaginal septum ; va, vagina. (From Parker's Zootomy.)

This almost wraps

round the ovary, and thus prevents the ova from straying in
the wrong direction. Moreover, the ovary itself is often so
arranged that it can easily be withdrawn into a pocket of
the peritoneum, from which the obvious exit is by the gaping
mouth of the oviduct. This disposition of the generative parts
is still further modified in a few animals, such as the Eat1 and
the Kinkajou.2 In these animals the mouth of the oviduct
actually opens into the interior of a closed chamber which con-
tains the ovary. In this case there is but one route for the

1 Robinson, Studies Biol. Lab. Owens Coll. ii. 1890, p. 35.
3 Beddard, Proc. Zool. Soc. 1900, p. 667.

74

OVIDUCTS OF MARSUPIALS

FIG. 48. — Female urino - genital
apparatus of various Marsu-
pials. A, Didelphys dorsigera
(young) ; B, Trichosurus ;
C, Phuscolomys wombat. B,
Urinary bladder ; Cl,
" cloaca " ; Fim, fimbriae ;
g, clitoris ; N, kidney ; Od,
Fallopian tube ; Ot, aper-
ture of Fallopian tube; Ov,
ovary ; r, rectum ; Sp, sep-
tum dividing vagina ; Sug,
urino - genital sinus ; Ur,
ureter ; Ut, uterus ; Ut',
opening of the uterus into
the median vagina ( VgB) ;
Vg, lateral vagina ; Vg', its
opening into the urino-genital
sinus ; t (in B), point of
approximation of uteri ; t (in
C) and *, rectal glands. (From
Wiedersheim's Comparative
Anatomy.)

extruded ova to follow. This series of steps in the perfecting
of the mode of safe extrusion of the ova is highly interesting,

THE BRAIN 75

and is a piece of evidence in favour of the high position of the
mammals.

The oviducal apparatus of the mammal is more specialised
than that of lower vertebrates. It is most simple, as might be
imagined, in the egg-laying Monotremes, where, indeed, it is on
the same level as that of reptiles. But in the Eutheria the
fimb dated mouth of the oviduct passes into a narrow and wind-
ing tube, the Fallopian tube ; this widens into a uterus, and the
two uteri combine into a single tube in the higher forms. They
are called the Monodelphia on this account. In the Marsupials
the uteri are distinct though they often join above, and from
this junction depends a median " uterus." After the uterus or
the uteri follows in every case a single vagina.

The testes of the Mammalia, like those of other vertebrates,
occupy primitively a position within the body cavity precisely
corresponding to that of the ovaries. And in the lowly-organised
Monotremata, and some other forms, such as the Whales, they
retain that primitive position within the body. It is, however,
distinctive of the Mammalia as opposed to lower vertebrates that
the testes descend later into a scrotum, which is simply a pro-
trusion of the skin of the body surrounded by muscles, and, of
course, containing a section of the body cavity in which lie the
testes. The penis of the Mammalia, represented by the clitoris
and associated structures in the female, is of a structure entirely
peculiar to this group.

The Brain. — Inasmuch as Professor Wiedersheim has said
with perfect truth that " the brain of the extinct Ungulate
Dinoceras shows so striking a likeness to that of a lizard that
one would be compelled to explain it as that of a lizard without
a knowledge of the skeleton," it is clear that to define the
mammalian brain is a difficult matter. The existing Mammalia,
however, all possess brains which can be readily distinguished
from those of vertebrates lying lower in the scale. They are
of relatively large size, brought about mainly by the dimensions
of the cerebral hemispheres, which have an importance in this
class of vertebrates that they have not elsewhere. Coupled
with this large size of the hemispheres is a more elaborate
system of transverse commissures uniting the two ; and this
culminates in the higher Mammalia, where the corpus callosum
attains a large size and great physiological importance. A

CONVOLUTIONS OF THE BRAIN

very marked feature, moreover, of the mammal's brain is the
development of regular fissures upon its surface, which fissures
are only absent from Ornithorhynchus, various small Eodents,

B

FIG. 49. — Brain of Dog. A, ventral ; B, dorsal ; C, lateral aspect. B.ol, Olfactory lobe ;
Or.ce, cmra cerebri ; Fi.p, great longitudinal fissure ; Hit, HH1, lateral lobes of
cerebellum ; Hyp, hypophysis ; Med, spinal cord ; NH, medulla oblongata ; P»,
pons Varolii ; VH, cerebral hemispheres ; Wu, middle lobe (vermis) of cerebellum ;
I-XII, cerebral nerves. (From Wiedersheim's Comparative Anatomy.)

Bats, and Insectivores, among living mammals. It is some-
times, but erroneously, said that the more complicated the
fissures of the brain are, the higher in intelligence and " zoological
position " is the possessor of that brain. Instances can un-
doubtedly be quoted to support such a view ; but they are

OPTIC LOBES

77

merely selected cases, which do not indicate a wide applicability
of such a generalisation. Thus it is true that the brain of a Man
is more elaborate in its furrows and convolutions than is that of
a Cat. The real fact of the matter is, that the complexity of the
brain from this point of view increases with the size of the animal
within the group.

The Gorilla and the Chimpanzee have a more furrowed brain
than has the little Marmoset ; the Bear a more complicated
brain than the Weasel, etc. The most highly -convoluted brains
of all mammals are those of the Elephants, and there does not

c.A~' ol'

%&**

. NC.THOS

Z ', \fwf\ \\ <^N

o.ch. ^nJ- ptyXmxo. p.va. v.vn

f-fo. rJcJ,,?

FIG. 50. — Lepus cuniculus. Longitudinal vertical section of the brain. (Nat. size.)
a.co, Anterior commissure ; b.fo, body of the fornix ; cb, cerebellum, showing arbor
vitae ; c.c, crus cerebri ; c./t1, parencephalon or cerebral hemisphere ; e./i.2, temporal
lobe ; C.TH#, corpus mammillare ; cp.d, corpus caljosum ; /.?«, foramen of Monro ;
inf, infundibulum ; l.t, lamina terminalis ; ly, lyra ; m.co, middle commissure ;
m.o, medulla oblongata ; o.ch, optic chiasma ; o.l1, o.P, corpora quadrigemina or
optic lobes ; olf, olfactory lobe ; p.cn, posterior commissure ; pd.ym, peduncle of the
pineal " gland," pn; p.fo, anterior pillar of the fornix ; pty, pituitary body ; pv.a,
pons Varolii ; sp.lu, septum lucidiim ; IA, fourth ventricle ; vl.ip, velum interposi-
tum ; v.vn, valve of Vieussens ; II, optic nerve. (From Parker's Zootomy.)

seem in the Ungulates to be so marked a relation between size
and abundance of fissures as there is among other mammals.
A regular plan of the fissures can be detected with certainty
for each group considered by itself; but it is not so easy to
homologise the details of arrangement from group to group.
This is so far in accord with the view that the existing groups
of mammals have diverged from each other ab initio.

Another marked characteristic of the mammalian as opposed
to other brains is the relatively small importance in size and yet
the fourfold nature of the optic lobes. What was the case with
the optic lobes of the early Ungulates is difficult to understand,
on account of the fact that the casts are necessarily imperfect.

7 8 GEOGRAPHICAL RANGE

CHAP.

Altogether the enormous progress in the complexity of the
brain from the early Tertiary mammals down to the present, is
one of the most remarkable revelations of palaeontology. It
goes perhaps some way in explaining the remarkable diversity
in mode of life exhibited by the mammals as compared, for
example, with the birds, whose brains have not diverged so much
or in so many directions from the primitive form.

The present Distribution of the Mammalia. — In the follow-
ing pages some of the principal facts in the geographical range
of the orders, families, and many of the genera of Mammalia
will be given. It has been justly observed by Mr. Sclater
that the habitat of an animal is as much a part of its
definition as is its structure or external form. No systematic
account of the Mammalia would therefore be complete without
such geographical facts. But that branch of zoology which
is concerned with the past and present distribution of animals
is wider in scope than this. Zoogeography deals not only
with the actual facts in the range of animals, but with the
inferences as to past changes in the relations of land and sea
which the facts seem to indicate, and with speculations as to
the place of origin of the different groiips, of which more than
hints are sometimes given by their past and present distribution.
In addition to this, the earth can be mapped out into provinces
and regions which are definable by their animal inhabitants.
In the present volume, dealing only with the Mammalia, it will
be obviously impossible to enter fully into the entire subject
of zoogeography. All that will be attempted is a brief general
survey of the science so far as it can be illustrated by the
Mammalia. For fuller knowledge the reader is referred to the
treatises mentioned below.1

There are certain facts in the distribution of animals which
are commonplaces of knowledge, but which may be set forth
with definiteness. Everybody knows that an animal has a given
range : Elephants, for example, are found in India and certain
adjacent parts of Asia, and again in Africa ; the Ehinoceroses
have roughly the same range ; the Tiger is limited to Asia ; the

1 Wallace, The Geographical Distribution of Animals, 1876. Heilprin, Tlie
Distribution of Animals, Internal. Scientific Series, 1887. Beddard, A Text-book
of Zoogeography, Cambridge Natural Science Manuals, 1895. Lydekker, Geographi-
cal History of Mammals, Cambridge Geographical Series, 1896. W. L. and P. L.
Sclater, The Geography of Mammals, Kegan Paul and Co. 1899.

ii BARRIERS TO DISTRIBUTION 79

Jaguar to America, and so forth. The entire expanse of country
which is inhabited by an animal is called its area of distribution.
Such areas are larger or smaller. The Lion ranges over the whole
of Africa, a small part of India, and some neighbouring countries ;
on the other hand, the Insectivore Solenodon is limited to Cuba
and Hayti, a separate species to each. Among other groups of
animals are instances of an even more restricted range. There

O

are humming-birds confined to the slopes of a single mountain,
and fishes limited in their range to a single small lake.

A species may be found everywhere within the area of its
distribution, or it may be confined to a number of limited tracts
within that area. In this case it is usual to speak of " stations."
In such cases the species in question is generally suited to some
particular kind of environment. Thus the Otter and other
aquatic mammals will only be found where there is water; and
intervening tracts of waterless country will contain no Otters.
Goats and Chamois live only upon mountains ; the intervening
plains are destitute of them. This discontinuity of distribution
within the area is very general. But a discontinuity of area is
also seen — not so commonly however ; and, indeed, when it does
occur, it is a matter of a genus and not of a species. Thus the
Tapir is found in the East Indies on the one hand and in South
and Central America on the other, being absent in the inter-
mediate tracts.

It is clear that tracts of country eminently suitable for the
housing of a particular mammal do not always possess that kind,
or even an allied form. Africa, for example, possesses no arboreal
Anteaters ; there are no Anteaters at all (of the order Edentata)
in Australia, though there are plenty of ants for them to feed upon,
and tropical conditions of climate prevail. But as in these cases
the inference may be denied on the grounds that no experiments
exist to prove or to disprove the assertion, the matter may be
better emphasised by such cases as the introduction of the Eabbit
into Australia, and various mammals, such as Goats, into oceanic
islands. The plague caused by the former is a matter of notoriety.
But although climate and conditions and animal inhabitants do
not march accurately together, there is certainly some connexion
between temperature and the range of animals. Mr. Lydekker
writes on this point as follows : " The llama -like animals, re-
spectively known as vicunas and guanacos, are met with in

8O THE SEA AS A BARRIER CHAP.

company on the highlands of the Cordillera in Peru and Ecuador,
but as we go farther south the latter are found on the plains of
southern Argentina and Patagonia, as well as on the island of
Tierra del Fuego at the sea level. Here then is a clear proof
of the intimate connexion existing between temperature and
station ; the guanaco being an animal which can only live in
cold or temperate climates, finds suitable conditions for its
existence in tropical latitudes solely at a height of so many
thousands of feet, although farther south it is able to thrive at
the sea level." This, however, cannot be pushed too far — the
world cannot be mapped out into areas bounded by parallels of
temperature as was once attempted — since there are plenty of
cases like that of the Tiger, which is as much at home in a
tropical jungle as on the icy plains of Northern Asia.

Seeing that there are in many cases no climatic barriers to
the spreading of a given race of animals over a larger area of
distribution than it actually occupies, it becomes important to
inquire why there are so many cases of restriction in range.

It is possible to see, at any rate, three causes which are
responsible for a large number of such cases. In the first place,
a given species of animal must have originated at a certain spot ;
its multiplication in individuals must always be a slow matter,
since enemies, and untoward events generally, would conspire
to check the natural multiplication by geometrical progression.
A long time might therefore elapse before the species greatly
extended its range. A restricted distribution may therefore,
in some cases, mean a modern race. In the second place,
there are definite physical barriers which check the migra-
tion of species. The terrestrial Mammalia cannot cross wide
arms of the sea ; that they can and do swim for considerable
distances has been, proved in several instances ; but, as has
been pointed out, it is unlikely that a purely terrestrial mam-
mal would voluntarily swim out into an unknown sea. And
then if it did, and successfully reached the opposite side,
nothing would happen unless it were a pregnant female ; or, if
not pregnant, till a male swam very soon afterwards in exactly the
same direction. Many travellers have told of floating islands,
formed of torn-up trees and brushwood, which have been seen
at the mouths of large rivers, with animal passengers upon them.
These are, however, so much at the mercy of currents and storms,

ii MAMMALS OF ARCTIC REGION 8 1

that but little reliance can be placed on them as a means of
transit ; besides, here again, two individuals, or a pregnant female,
would be required to effect a settlement on a foreign shore.
The existence of oceanic islands is often urged as a proof of this
inability to cross tracts of sea ; even those which are com-
paratively near an extensive continent, such as, for example,
Fernando Noronha in the Atlantic, are destitute of mammals
(except, indeed, the ubiquitous Mouse, which is believed to have
been carried there, often in company with the equally widely-
spread Hat, in ships). This argument, however, is not so conclusive
as might appear ; it doubtless is in the case of far-distant islands.
But the size of the islands has to be taken into account. For
there are islands, such as the Galapagos, or, to take a less con-
tested instance, some of the islands of the Malagasy Archipelago,
undoubtedly continental, which have an exceedingly reduced
number of mammals. An area of a certain size seems to be a
necessity.

The converse of this is in many cases easy to show, that is,
the wide range of animals when there are no marine barriers to
stop their spreading. John Hunter, the celebrated anatomist
and surgeon (not often quoted, however, as an authority upon
geographical distribution), observes : •" It is a curious circum-
stance in the natural history of animals to find most of the
northern animals the same both on the continent of America and
what is called the Old World, while those of the warmer parts
of both continents are not so. Thus we find the bear, fox, wolf,

elk, reindeer, ptarmigan, etc., in the northern parts of both

The reason why the same animals are to be found in the northern
parts is the nearness of the two continents. They are so near
as to be within the power of accident to bring the animals,
especially the large ones, from one continent to the other either
on the ice or even by water. But the continents diverging from
each other southward, so as to be at a very considerable distance
from each other even beyond the flight of birds, is the reason
why the quadrupeds are not the same."

There is no doubt, in fact, that the ocean is the most in-
superable of all barriers to the dispersal of mammals. In a less
degree mountain ranges and deserts are also barriers. The
Desert of Sahara is a striking instance to the point ; it separates
two exceedingly different faunas.

VOL. X G

82 RANGE OF MARSUPIALS CHAP.

A third cause of more or less limited range is the barrier
due to competition. If the ground is already taken up, there is
no room for new immigrants. There is obviously a limit to the
number of Antelopes or Deer that can graze upon a given tract
of grassy plain. These two groups of Ungulates illustrate the
matter well: the Antelopes are African and Indian, especially
the former, while Africa has no Deer at all ; America, on
the other hand, has plenty of Deer but no Antelopes, save the
Prong-horn. The more nearly akin the two species or groups
of species are, the fiercer will be the competition ; for a near
kinship will at least often imply similar habits, the need for
similar food, and other likenesses which will prevent both from
successfully occupying the same tract of country. The remark-
able fauna of Australia is believed to afford an example of this.
In that country the prevalent inhabitants are the Marsupials.
The Monotremes are found there also, and nowhere else save in
New Guinea and Tasmania. The remaining mammals are in-
conspicuous ; they embrace a few Kodents and Bats, and the
doubtfully indigenous Dingo -dog. Now the Marsupials are
fitted to every variety of life. We have the grazing Kangaroos
and Wallabies, the burrowing Wombats, the arboreal Phalangers,
and the carnivorous Dasyures. In the second place, it is an
unquestioned fact that the Marsupials are an older race than
are the existing Eutherian mammals ; they were the dominant
mammals during the Secondary epoch. At that time they were
more widely distributed than at present. In most parts of the
world they are now absent, since they have been successfully
ousted by the more highly organised groups of Eutheria. But at
that period, when the higher Eutheria were in the ascendant,
Australia and the islands to the north became cut off from Asia,
and thus became freed from inroads of Eutheria, which were
partly prevented by the physical barrier of the sea from effect-
ing a settlement, and partly perhaps prevented owing to the
ground being already taken up by the Marsupials. Likeness
of habit gave the older inhabitants victory in the struggle for
existence.

The general statements that have been here made are in
accord with current opinion upon the factors of geographical
distribution. But the past range of animals appears to be
less consonant with the received views. In the Tertiary

ii DISTRIBUTION IN THE PAST 83

period, groups of animals had often a far wider range than
at present. To-day the Rhinoceroses are limited to Asia and
Africa, and to quite limited parts of the former continent.
In the past, these animals were abundant in Europe and
North America. Wild Horses now have a range which is not
widely different from that of the Rhinoceroses, save that they
extend into the more northern regions of Asia. Their remains
are abundant both in North and South America. The Hippo-
potamus, now confined to Africa, once ranged over Europe,
Madagascar, and India. There were plenty of American and
European Lemurs. Elephants were nearly world-wide in their
range ; and, in short, restricted distribution seems to be on the
whole a characteristic of animals of the present day.

These statements, however, though perfectly true, must not
lead to erroneous inferences. It is rather impressed upon the
reader, in books which contain sections dealing with geographical
distribution, that animals on the whole occupy more restricted
areas at present than in the past. There are, however, plenty
of examples of groups of extinct creatures which had, so far as
we know, quite a restricted range. Thus the Toxodonts were
purely South American, as were the Glyptodonts and some other
forms. And, on the other hand, the Cervidae of to-day are as
widely, if not more widely, distributed than at any other time.
The Hares and Rabbits are now nearly universal in range ; the
Cats almost so. We meet with Bovidae, even excluding the Sheep
and Goats, in all the four quarters of the globe, excluding only
South America and, of course, Australia. The Camelidae are still
common to both the Old and the New Worlds.

During certain periods of the Tertiary epoch it is true that
there was more similarity between Europe and North America
than there is at present. It would have been quite necessary to
unite both into a Holarctic area, such as is now insisted upon by
many ; but the reasons for this union would then have been
stronger. The fact is, however, that the closer resemblances were
due to the larger number of families of animals which existed
then than now ; these have decayed away from both continents,
and allowed the unlikenesses between the mammalian fauna of
both to become evident. But the likenesses which still sur-
vive have led many to associate the two regions closely together.

So far as the history of a genus or family or larger division

84 DR. BLANFORD'S REALMS CHAP.

can be traced, it results as a conclusion that from a given area of
origin the group in question migrated in all directions where
possible to a varying degree ; it then died out in intervening
tracts, or was left only in a certain part of its former and more
extensive area of range.

Zoological Regions. — Seeing that each species of animal has
its own definite range, it is clear that the earth's surface can be
apportioned into divisions which are characterised by their
animal inhabitants. We shall divide the earth into realms,
which are the largest divisions ; then into regions ; and
finally into subregions. It must be borne in mind that the
various groups of the animal kingdom are of different ages,
geologically speaking, and have therefore had less or more time,
as the case may be, to settle down into their present distribution,
and that different animals differ greatly in their rate of multi-
plication, their power of migration, and their susceptibility
to the effectiveness of various natural and other barriers to
distribution. It is not, therefore, possible to divide the
world into realms and regions which shall express the facts of
distribution of the entire animal kingdom. Such divisions,
which are common in text-books of zoology having but a small
section devoted to zoogeography, are at best mere approximations
and averages ; no good is gained by taking such a comprehen-
sive view of the matter, as the essential object of subdividing
the earth's surface is thereby lost sight of. The zoogeographical
division of the earth which will be adopted here is that origin-
ally recommended by Dr. Blanford, and now accepted by a
number of authorities. There are three " realms," to which a
fourth may perhaps be added — though on negative grounds, and
merely for the purpose of emphasising the parts of the world to
which mammals have not gained access. The realms are again
divisible into regions, at least in the case of one of them, and
the regions may be again separated into more or less distinct
subregions or provinces. The three primary divisions or realms
which contain mammals are the Notogaean, including Australia
and certain islands to the north of it ; the Neogaean, or the
South American continent and Central America; the Arcto-
gaean, including the continents of North America, Europe, Asia,
and Africa, together with the adjacent islands, such as the
West Indies, East Indies (exclusive of those which fall within

n NOTOGAEA, NEOGAEA, AND ARCTOGAEA 85

the realm of Notogaea), and Madagascar ; and finally, the realm
of Antarctogaea or Atheriogaea, which embraces New Zealand,
the Antarctic continent, and a series of islands such as South
Georgia and Kerguelen, and possibly even the extreme south of
Patagonia. This latter quarter of the globe will need no further
reference, as it has no truly indigenous terrestrial mammalian
inhabitants. We cannot include the Bats in this statement, as
their distribution is due to different powers of extending their
range, and to different barriers from those which govern the
range of other groups of mammals.

(1) Notogaea.1 This realm is characterised by the exclusive
possession of the Monotremes : — that is to say, one of the two
primary divisions of the Mammalia is absolutely restricted to
this area. It contains, moreover, the vast majority of the
Marsupials. Further, the realm of Notogaea is to be dis-
tinguished by the entire absence of the higher mammals, with
the exception of a few small Eodents. (The Bats are ignored
for the reasons stated, and the Dingo is believed to have been an
importation.) It cannot be disputed that this is a very distinctly-
marked area of the earth's surface.

(2) Neogaea. The continent of South America has no
Monotremes and only a few Marsupials, all of which, with the
exception of Caenolestes, belong to the Polyprotodont division of
that order, and to a peculiar family, Didelphyidae. The recent
discovery of other fossil Marsupials, however, to some extent
favours Huxley's view that Neogaea and Notogaea form one
realm as opposed to the rest of the world. Besides this,
Neogaea possesses the Edentata, which are found nowhere else ; —
that is, the division of the Edentata to which the name is
now restricted by some authorities. It is also characterised by
the nearly entire absence of the important order of Insectivora ;
and, as minor marks of distinction, by the absence of Antelopes,
Oxen and Sheep, of the Ichneumon tribe, of Horses, and of Lemurs.
It has the exclusive possession of the Hapalidae and Cebidae,
and of several families of Eodents.

(3) Arctogaea. This vast realm is clearly capable of sub-
division into four regions, which will be considered in detail
later. In the meantime the points of likeness between these sub-
divisions is more marked than are either the resemblances or the

1 This term is sometimes used in a wider sense ; cf. vol. viii. p. 74.

86 RANGE OF EDENTATES CHAP.

differences of any one of them to either of the two realms which
have just been defined. The two realms that have been discussed
retain their distinctness from each other and from Arctogaea for
a considerable way back into the Tertiary period. It is not until
we reach very early Tertiary times that Edentates are met with
in North America ; and then it cannot be regarded as absolutely
settled that the Ganodonta are really the forerunners of the
Armadillos, Sloths, etc. Nor do we find Marsupials in Europe
until far back in time, and at a corresponding period in North
America. Indeed the fauna of South America in late Tertiary
times was even more distinct than it is now ; for then we had
confined to that region the Toxodonts, Glyptodonts, Macvauchenica,
and other forms, while in Australia there were still Marsupials.
In late Tertiary times Europe and India were by no means
so distinct from Africa as they are to-day. North America does
not resemble the Old World quite so much as the subdivisions
of the Old World resemble each other ; but, as will be pointed
out later, there are and were very substantial agreements. The
Elephants, Ehinoceroses, Giraffe, Hippopotamus, Orycteropus, are
now distinctively African or Indian animals ; but all these
genera, or at least families (in the case of the Giraffe), have
occurred in Europe during quite recent times. Lycaon indeed,
now confined to Africa, is thought to have had a European
origin from its occurrence in caves there. The Hyaena and the
Lion, certain members of the Horse tribe, Apes, and other
animals, were also but are not now European.

India again, and the Oriental region generally, once possessed
the Hippopotamus, the Chimpanzee, Giraffidae, the Antelopes,
Cobus, Hippotragus, Strepsiceros, and Orias, which are now purely
African animals. It shares at present with the Ethiopian region
the Catarhines, including the Anthropoid Apes, the Lemurs, Tra-
gulina (the genus Dorcatkerium is also known from fossils in
India), Manis, Hyaena, the Cheetah, Elephant, Ehinoceros, and the
Eatel. There is, in fact, no order of mammals which is now
absent from one of these three regions though present in the
others, save the Lemurs, and they occurred in past times in
Europe. The Tapir of India is known fossil in Europe, and the
latter continent had its Monkeys and even Anthropoids. On the
other hand, North America is more distinct. It has no Lemurs,
Apes, Elephants, Ehinoceroses, Tapirs, Old World Edentates (Effo-

ii MR. SCLATER'S REGIONS 87

dientia), Viverridae, Horses, or Antelopes, excepting Antilocapra,
a type of a separate division of Bovidae. But since several of
these groups have been represented in recent times, no primary
line of division can be profitably drawn.

Arctogaea as a whole may be characterised by both negative
and positive characters. As negative features may be mentioned ;
— the entire absence of Edentates (Necrodasypus of Pilhol is
rather doubtful, see p. 164, n.), though a few crept up into the
Nearctic region from Neogaea during past times ; and of Hapalidae,
Cebidae, and Marsupials, except an Opossum in North America.
This realm has, on the other hand, all the Lemurs, all the
Insectivores with the exception of the West Indian Solenodon,
all the Proboscidea, Rhinoceroses, Horses, Deer, Antelopes, the last
group including the Oxen and a variety of other important
families. It is in fact the headquarters of all the Eutheria with
the exception of the Edentata and Marsupials.

The subdivisions of this realm have been variously effected.
The classical subdivisions are of course those of Mr. Sclater, who
would recognise (1) the Nearctic, North America ; (2) the Palae-
arctic, including Europe, Northern Asia, and Japan ; (3) the
Oriental, including Asia south of the Himalayas and the islands
of the Malay Archipelago as far east as the Australian region ;
and (4.) the Ethiopian, i.e. tropical Africa and Madagascar. Some
would alter this by uniting America and the north of the Old
World into a Holarctic region, separating off the southern parts
of the North American continent into a Sonoran region. To
some, the claims of Madagascar to form a separate region are
convincing. To distinguish the boundaries of the several regions
is a difficult task ; they dovetail into each other on the frontiers
with the complex curves of a puzzle-map. The difficulty has
been grappled with by the suggestion of intermediate transitional
•areas ; but this proceeding really doubles the difficulty, for there
are then two frontiers to delimit in each case instead of
only one. The animal inhabitants must be expected to mingle
somewhat at the lines of junction of one region with another.

The Sonoran region does not appear to us to have great claims
to recognition. It shows a mingling of southern with northern
forms exactly as might be expected. An Armadillo and Didelphys
have, as it is believed, invaded it from the Neogaeic realm ; it
possesses also the South American genera, Dicotyles, Nasua, Cone-

88 NEARCTIC AND ORIENTAL REGIONS CHAP.

patus, Sigmodon. On the other hand, the Sonoran genera Antilo-
capra, Cynomys, Procyon, and the Insectivora Blarina and Scapanus,
extend further north. Peculiar to this region are only six genera
of Eodents, which seems an insufficient reason for raising the
Sonoran province to the dignity of a region. Considered from the
point of view of numbers of peculiar forms, the Thibetan subregion
has more claims to distinction as a region ; for confined to that area
we have the genera Nectogale, Aeluropus, Eupetaurus, Pantholops,
Budorcas ; while by slightly extending its limits, a number of
other peculiar forms might be added. Madagascar has distinctly
more claims to regional division. Absolutely confined to it are
eleven of the seventeen existing genera of Lemurs, the family
Centetidae among the Insectivora, which contains seven genera,
and another recently discovered and peculiar genus, Geogale ; it
has six peculiar genera of Viverridae ; it has five peculiar genera
of Eodents. In addition to this it is negatively characterised
by the absence of the following typical African animals, Felidae,
Proboscidea, Ehinocerotidae, Equidae, Monkeys, etc. It seems
to be impossible to avoid allowing the rank of a region to this
part of the world.

In separating the Nearctic from the Palaearctic region, stress
must be laid rather upon the absence of Asiatic and European forms
from North America than upon the existence in the northern half
of the New World of many peculiar forms. Peculiar to the Nearctic
are the Goat genus Haploceros, the Eodents Erethizon, Zapus, and
the family Haplodontidae. The Mole genus Condylura is also
restricted to this part of the New World. Even so it has more
peculiar forms than the Sonoran. If we add to this the absence
of Horses, Antelopes except Antilocapra, Pigs, Hyaenas, etc., there
are strong grounds for retaining this division. It must be
agreed, however, that it comes rather nearer to the Eurasian
district than the latter does to the Oriental.

The Oriental region has many characteristic animals. It has
among the Anthropoid Apes the Orangs and Gibbons; of Old World
Apes it has confined to its own area the genera Semnopithecus
and Nasalis. Of Lemurs there are Loris and Nycticebus, and
Tarsius, representing a family of that order, or even a sub-order.
The Galeopithecidae are entirely Malayan. There are many
Eodent, Carnivorous, and Insectivorous genera; the Ehinoceroses
and the Elephant of this region differ from those of Africa.

ii THE ETHIOPIAN REGION 89

Tragulus concludes a sample from a very rich list of peculiar
forms.

The Ethiopian region has also its Anthropoids, the Gorilla and
the Chimpanzee, but they belong to genera or a genus different
from those which include the Oriental forms. There are five
peculiar genera of Cercopithecidae. The Lemurs restricted to
this region are Gcdago, Perodicticus and Arctocebus. The peculiar
Insectivorous families Macroscelidae and Chrysochloridae are
only found here, besides many other peculiar genera. Africa is
especially the home of Antelopes, and the Giraffe is not found
now outside its borders. The Elephant and the Rhinoceroses are
of different species from those of India. There are many peculiar
Kodents and Ungulates.

CHAPTER III

THE relationship of Mammals to Vertebrates lying below them
in the scale, their origin in fact, is a much-debated question,
with many attempted solutions. To enter into this large
question in detail would involve a great deal of useless state-
ment of arguments founded upon misleading or upon quite in-
accurate " facts." It will perhaps be sufficient if we reflect here
the current view most in vogue at the present, i.e. that which
would refer the Mammalia to reptiles belonging to the extinct
Permian and Triassic group of the Theromorpha (also called
Anomodontia). These have been explored lately to a very large
extent, and chiefly by Professor Seeley.1 The very fact that a
genus Tritylodon, only known by the forepart of the skull, has
been called Mammalian and Anomodont by various authors,
shows at least the difficulty of differentiating the two groups when
the material for study is imperfect. As a matter of fact
these Theromorpha are without doubt reptiles ; they show, for
example, a lower jaw formed out of several distinct pieces, of
which the articular articulates with a fixed quadrate on the
skull. They possess the characteristic reptilian bones, the
" transverse," the pre- and post-frontals, and there are various
other points of structure which leave no room for doubt as to
their truly reptilian nature. There are, however, numerous
indications of an evolution in the mammalian direction in all
parts of the skeleton, to the more important of which some
reference will be made here. It may be as well to clear the

1 A series of papers in the Phil. Trans, for 1888-96, of which a useful abstract
by Professor Osborn was published in the American Naturalist, 1898, p. 309 ; see
also Canibr. Nat. Hist. viii. 1901, p. 303.

CHAP, in THE THEROMORPHA pi

ground by mentioning the fact that among the Theromorpha
four distinct types of reptiles are included, which are considered
to form four orders, i.e. the Pareiasauri, -the Theriodontia, the
Anomodontia (Dicynodontia), and the Placodontia.

The first of these divisions includes what seem to be basal
forms. These reptiles show numerous points of likeness to the
Amphibian Labyrinthodonts.1 On the other hand the third divi-
sion, that of the Dicynodontia, are highly -specialised Thero-
morpha, from which no further evolution would appear to have
been possible. Thus the dentition was either completely lost,
or reduced to tusks as in Dicynodon. We need not therefore
concern ourselves in the present volume with these Anomo-
donts. It is with the Theriodonts that our business lies.
The very name, be it observed, is aptly chosen on the hypo-
thesis to be explained here ; but it is not only in the teeth
that these reptiles show likenesses to the Theria or Mammals,
but in almost every feature of their organisation. Unlike other
reptiles, the Theromorpha in general were lifted comparatively
high above the ground on legs of fair length and of mam-
malian relationship in the position of the segments of the
limbs. The typical reptile grovels upon the earth with legs
sprawling out, as indeed the very name suggests. One bar to
the Theriodonts being on the direct line of mammalian ancestry
has been urged as a preliminary difficulty, and that is their
large size. The earliest undoubted mammals were small
creatures, comparable to a Eat or a Mouse in size ; whereas a good-
sized Bear or a Wolf is a better standard of size for some of the
best-known genera of Theriodonts. It has, however, been quite
permissibly suggested that living in company with these large
Theriodonts were less obtrusive genera, from which the mammals
might have sprung. It is so familiar a fact that a given group
of animals generally contains giants, dwarfs, and members of
intermediate size, that this suggestion may almost be accepted
as a fact. It need at least present no difficulties to us in our
comparisons.

The most salient " mammalian " feature of the Theriodonts
is the heterodonty of the teeth, the pattern of the " molars," and
the limited number which constitute the series. The fact, too,
that they are limited to the dentary bones below and to the

1 Cf. vol. viii. p. 82.

92 TEETH OF THEROMORPHA CHAP.

maxillae and the premaxillae above, is a sine qua non for
mammalian comparison. In the more basal Theromorpha the
teeth are not so limited in position. Finally, to complete the
remarkable mammalian resemblance of the teeth of these reptiles,
it must be mentioned that in Tritylodon and Diademodon the
roots of the molars, as we may fairly term them, though not
actually divided after the mammalian fashion, were deeply
marked by a groove, which suggests an incipient division or a
fusion of two distinct roots. Some of these facts of structure
may now be considered in further detail. As to the incisors
and canines, it is sufficient to say that the numbers of the former,
and the shape of the latter, are in perfect consonance with a
derivation of the Mammalia from this group. The molar series
can be divided into premolars and molars, at least in so far as
regards their shape ; for the anterior teeth are often smaller and
less complicated than those which follow, as is often the case with
the two series in mammals. The molar series also consist of teeth
in close apposition to each other and separated from the canines
by a diastema, which is a character of mammalian teeth. The
fact that in the reptile Cynognathus and the mammal Myr-
mecobiiis there are nine of these molar teeth in each half of each
jaw is perhaps not a point upon which it is desirable to dwell
with too much weight ; but the general fact that the molars are
further reduced in some genera of Theriodontia than in that
which has been mentioned, is clearly a matter of significance
when the ancestry of the mammals is under consideration.

The most interesting fact about the molar series in the
Theriodontia is that we meet with the two types of molars that
occur in the mammals. Cynognathus and other genera have
molars which consist of a main cusp, and of one cusp before and
one after the main cusp ; in fact these teeth are triconodont as
in certain early mammals, a state of affairs which is believed
by the " trituberculists " (see p. 56) to have preceded the
tritubercular tooth. There are also " multitubercular " teeth,
especially well developed in Tritylodon, where they exactly
resemble those of certain Multituberculata, and whose structure
originally led to the placing of Tritylodon among the mammals of
that group. If there is any question about the mammalian nature
of this fossil, there remain several other Theriodontia in which
the multituberculism is well marked. It is so in Trirhachodon

SKULL OF CYNOGNATHUS

and in Diademodon for instance. This incidentally lends some
support to the idea that the Mammalia have been evolved from
two sources, a way of looking at the origin of the group that
will coincide with the views of some authors like the late Dr.
Mivart, and will at the same time reconcile the trituberculists
and the multituberculists. For we should then assume that
the Eutheria and Triconodontia had originated from some such
form as Cynognathus ; and the Multituberculata and the ex-
isting Monotremes from some form like Diademodon. It is not
of great use to point out that Diademodon is really pf the tri-
tuberculate pattern, because in its molars, though multituberculate,
the trituberculate main cones can be recognised ; for that state
of affairs could just as well have been brought about by a
reduction from the multituberculate type. The skull of these
Theriodonts shows some well-marked approximations to the
mammalian type. There is in the first place a commencing
consolidation and reduction of the individual bones, which is so
distinguishing a feature of the mammalian skull as opposed to the
skull of lower vertebrates. In Cynognathus the postorbital is
fused with the jugal, and the supratemporal with the squamosal,
forming apparently one bone. In the lower jaw the splenial is
often reduced to the thinness of paper, thus indicating a com-
mencing disappearance. In many Theromorpha the squamosal
shares largely in the formation of the articular facet for the
lower jaw, obviously an important mammalian characteristic ;
this is brought about by the reduction of the quadrate, which
latter bone, moreover, acquires in certain particulars the appear-
ance of the mammalian malleus, with which it is, according
to many, homologous. But this subject has been already dealt
with on page 26. A very pronounced likeness to the mam-
malian skull is that there are two occipital condyles. That
this has been brought about by the further development of
a tripartite condyle such as occurs in tortoises, and that by
the suppression of the basi- occipital part, does not affect the
resemblance to the mammalian skull ; in fact it explains the
origin of two condyles from the typical reptilian single con-
dyle, and disposes of the necessity for believing, with Huxley
and others, the Amphibia to be on the main line of mam-
malian evolution on account of their two condyles. The
general aspect of the skull in Cynognathus has been com-

94 SHOULDER GIRDLE OF CYNOGNATHUS CHAP.

pared to that " of Thylacinus or Dissacus." No one can ex-
amine the actual sketches of the skull of that Theriodont
without endorsing that opinion. As a curious detailed point
of likeness to certain Mammalia may be mentioned "a small
descending process of the malar bone, which may be a diminu-
tive representative of the descending element of the malar
seen in Motherium, Nototlierium, Diprotodon, Macropus, certain
Edentata, such as Glyptodon, Megatherium, Mylodon, Brodypus,
but unparalleled so far as I am aware in fossil reptiles." (Osborn.)
The zoologist cannot help being impressed with the significance
of small details of similarity, which do not seem to be due in any
way to surrounding conditions of life, and thus referable to mere
convergence, like the fish-like form of Whales and Seals.

The rest of the skeleton of the Theriodontia is by no means
so well known as the skull and teeth. But from what is known,
other mammalian characters can be pointed out. Perhaps the
most striking mammalian feature is to be found in the scapula
of Cynoynatlius. It is in this creature somewhat narrow and
elongated ; but it has a well-marked spine, ending in a hooked
acromion. Now it is to be noted in support, so far, of the
diphyletic origin of mammals, that in the Monotreme, as in
Whales indeed, the spine forms the anterior border of the scapula,
and is coincident with it, there being thus no prescapula at
all in the Monotreme, and only a trace of it in certain Whales.1
Whether the multituberculate Tritylodon or Diademodon had
a scapula after the Monotreme pattern is not known ; but it is
clear that the scapula of the triconodont Cynoynatlius is quite after
the pattern of the Eutherian scapula. Furthermore, Professor
Seeley is of opinion that the coracoid was relatively small, and
indeed smaller than the same bone in Edentates, and a fortiori
than in Monotremes. Another fact of structure which points
also, possibly, in the direction of a diphyletic origin for the
Mammalia, is the double-headed ribs of Cynoynathus. As is well
known, the ribs of the Monotremata have only the central head,
the capitulum.

As a general mark of affinity with mammals the reduction of

the intercentra in ,Cynognathus may be noted, and also the

existence of a small though perfectly obvious obturator-foramen,

separating the pubis from the ischium. There are further details

1 It may be necessary to exclude the Whales from the comparison.

in THE ORIGIN OF MAMMALS 95

which tend in the same direction. And we shall probably not
go far wrong in the present state of our knowledge if we
assign the origin of the mammals to some type which would be
included in the order Theriodontia or at least in the sub-class
Theromorpha.

CHAPTER IV

THE DAWN OF MAMMALIAN LIFE

THE animals that we considered in the last chapter, though show-
ing certain unmistakable likenesses to the mammals, are never-
theless unquestionably not mammals but reptiles. In the
Triassic strata, however, we first meet with the remains of
undoubted mammals. The Mammalia first appeared upon the
earth in a tentative and hesitating way : they had not cast off
many of the characters of their supposed reptilian forefathers ;
they shrank from observation and destruction by their small size,
and apparently, so far at any rate as their teeth afford a clue, by
an omnivorous diet. The world abounded at that period in large
and carnivorous reptiles, which may indeed have been the
principal enemies with which the first mammals had to cope.
These early mammals lingered on to so late a period as the
Eocene ; but the majority of the genera were Triassic, Jurassic,
and Cretaceous. Certain of the primitive mammalian forms have
been referred to the Marsupials, and their resemblances to the
Monotremata have also been pointed out. The current view of
the present time, however, is that they form a special order,
which may possibly have embraced the ancestors of both
Marsupials and Monotremes ; for it is reasonable to explain in this
way the combination of characters of these two orders which they
present. For this group the rfame Allotheria has been proposed
by Marsh, and Multituberculata by Cope ; the latter term is the
less suitable, in that the Monotremata (Ornithorhynchus) are also
" multituberculate." The group is known in a very imperfect
fashion. The remains are but few and fragmentary ; and for the
most part we have only a few teeth to speculate upon. This is
natural enough, for the harder teeth might easily be supposed to

CHAP, iv FOSSIL JAWS 97

have resisted the decay which would more readily affect the
softer bones. Where there are bones it is frequently the lower
jaw alone which has been preserved for us — a bone which has also
been preserved in the case of some of the contemporary Marsupials.

It has been pointed out (from the observation of dead dogs
floating in canals) that the lower jaw is occasionally detached
from the carcase. It is the most readily separable part which
contains a skeleton. It may be, therefore, that the remains of
these early mammals, floating down some river to the sea, may
have lost their jaws while in the river, or at furthest in the
shallow waters of the sea, the rest of the carcase floating out to
a greater distance, and being finally entombed in the stomach
of some carnivorous fish, or in the mud at the bottom of a deep
ocean, which has never since seen the light.

The characters of this group are really more those of the
Monotremata than of the Marsupials. The undoubted likeness
which their molar teeth show to the temporary teeth of the
Platypus have already been commented upon. Like the Mono-
tremes the Allotheria appear to have possessed a large and
independent coracoid ; the evidence for this rests upon the
discovery of the lower end of a scapula of Camptomus, a Cretaceous
genus from North America upon which there is a distinct facet
for the articulation of what can have been nothing else than a
coracoid. On the other hand they differ from the Monotremata
by the presence of incisor teeth which were Rodent-like in form,
and not very different from those of certain Marsupials. This
point of difference cannot be regarded as of very first-rate im-
portance ; no one would relegate the Sloth and the Armadillo to
different orders on account of their tooth differences, which are
about on a par with those to which we have just referred. It
seems indeed likely that it will be ultimately necessary to rub
out the boundary line which now divides the Allotheria and the
Monotremata.

The Plagiaulacidae are unquestionably mammals, and they
are placed by most naturalists in this at present uncertain group
of Multituberculata, which will be retained here in deference to
the distinguished authorities who have instituted the group,
though there are but few characters by which it can be defined.
This family though appearing in the Trias, extends down in time
to the Eocene. The type-genus, that which has given its name to

VOL. x H

98 JURASSIC MAMMALS

the family, is Plagiaulax. As it is not Triassic, the consideration
of its characters will be deferred until later. Microlestes is a
Khaetic genus, known from rocks in Germany and England ;
but it is entirely based upon molar teeth. M. antiquus has a two-
rooted molar of an elongated form with a row of tubercles on either
side of a median groove, which traverses the long axis of the
tooth. To some extent the teeth of the ancient form resemble
those of Ornithorliynclms. Microlestes has been sometimes spoken
of as a Marsupial, but Mr. Tomes * has found that it does not
show one very universal character of the Marsupial teeth :
it has not those continuations of the dentinal tubes which
traverse the enamel in all Marsupials that have been examined
with the sole exception of the Wombat.

The rarity of the remains of mammals in these earliest rocks
of the Secondary epoch has been accounted for in another way
from that which has been suggested above. It may be that the
group Mammalia was not evolved in Europe at all, and that the
stray remains which have been found in that continent represent
the fragmentary remnants of a few scattered immigrants which
heralded the later invasion of more numerous genera during the
Jurassic period.

The Mammals of the Jurassic Period. — Some of the Allo-
theria or Multituberculata described in the last section occur in
the rocks of this early part of the Secondary epoch. They are
doubtful in position, as already stated ; some of them indeed,
as for instance Tritylodon and Dromatherium, are possibly not
mammals at all, while the remainder probably belong to a non-
existent order of mammals. Along with these dubious creatures
are the fragmentary remains of small animals which are not
merely mammals, but in all probability definitely Marsupials.
It is true that here again we have little beyond lower jaws
and teeth to deal with ; so that there may be less certainty
in referring them to the Marsupials than appears to be the
opinion of the majority of Palaeontologists.

Professor Osborn in fact considers that the Mesozoic mammals
consist of three groups : (1) The Multituberculata, including
the Bolodontidae, Stereognathidae, Plagiaulacidae, Polymasto-
dontidae, and possibly the Tritylodontidae (which, however, are
regarded by him and by others as more probably reptiles of the
1 Dental Anatomy, 5th ed. 1898, p. 304.

iv PLAGIAULAX 99

Theromorphous group). (2) The Triconodonta, which were
Marsupials, though in all probability with a complete succession
of teeth and with an allantoic placentation. This group will
include the genera Phascolotherium and Amphilestes, as well as
Triconodon and Spalacotherium. Finally we have (3) the Tritu-
berculata (or Insectivora Primitiva) with the genera Amphitherium,
Peramus, Amblotherium, Stylacodon, and Dryolestes.

We shall take these three groups in order. The Multituberculata
have already been to some extent defined, if such a word can be
used to express the summation of the very scanty information at
our disposal. Of this group, Plagiaulax is a genus which occurs in
the Purbeck beds ; it is only known by lower jaws implying an
animal of the size of a Eat or rather smaller. The jaws have in
fronb a large incisor which looks Rodent-like, and also like those of
the Diprotodont Marsupials ; but it is held that these teeth did not
grow from persistent pulps, and there is in any case no anterior
thickened coating of enamel. Canines are absent ; the diastema
is followed by four premolars increasing progressively in size and
possessing somewhat complicated grinding surfaces. These surfaces
are formed by several obliquely-set ridges. The succeeding teeth
are termed molars on account of their difference in structure, and
there are but two of them on each side. The molars are of a
pattern common in the Multituberculata ; the centre is hollowed,
and the raised rirn is beset by tubercles. Other Jurassic genera
of Multituberculates are Bolodon, Allodon, and Stereognathus.
All of these possess the same multituberculate molars.

Of the Triconodonta the type-genus is Triconodon. This genus
is better known than most Jurassic mammals, since both the upper
and the lower dentition have been described. It appears to have
possessed the typical Eutherian dentition of forty-four teeth, to
which a fourth molar is added in some species. The great differ-
ence between the molars and premolars argues a complete tooth-
change. The genus is American as well as European.

Spalacotherium has more molars, five or six.

Phascolotherium bucklandi, on the other hand, is a much
older type in the form of its teeth. There are, however, not so
many of them as in Amphitherium ; Phascolotherium has but two
premolars and five molars, making a total of forty-eight teeth.
The teeth are of the triconodont form, the three cusps being in
line, and the middle one the largest.

TOO STONESFIELD MAMMALS CHAP.

Amphilestes has teeth of the same pattern but has more of
them, the premolars and molars being respectively four and five.
All these animals had the lower jaw inflected. Whether they
are all Marsupials or not, it is clear that Phascolotherium and
Amphilestes should be united and placed away from Amphitherium
on account of the more primitive form of their teeth.

We next come to the Trituberculata.

Among the most celebrated of these remains are a few jaws
discovered in the Stonesfield slates near Oxford, and examined by
Buckland, Cuvier, and some of the most eminent naturalists of the
beginning of the last century. These jaws have been lately sub-
mitted to a careful re-examination by Mr. Goodrich,1 who has
increased our knowledge of the subject by exposing from the
rocky matrix in which the jaws lie fresh details of their structure ;
it is probable therefore that now all that there is to be learnt
from these specimens has been recorded.

Amphitherium prevostii was a creature about the size of a Eat.
Its jaw was first brought to Dean Buckland about the year 1814,
and described six years later. Buckland thought the jaw to be
that of an Opossum, an opinion in which Cuvier concurred. The
jaw, however, is marked by a groove running along its length, and
this groove was regarded by de Blainville as evidence of the com-
position of the jaw out of more than one element, which would
naturally lead to its being regarded as the jaw of a reptile.2 This
species and another named after Sir Eichard Owen have a dental
formula which, Like that of the Marsupials, is large as compared with
that of the Placental mammals ; it runs : I 4, C 1, Pm 5, M 6 — i.e.
64 teeth altogether. This is a larger number than we find in any
existing Marsupial. But as in Marsupials, and in certain Insectivora
also, the angle of the jaw is inflected. These teeth are of the
tritubercular pattern with a " heel." They are in fact closely like
those of the living Myrmecolius ; but not, it should be remarked,
unlike those of certain Insectivora.

The Mammals of the Cretaceous Period. — At one time
there was a totally inexplicable gap between the Jurassic and
the basal Eocene, a series of strata which occupy an enormous
expanse of time in the history of the earth having appeared to

1 "On the Fossil Mammalia from the Stonesfield Slate," Quart. Journ.
Sci. xxxv. 1894, p. 407.

2 This groove has been found in the existing Myrmecobius, see p. 154.

EARLY TERTIARY MAMMALS IOI

be devoid of mammalian remains. This gap, however, has been
filled up by the discovery of mammalian remains in the North
American Laramic formation, which seems to be clearly of Creta-
ceous age. Furthermore, it is held by some that the Purbeck beds
are more properly to be placed with the Cretaceous, which would
then necessitate the consideration under the present heading of
some of the types already dealt with ; and if, as is suggested in
the following section, the lowest so-called Eocene beds are really
referable to the Cretaceous, there is no lack of mammalian remains
in that period. And, moreover, it was in that case the Creta-
ceous period which witnessed the evolution of the existing orders
of Placental mammals. Otherwise the mammalian remains of the
Cretaceous agree with those of the Jurassic. We find remains
of the Multituberculata in fragments of Plagiaulacidae and
Polymastodontidae. Ptilodus is a genus which has two pre-
molars; and Meniscoessus is another multituberculate from the
same Laramic formation. The other detached fragments of
mammals are thought by Osborn to represent both Placentals
and Marsupials.

The Mammals of the Tertiary Period. — Unless the lowest
beds of the earliest Tertiary period, the Eocene, such as the
Torrejon of North America, should be in reality referred to the
Cretaceous, there is no evidence that the modern groups of
Mammalia existed before the present epoch of the earth's history.
It is probable, however, that the Eutheria as a group were Meso-
zoic. The fossil jaws that have been considered in the last chapter
may quite probably be primitive Eutherians, or even divisible, as
believed by Professor Osborn, into Marsupials and Insectivores.
In the Tertiary, however, apart from the question as to the
nature of the Puerco and Torrejon formations, and as to certain
South American strata whose fossil contents have been investi-
gated by Professor Ameghino, we find the first traces of mammals
definitely referable to existing orders, or to be distinctly com-
pared with existing orders. Since, however, representatives of
types which have obvious relationships to modern types appear
in considerable profusion in the very earliest strata of the
Eocene, it seems clear that much remains to be discovered in
beds earlier than these. Confining ourselves, however, to facts
and to comparisons which can be made on more than a few
lower jaws and scattered teeth, which is practically all that we

102 PRIMITIVE UNGULATA CHAP.

possess of earlier mammals, we must arrive at the general con-
clusion that two of the existing larger groups of the Eutherian,
non-Marsupial, mammals were differentiated at quite the begin-
ning of the Eocene, and were represented by forms from which
it is possible to derive at least the existing Carnivora, Insectivora,
Artiodactyla, and Perissodactyla. These were the Creodonta and
the Ungulate Condylarthra. In addition to these we may
enumerate as very early types the Lemuroidea, represented by
such forms as Indrodon in the New World, and (though later) by
Necrolemur, etc., in the Old World, and the Edentata, if we are
to allow as their ancestors the Ganodonta.

The early Eocene strata also contain representatives of at
least one order, the Amblypoda, which increased subsequently,
but has died out without descendants, unless we are to believe
with some that the Elephants are to be derived from these
Eocene " pachyderms." In later Eocene times the great majority
of the existing orders, and even subdivisions of orders, are to be
met with ; and there are in addition such totally extinct orders
as the Typotheria, Ancylopoda, and Tillodontia. Coupled with
this gradual specialisation in the orders of Eutherian mammals,
there is naturally a vast increase in the number of generic and
family types. This culminates perhaps in the Miocene, from
which time there has been a gradual decline in mammalian
variety, so that it is justly said that we live now in an epoch
which is impoverished of mammals. This gradual decay has
persisted until to-day, as is witnessed by the extinction of the
Ehytina and the Quagga, and the growing rarity of the White
Ehinoceros and the American Bison.

The early Eutherian stock consisted of small mammals
with small heads and slender, long tails. The limbs were
pentadactyle, ensheathed in claws or broader hoofs. The fore-
limbs may have been partly prehensile. The teeth were forty-
four, completely differentiated into incisors, canines, molars, and
premolars ; and there appears to have been a complete diphy-
odontism. The canines were not greatly enlarged, and no
diastema separated any of the teeth. The molars were bunodont
or of a more cutting pattern, with some five or six tubercles.
These animals were, moreover, very small-brained. This early
stock is represented by Creodont and Condylarthrous animals,
the exact boundaries between which are hardly marked in the

iv CENTRES OF RADIATION 1 03

very early types. Professor Osbom has argued that from this
early Eutherian stock there were two waves of progress, or, as he
expresses it, " two great centres of functional radiation." *

The first was largely ineffective, the second has produced all
the Eutherian orders of to-day. These two divisions are termed
by him " Mesoplacentalia " and " Cenoplacentalia." The first
division embraces the Amblypoda and their descendants the
Coryphodonts and Dinocerata, many of the Condylarthra, the
bulk of the Creodonts and the Tillodonts. These creatures
persisted for a time, but died out in the Miocene. They were
mainly distinguished by the smallness of their brain ; the great
specialisation of structure which they exhibit having left that
organ unaffected, and therefore tending in the long run to render
them unable to cope with changes in the inorganic and organic
world. The successful division of the primitive Eutheria com-
prises the groups which exist at the present day, and is not
connected directly with those small-brained Mesoplacentals ; it
has apparently originated, however, from the least specialised of
their ancestors. Professor Osborn thinks, moreover, that the Lemurs
and the Insectivores are persistent descendants of the earlier
wave of Eutherian life. It appears in fact as if Nature had
created the existing Ungulate, Unguiculate, and other types on a
defective plan, and, instead of mending them to suit more
modern requirements, had evolved an entirely new set of
similarly -organised types from some of the more ancient and
plastic forms remaining over. The Marsupials may be the only
group of the early wave remaining, and they have been able to
hold their own for the geological reason that Australia was
early cut off from communication with the rest of the world.
That they are disappearing seems to be shown by their gradual
diminution as we pass from Australia towards the continent of
Asia, through the islands of the Malay Archipelago. Com-
petition has here decimated them, as it may do in the remote
future in Australia.

It is often said, but with some looseness of statement, that
ancient quadrupeds are huger than their modern representatives.
This statement is partly true in fact, but largely wrong in
implication. For it suggests that — and the suggestion is often
expressed in books that are not authoritative — huge animals

1 Trans. New York Acad. Sci. xiii. 1894, p. 234.

104 SECULAR INCREASE IN SIZE CHAP, iv

have left a dwarfish offspring ; that there were giants of
old, and that there is a puny race to-day. As a matter of
fact, the study of the gradual evolution of the early Tertiary
Mammalia into their descendants of later times shows very
plainly the truth of this interesting generalisation : That the
primitive types were all small creatures, and that in those
instances where we can trace a pedigree, there was a gradual
increase in size up to a point where greater increase led to
extinction. "We point out later on a number of facts illus-
trating this matter in detail. It has been ascertained, for
instance, that the pedigree of the Horses, the Camels, the Rhino-
ceroses, and many other groups, commences with small forms
and culminates in large ones. It may be urged that such
animals as the Tapir are to-day smallish forms, and that related
to them in the past were the gigantic Titanotheres ; but in this
and similar cases it will be found that the extinct giants were
not in the direct line of pedigree, but represented side-branches
which waxed huge on their own account and then disappeared.

CHAPTER V

THE EXISTING OllDERS OF MAMMALS
PROTOTIJERIA — MONOTREMA TA

APART from those creatbres whose fragmentary remains have
been considered in the last chapter, and which belong to the
earliest of mammaliferous strata, the remains of Mammalia are
all referable to existing orders. In the pages which follow we
shall therefore deal with the actual representatives of living
families side by side with their extinct relatives. The existing
orders of Mammalia, together with those of their fossil allies, can be
plainly divided into two great subdivisions, or, as we shall term
them, sub-classes ; the Mammalia as a whole being termed a class
of the Vertebrata comparable with the class Eeptilia, etc. It
has been usual, owing to the initiative of Professor Huxley, to
divide the Mammalia into three divisions of primary importance.
We shall adduce reasons later for not accepting this mode of
division, but that which allows of only two primary divisions.
These two divisions are (1) Prototheria and (2) Eutheria.
Whether the Multituberculata, Trituberculata, and Tricono-
donta, considered in the last chapter, are really to be distributed
among these two sub-classes is a matter upon which it is possible
to form an opinion, but not to dogmatise. The Prototheria
stand at the base of the mammalian series, and present many
likenesses to the Sauropsida ; the Eutheria are the animals
which are most fully differentiated as mammals. We shall
commence with

SUB-CLASS I.— PROTOTHERIA.

To this group belongs the order Monotremata, and possibly
also the so-called Allotheria or Multituberculata. As, however,

106 PROTOTHERIA — MONOTREMATA CHAP.

the latter are only known from very fragmentary remains,
which are not sufficient to determine the systematic position of
the animals of which they are fragments, I have not thought
it worth while to attempt a serious definition of the order
Multituberculata. I have introduced a short account of the
principal facts which are known concerning the creatures
grouped together under this name into the historical sketch of
the progress of mammalian life in Chapter IV. As to the
Monotremata, there is no question that they are entitled to rank
in a group equivalent to that including all other mammals of
which we have sufficient knowledge to construct a classificatory
scheme. There have been, indeed, naturalists, such as Meckel,
who would altogether deny the mammalian rank of these
creatures.

The Monotremata or Ornithodelphia may be thus defined : —

Mammalia with no teats, but with a temporary pouch in
which the young are hatched, or to which they are transferred
after hatching, and into which open the ducts of the mammary
glands. An anterior abdominal vein, or at least the membrane
supporting it, persists throughout the abdominal cavity. Heart
with an incomplete and largely fleshy right auriculo-ventricular
valve. Brain without a corpus callosum. Shoulder girdle with
a large coracoid reaching the sternum ; clavicles and an inter-
clavicle present. There are " marsupial " or epipubic bones
attached to the pelvis. Vertebrae with no epiphyses for the
most part. Ribs with only capitulum and no tuberculum.
Mammary glands of the sudoriparous and not the sebaceous
type of epidermic gland.1 Oviparous, with a large-yolked and
meroblastic ovum, enclosed within a follicle of two rows of cells.

To call these animals Mammalia is of course an abuse of the
meaning of that word in one sense, but it is not in another ;
since the pouch of these Monotremes is, as has been explained
elsewhere (p. 16), the real equivalent of a teat, and not of the
pouch of the Marsupials.

The most salient characteristic of this group of mammals in
the estimation of their position in the vertebrate series is not
so much the fact that they are oviparous as that the eggs are
large-yolked, and develop therefore, so far as regards their early
stages, after the fashion of the egg of a reptile. The laying
1 Gegenbaur, Zv,r Kenntniss der Mammarorgane der Monotremen, Leipzig, 1886.

v EGGS OF PLATYPUS IO/

of eggs, or at least ovoviviparity, would follow from the struc-
ture of the egg, since the abundance of yolk would do away
with the necessity for a placenta. That the eggs had this
Saurian characteristic was first definitely made known by Pro-
fessor Poulton l for Ornithorhynchus, and his results were con-
firmed later for Echidna? The structure of the eggs has,
however, already been dealt with on p. 72. The fact that
these animals lay eggs appears to have been known for a very
long time, though rediscovered so lately as 1884 by Mr.
Caldwell.3 In connexion with the structure of the ova, the
ovaries themselves and the oviducts are built upon the Saurop-
sidan plan. In the male the testes retain the primitive ab-
dominal position. The fact that the urinary and genital
products escape by means of their ducts into a chamber which
also receives the end of the alimentary tract is not a distinctive
feature of this group, inasmuch as it is seen in the Marsupials,
and also in certain low Eutheria, such as the Beaver and other
Rodents, and a few Insectivores. As to external features, the
Monotremata show certain archaic characters. The unspecialised
arrangement of the mammary glands has already been described.
These animals are plantigrade, if the term may be used also to
describe the aquatic Ornitliorhynchus. The ears are absolutely
destitute of a conch. The remarkable spur upon the hind-legs
furnished with a gland, which is more marked in the male, and
indeed disappears in the female of Ornithorhynchus, is a structure
which argues the specialised condition of these two modern
representatives of what must have been a large order in the past.
The skeleton shows numerous ancient characteristics. In
the skull there is no demarcation of the orbit from the temporal
fossa, a feature widely found in archaic mammals. The tympanic
remains as a slender ring, there being no auditory bulla formed
either from this or from any other bone. The malleus and
incus are large, and thus reminiscent of the quadrate and
articular bone of reptiles. In the lower jaw the absence of
a marked coronoid process, and the absence of a firm ossification
at the meeting of the two rami, may be a primitive state of

1 Quart. Journ. Micr. Sci. xxiv. 1884, p. 124.

2 Beddard, Proc. Roy. Phys. Soc. Edinb. viii. 1885, p. 354.

3 See Phil. Trans, clxxviii. 1887, where the literature of the subject is fully
cited.

io8

SKULL OF ECHIDNA

CHAP.

affairs. It must be remembered, however, that the Cetacea
show the same characters, though it is possible that they too
are developed from a low mammalian stock. In the vertebral
column we find the typical mammalian seven cervicals ; but
those characteristically mammalian structures the epiphyses are

oc.cond

Fio. 51. — Ventral view of skull of Echidna aculeata, and right half of mandible, any,
Angle of mandible ; aud.oss, auditory ossicles ; cond, condyle of mandible ; cor,
coronoid process ; max, maxilla ; oc.cond, occipital condyle ; pal, palatine ; p.max,
premaxilla ; pt, pterygoid ; sq, squamosal ; ty, tympanic ring. (After Parker and
Has well.)

totally absent in Echidna, and only to be seen in the tail-region
in Ornitliorliynchus. In having only the capitular head to the
ribs, these mammals are evidently far removed from all other
mammals, and are even more reptilian than the Theromorphous
reptiles. The large clavicles and the interclavicle (Fig. 52,

INTESTINE AND BRAIN

1O9

p. 109) are characteristic of the group, and the latter bone is
peculiar to the Monotremata among mammals. So, too, is the
large coracoid. In the scapula there is a spine which coincides
with the anterior border of that bone. The arrangement of the
muscles in this region proves conclusively that this projection is
the homologue of the spine and the acromion of other mammals.
Here, again, we have a point of likeness to the Cetacea.1 In
the pelvis the acetabulum is perforate (in Echidna), as in
Sauropsida.

Considering the numerous very
archaic features which the general
structure of this group displays,
it is surprising to find how typi-
cally mammalian they are in
certain other peculiarities. The
mammalian diaphragm, one of
the distinguishing features of the
class, is perfectly normal in the
Monotremata, The alimentary
canal shows no great divergences
from the normal structure. The

Stomach is almost globular, with FIG- 52.— Side view of right half of the

shoulder girdle of a young Echidna
(Echidna aculeata). x 1. «, Acro-
mion ; c, coracoid ; cb, coracoid border ;
cl, clavicle ; ess, coraco- scapular
suture ; ec, epicoracoid ; gb, glenoid
border ; gc, glenoid cavity ; ic, inter-
clavicle ; pf, postscapular fossa ; ps,
presternum ; s, spine ; ss, suprascapu-
lar epiphysis ; ssf, subscapular fossa.
(From Flower's Osteology.)

a projecting pyloric region in
Ornithorliynchus ; the intestine
is divided into a " small " and
" large " intestine by a slender
caecum. The liver has the sub-
divisions that this organ usually
shows in the Mammalia. How-
ever, the presence of the ventral mesentery and of the
abdominal vein in Echidna and Ornithorhynchus has already
been mentioned as a distinctive character. The peculiar and
apparently partly primitive valve of the right ventricle has been
described above (see p. 66). The brain is in most respects
mammalian in its characters, but naturally shows some important
differences. Dr. Elliot Smith, who has most recently studied
this question,2 is of opinion that the size of the cerebral hemi-
spheres is not at all reptilian ; indeed, it " greatly exceeds that of

1 Muscular insertions and attachments do not, however, altogether support the
comparison. 2 Journ. Anat. Phys. 1899, p. 309.

I IO

FOSSIL ECHIDNAS

CHAP.

many other mammals." In Echidna, too, but not in Ornitho-
rhynchus, the hemispheres are well convoluted, though the arrange-
ment of these convolutions cannot be brought into line with what is
known concerning the convolutions upon the hemispheres of other
mammals. It had been stated that in these animals, at least in
Echidna, there were only two optic lobes, as in lower vertebrates,
instead of the mammalian four. The late Sir ~W. H. Flower set
this matter at rest,1 and showed that Echidna was in this respect
typically mammalian. The absence of the corpus callosum is

one of the principal features separ-
ating the Monotremes from other
mammals.

The Monotremata are repre-
sented to-day by two types, Ornitho-
rhynchus and Echidna, which are
no doubt worthy of being placed in
separate families. Fossil remains
of the group (apart from the prob-
lematical Multituberculata) are only
known from Pleistocene times in
Australia, and consist of the bones
of a large species of Echidna, and
some fragments of Ornithorhynchus,
FIG. 53.— Braiu of Echidna aculeata, indicating a smaller animal than

dorsal view. (Nat. size.) (From .-, livintr Platvmm
Parker and Haswell's Zoology.)

Fam. 1. Echidnidae. — This

family contains two genera, of which Echidna is the older and
much the better known. The skin is abundantly covered with
spines, with which are mingled hairs. The snout is tapering,
the tail rudimentary, and the fingers and toes five in number.
The spur and gland upon the calcaneum are smaller than in
Ornithorhynchus. The claws are very strong, serving to tear
open the ants' nests, upon the inhabitants of which the Echidna
feeds, licking them up with a long extensile tongue like that of
Myrmecophaga. In relation to this habit the salivary glands are
enormously developed, and indeed the animal has been con-
founded with Myrmecophaga,2 as the vernacular name " Australian
Anteater " exemplifies.

1 Proc. ZooL Soc. 1864, p. 18.
2 Myrmecophaga aculeata was the name given by Shaw.

v SPECIES OF ECHIDNA III

In the skull the Echidna differs from Ornithorhynchus in
the greater extension backwards of the palatines, and the larger
size of the pterygoids. The extent and relations of these bones
to each other is not at all unlike that which obtains in many
Whales. The premaxillae show traces of the same divergence
followed by convergence of their ends that is seen in the
Platypus. There are only sixteen pairs of ribs, and either
three or four lumbar vertebrae. Echidna has no trace of teeth,
and there are no horny pads which take their place ; the mouth
is as edentulous as in the true American Anteaters. The brain
(Fig. 53) is marked by sulci, contrary to what we find in
Ornithorhynchus. The genus has been divided into three species,

FIG. 54. — Australian Anteater. Echidna aculeata.

but it is doubtful whether more than one can be allowed, which
ranges from Australia through the Papuan region. While there
is but one species of true Echidna, a New Guinea species must
clearly be referred to a distinct genus Proechidna} This animal
is to be distinguished by the fact that there are usually but
three toes on each foot. But there are copious rudiments of
the other phalanges, upon which claws are sometimes developed.
The beak is curved downwards, and the back is rather arched ;
the whole animal has the most singular likeness to an Elephant !
The ribs are increased by one pair, and there are four lumbar
vertebrae. The one species is named P. Iruijnii. The Hon. W.
Pvothschild 2 distinguishes a form P. nigroaculeata, which is allowed
by Mr. Lydekker.

1 Zaglosms has apparently priority as a name ; but Proechidna is better known.
2 Proc. Zool. Soc. 1892, p. 545.

I I 2 FOOD AND HABITS

The Echidna feeds like anteaters, by thrusting its tongue
into an ant-hill-, and waiting until it is covered with indignant
and marauding ants, which are then swallowed. But this
animal also devours worms and insects, which are extracted from
their hiding-places by the tongue. It is mainly nocturnal, and
prefers the seclusion of the densest scrubs of the bush, or rocky
spots where it is free from intrusion. Dr. Semon did not find
that the spur of this animal was used at all in self-defence ;
but he thinks that possibly the weapon may be used, in the
breeding season only, in the combats of the males for the females,
when perhaps, as has been shown to be the case in Omithorkyn-
chus, the gland attached to it produces a poisonous secretion.

The egg, as it appears, is transferred to the pouch by the
mouth of the mother ; the shell is broken by the emerging-
young one, which has an egg-breaking tubercle on its snout
for this purpose ; the mother removes the shell. When the
young has attained a certain size, the mother removes it from
the pouch, but takes it in from time to time to suckle it. When
on her nightly rambles the young one is left in a burrow dug
for the purpose. Dr. Semon was able, from his own observations,
to substantiate this act of intelligence on the part of the
Echidna. It is well known that the temperature of the Mono-
tremes is less than that of higher mammals ; in addition to this
fact Dr. Semon found that the range of variation of tempera-
ture in the Echidna was as much as 13 degrees or more. It is
thus intermediate between the " poikilothermal " reptiles and the
" homoeothermal " mammals.

Fam. 2. Ornithorhynchidae. — There is no need to attempt to
define this family, since it contains but one genus Ornithorhynchus,
with but one species, 0. anatinus. The general aspect of the
animal is well known. It is covered with dense fur of a blackish
brown colour ; the limbs are short and five-toed, the toes being
webbed. The tail is longish and broad, being flattened from
above downwards. The webbing on the anterior toes consider-
ably outdistances the tips of the claws, as in the Seals. But this
is not the case with the hind-feet. The "beak," which is broad
and flat, and does actually suggest that of a duck, is not covered
with horn, as is often stated, but with a fine, soft, sensitive, naked
skin, which abounds in sense-organs of a tactile nature. As to
characters derived from the skeleton, Ornitkorhynchus has seven-

TEETH OF PLATYPUS I I 3

teen pairs of ribs and only two lumbar vertebrae. The skull is
expanded in front, and the bill is supported by two, at first
diverging, and then converging, premaxillae. Between them is
the famous " dumb-bell shaped bone," which is believed to be the
representative of the reptilian prevomer. The pterygoids are
smaller than in Echidna, and the hard palate does not extend
so far back as in that genus. The brain of this genus is
smooth.

The discovery of the real teeth of Ornithorhynchus only dates
from the year 1888, when they were found by Professor Poulton l
in an embryo. Later Mr. Thomas found 2 that the teeth persist

FIG. 55. — Duck-billed Platypus. Ornithorhynchus anatinus. x £.

for a considerable portion of the animal's life, and are only shed,
like milk teeth, " after being worn down by friction with food
and sand." We have already (p. 98) called attention to the
general similarity of these teeth to those of certain of the
earliest Mammalia and of mammal-like reptiles. The teeth are
all molars, and they are either eight or ten in number. They
are replaced by the horny plates of the adult animal ; but the
mode of replacement is curious. The plates are developed from
the epithelium of the mouth, but round and under the true
teeth ; the epithelium of the mouth grows gradually under the
calcified teeth, a method of growth which has possibly some-
thing to do with the shedding of the latter. The hollows and

1 Quart. J. Micr. Sci. xxix. 1888, p. 353.

2 Proc. Roy. Soc. xlvi. 1889, p. 127. See also Stewart, Quart. J. Micr. Sci. xxxiii.
1892, p. 229.

VOL. X J

nas.

FIG. 56. —Skeleton of
male Ornithorhyn-
chus. Ventral view.
The right fore - limb
has been separated
and turned round so
as to bring into view
the dorsal surface of
the manus.- The
lower jaw is removed.
ace. tars. Accessory
tarsalbone supporting
the spur ; ant.pal.for,
anterior palatine fora-
men ; asi, astragalus ;
ati, atlas ; ax, axis ;
bs.oc, basi - occipital ;
bs.sph, basi-sphenoid;
calc, calcaueum ; cbd,
cuboid ; cerv. rb, cer-
vical rib ; doc, clavi-
cle ; cond.for, fora-
men above inner cou-
dyle of humerus ; cm;
coracoid ; nm, cunei-
form of carpus ; dent,
horny dental plate ;
ect.cun, ecto-
cuneiform ;
ent.cvn, en-
to - cunei-
form ; ep.co,
epicoracoid ;
epist, epi-
sternum ;
ep.pb, epipubis ; fb,
fibula ; fern, femur ;
for. mag, foram en mag-
num ; glen, glenoid
cavity of shoulder-
joint ; glen, glenoid
cavity for mandible ;
hum, hxi m erus ;
in.cond, inner con-
dyle of humerus ;
inf.orb.for, points to
position of infra-orbi-
tal foramen ; infr.proc,
inferior processes of
caudal vertebrae ;
int.rbs, intermediate
ribs ; isch, ischium ;
mag, magnum of car-
pus ; max, maxilla ;
max/or, maxillary
foramen ; metat.I,
first metatarsal ;
metat. V, fifth meta-
tarsal; nas.cart, nasal
cartilage ; obt, obtu-
rator foramen ; ol,
olecranon ; out.cond, outer condyle of humerus ; pal, palatine ; pat, patella ;
post.pal.for, posterior palatine foramen ; pr.max, premaxilla ; pr.st, presternum ;
pter, pterygoid ; pub, pubis ; rod, radius ; scap, scapula ; scaph, scaphoid of tarsus ;
scaph.lun, scapho-lunar ; ses, sesamoid bones of wrist and ankle ; sp, tarsal horny spur ;
sq, squamosal ; tib, tibia ; trd, trapezoid ; trm, trapezium ; tym.c, tympanic cavity ;
uln, ulna ; unc, unciform ; vom, vomer ; x, dumb-bell shaped bone : zyg, zygomatic
arch ; /- V, digits 01 mauus ; V, foramen for fifth nerve. (From Parker's Zoology.)

CHAP, v DR. SEMON'S OBSERVATIONS 1 1 5

grooves in the plates are the remains of the original alveoli of
the teeth.

The Duck-billed Platypus is, as every one knows, an aquatic
animal. It is not found all over Australia, but is limited to the
southern and eastern parts of that continent, and to Tasmania.
The animal excavates a burrow for itself in the bank of the slow
streams which it frequents. The burrow has one opening below
the water and one above ; and it is of some length, twenty to
iifty feet. The Platypus feeds upon animal food, chiefly " grubs,
worms, snails, and, most of all, mussels." These it stows away
when captured into its capacious cheek-pouches. The food is
then chewed and swallowed above the surface as the animal
drifts slowly along. Dr. Semon, from whose work, In the
Australian Bush, this account of the animal's habits is quoted,
thinks, that in the nature of the food of the creature the ex-
planation of the loss of the teeth is to be found. He is of
opinion that for cracking the hard shells of the mollusc Corbicula
nepeanensis, upon which Ornithorhynchus mainly feeds, the horny
plates are preferable to brittle teeth. Ornithorhynchus is appar-
ently not eaten by the natives by reason of its ancient and fish-
like smell. Besides, it is hard to catch on account of its diving
capacities, which are aided by an acute sense of sight and of
hearing. When the Duck-bill was first brought to this
country it was believed to be a deliberate fraud, analogous to
the mermaids produced by neatly stitching together the fore-
part of a monkey and the tail of a salmon.

CHAPTER VI

INTRODUCTION TO THE SUB-CLASS EUTHERIA

SUB-CLASS IL— EUTHERIA

Definition. — Mammalia with teats. Mammary glands of seba-
ceous type. Heart with entirely membranous and complete right
auriculo- ventricular valve. Brain generally with a corpus cal-
losum. Coracoid much reduced and not reaching sternum. No
interclavicle. Vertebrae with epiphyses. Ribs double-headed.
Viviparous, with a small ovum.

In this group are included not only the Eutheria in the sense
of Huxley, but also his Metatheria. Though the Metatheria, or
Marsupials as we shall term them, undoubtedly form a most
distinct order of mammals, perhaps even a trifle more distinct
than most others, their differences from the remaining tribes are
not by any means so great as those which separate Ornitho-
rliynchus and Echidna from all other mammals. In his well-
known memoir upon the arrangement of the Mammalia,1 Pro-
fessor Huxley enumerated eleven characters as distinguishing the
Metatheria either from the Prototheria or from the Eutheria.
Of these only three were characters in which they approach the
lower mammals. According to his showing, therefore, the
preponderance of marsupial features are Eutherian. The three
characters of Protothenan type are (1) the presence of epipubes ;
(2) the small corpus callosuin ; (3) the absence of an allantoic
placenta.

The last of these can be dismissed, in consequence of the
recent discovery of an allantoic placenta in Perameles. The first
character is apparently a valid distinction between the Marsupials
1 Proc. Zool. Soc. 1880, p. 649.

EUTHERIA CORPUS CALLOSUM

117

and their mammalian relatives higher in the series ; but it is
not a character that should have been made use of by Huxley,
since he believed in the existence of a corresponding element in
the Dog. As to the corpus callosum (Fig. 50, p. 7V) being small,
that seems to be not more than a slight difference of degree.1
A number of other characters of secondary importance were added
by Huxley to the weight of evidence which led him to form a
group Metatheria for the Marsupials. Some of these, however,
are now known to be not evidence in that direction. For in-
stance he observed that no
Marsupial had more than a
single successional tooth. It
seems at the present moment
to be fairly clear that Marsu-
pials have a milk dentition
like other Eutherians, but

hip.com.

anl.com

jnecl

-ve.nl.3

section. ant.com, Anterior commissure ;
cbl, cerebellum ; c.mam, corpus mammil-
lare ; col.forn, column of the fornix ; c.qu,
corpora quadrigemina ; gang.hab, ganglion
habenulare ; hip.com, hippocampal com-
missure ; med, medulla oblongata ; mid.com,
middle commissure ; olf, olfactory lobe ;
opt, optic chiasma ; tub.olf, tuberculum
olfactorium ; vent. 3, third ventricle. (From
Parker and Haswell's Zoology.)

that only one of these teeth,
the fourth premolar, comes
to functional maturity. That
it is really one of a complete

milk Series is evidenced by FIG. 57._Brain of Echidna aculeata ; sagittal

the fact that this tooth is
differentiated contemporane-
ously with another series
formerly held to belong to
the so-called prelacteal denti-
tion.2 There still remains, of
course, the actual fact that
the milk dentition is not for the most part functional, but its
significance breaks down with these fresh discoveries. Of this
Professor Osborn has remarked : " The discovery of the complete
double series seems to have removed the last straw from the
theory of the marsupial ancestry of the Placentals." But Huxley
did not lay much stress upon this matter of the teeth, since he
observed that similar suppressions of the milk dentition were to
be found in many other mammals admittedly Eutherian.

Huxley regarded the peculiarities in the reproductive organs

1 Moreover, the " corpus callosum and the anterior commissure ... in ... Erin-
uceus and Dasypus are almost Monotreme-like."

2 See Wilson and Hill, Quart. J. Micr. 3d. xxxix. 1899, p. 427.

I I 8 MOLARS OF EUTHERIA CHAP.

of the Marsupials as " singularly specialised characters," in no
way intermediate in character. This view applies also to the
pouch, which, as already stated, distinguishes the adults of that
group. But the impossibility of using this last character as
one of any importance has been shown by the discovery of
rudiments of it in embryos of undoubtedly Eutherian mammals
(see p. 18).

Less stress is laid now upon the existence of four molars in

the Marsupials as

hiPf°m dividing them from

. ,
_ the higher mammals

cbl stsfSS^X than was formerly the

case. The total denti-

.aril.com .• <? .••

tion ol the group is

on the whole com-
posed of more numer-
ous individual teeth

mecl hyao C7£/* than in the typical

c.mam venl.9 Eutheria; but we have

FIQ. 58.— Sagittal section of brain of Rock Wallaby exceptions like the

(Petrogale penwittata). ant.com, Anterior commis- -«7-u0ipa <-!,„ ArTri,
sure ; cbl, cerebellum ; c.mam, corpus manimillare ;

c.qu, corpora quadrigemina ; cno; crura cerebri ; dillo PriodoiltcS, and

epi, epiphysis, with the posterior commissiire. ini- , , -..- ,

mediately behind it ; /.mow, position of foramen of til 6 Manatee ; Or

Monro ; hig.com, hippocampal commissure, consist- better because free

ing here of two layers continuous behind at the „ , „

spleueium, somewhat divergent in front where the irom tne Suspicion OI

septum lucidum extends between them ; ffypo, hypo- secondary multiplica-

physis ; med, medulla oblongata ; mid.com, middle .

commissure ; olf, olfactory lobe ; opt, optic chiasma ; tion, Otocyon and OCCa-

vent. 3, third ventricle. (From Parker and Haswell's 8ionaUy (according to

Zoology.) J ^ °

Mr. Thomas) Centetes.
In the last two there are at least sometimes four molars.

On the other hand, a few archaic characters of some import-
ance crop up here and there among the Marsupials, which are
sometimes held to point to a primitive ancestry. It has been
remarked that in Marsupials it is the fourth toe which is dominant
in size, whereas in Ungulates it is the third. An attempt has
been made to explain this on the view (reasonable enough in
itself) of a tree-living ancestry for the group. A greater develop-
ment of the fourth toe is, however, by no means a necessary
character of arboreal creatures ; the Primates themselves are an
exception. Nor is this prevalence universal among the Marsupials ;

vi POSITION OF MARSUPIALS 119

in Jlyrmecobius (alone) is the third toe the longest ; and no great
difference can be detected between the third and fourth toes in
the case of the genera Phascologcde, Didelphys, and some others.
Professor Leche compares the predominance of the fourth toe
with the hyperphalangeal condition in the fourth toe of the embryo
Crocodile, and considers it an archaic feature, not surpassed by
the ancient characteristics of the Monotremata. Again it has been
pointed out that in Phascologale and Perameles, the epistropheus
(axis vertebra) has a separate rib as in Ornithorhynclius. In the
third place, the likeness of the teeth of Myrmecobius to those of
Ornithorliynchus is an argument in the same direction, which is
furthermore supported by the great age (Mesozoic) of the Meta-
therian group, if we are right in regarding those extinct creatures
as Marsupials.

AVe may now mention certain facts which are not so generally
used. The partly primitive structure of the right auriculo-
ventricular valve in the Monotremata has no counterpart in
any Marsupial which has been dissected ; but there are traces
in the latter of the characteristic " ventral mesentery " of Orni-
thorhynchus and Echidna,} Mr. Caldwell's interesting observation
upon the segmenting egg of the Marsupial, the incompleteness
of the first segmentation furrow (reminding us of the meroblastic
ovum of the Mouotreme), may possibly not turn out to be so
exclusively Marsupial a feature as has been thought.

The balance of evidence thus points to the nearer relationship
of the Marsupials to the Eutherian mammals ; and their great
specialisation combined with certain evidences of degeneration
(disappearance in part of the milk dentition), and their age, point
to the fact that they are, at any rate, the descendants of an early
form of Eutherian. But they must have separated from the
Eutherian stock after it had acquired a definite diphyodonty and
the allantoic placenta, the two principal features of the Eutherian
as opposed to the Prototherian mammals.

Nevertheless it seems probable that the Marsupial tribe is
derived from some of the earliest Eutherians. And on this view
may be explained the retention of Prototherian characters.

The remaining Eutheria are obviously all to be referred to
one great division with the possible exception of the Whales,
whose affinities form one of the principal difficulties to the student
1 In Dendrolagus at any rate. See Proc. Zool. Soc. 1895, p. 132.

120 RELATIONSHIP OF CETACEA PHAP.

of this group. A short resume of what is at present thought
of the systematic position of this anomalous order is appro-
priate here. Albrecht went so far as to regard the Cetacea as
the nearest group of animals to the hypothetical Promammalia.1
But discounting his arguments by the removal of such of them
as relate to structure plainly altered by the singular mode of
life of these creatures, there is really a great deal to be said in
favour of his view.

The chief facts which argue a primitive position among
mammals for the Cetacea are perhaps : (1) the slight union of
the rami of the lower jaw ; (2) the occasionally rather marked
traces of the double constitution of the sternum ; (3) the long
and simple lungs ; (4) the retention of the testes within the
body-cavity ; (5) the occasional presence (in Bcdaenoptera) of a
separate supra-angular bone. These points, however, are but few,
and are not of such great weight as those which ought to be pre-
sent to establish a claim to separate treatment for the Cetacea
as opposed to the Eutheria. If this group of mammals can be
tacked on anywhere, it appears to us that the nearest relatives
are not, as is sometimes put forward, the. Ungulata or the
Carnivora, but the Edentata. There are quite a number of
rather striking features in which a likeness is shown between
these apparently diverse orders of mammals. The chief ones
are these: (1) the existence of traces of a hard exoskeleton, of
which vestiges remain in the Porpoise ; (2) the double articula-
tion of the rib of the Balaenopterids to the sternum, with which
compare the conditions obtaining in the Great Anteater ; (3) the
concrescence of some of the cervical vertebrae ; (4) the share
which the pterygoids may take in the formation of the hard
palate ; (5) the fact that in the Porpoise, at any rate, as in many
Edentates, the vena cava, instead of increasing in size as it
approaches the liver, diminishes.

Another group which is perfectly isolated is that of the
Sirenia. The alliance advocated by some with the Cetacea, and
quite recently renewed by Professor Haeckel, is contradicted by so
many important features that it seems necessary to abandon it.
The recent discovery of a fossil Sirenian jaw by Dr. Lydekker witli
teeth highly suggestive of those of Artiodactyla, may prove a clue.
A third group which is so isolated as to have been placed in a
1 Anat. Anz. i. 1886, p. 338 ; and see Weber, ibid, ii. 1887, p. 42.

vi CREODONTA AND MODERN MAMMALS 121

primary division, proposed to be called Paratheria, is that of the
Edentates. Probably th« group so called should really be
divided into the Edentata and the Effodientia, the latter con-
taining the Old World forms. Whether or not it be ultimately
shown that the Ganodonta are ancestral Edentates (sensu strictiori},
the connexion of the group with others is not at present plain.
The same is the case with the extensive order of Eodents. It
is true that the extinct order of the Tillodontia shows certain
Rodent-like characters on the one hand, and likenesses to Ungulates
on the other. Certain likenesses shown by such apparently
diverse animals as the Rabbit and the Elephant used to be
insisted upon by Professor Huxley. For the present, however, the
Rodents must remain as an isolated group with only very
dubious affinities to others. The remaining groups of existing
mammals are easier to connect. At first the differences between
a Cat and a Horse seem to be quite as wide as those which
separate any two of the higher Eutherian orders. But it seems
to become clearer and clearer, as palaeontological investigation
proceeds, that the bulk of the Ungulate and the Carnivorous,
Insectivorous, and perhaps Lemuroid stocks converge into the
early Eocene Creodonta. From the Lemuroid branch the higher
Primates can be derived. The only" Ungulates " which cannot
be fitted in with some reasonable probability, is the group of the
Proboscidea. But of the early forms of this division we have at
present no knowledge.

CHAPTER VII

EUTHEKIA MARSUPIALIA

Order I. MARSUPIALIA1

THE Marsupials may be thus defined : — Terrestrial, arboreal, or
burrowing (rarely aquatic) mammals, with furry integuments ;
palate generally somewhat imperfectly ossified ; jugal bone reach-
ing as far as the glenoid cavity ; angle of lower jaw nearly always
inflected. The clavicle is developed. Arising from the pubes are
well developed and ossified epipubic bones. Fourth toe usually
the most pronounced. Teeth often exceed the typical Eutherian
number of forty-four ; molars generally four on each side of each
jaw. As a rule but one tooth of the milk set is functional, which
is (according to many) the fourth premolar. Teats lying within
a pouch, in which the young are placed. Young born in an im-
perfect condition, and showing certain larval characters. There
is a shallow cloaca. The testes are extra-abdominal, but hang in
front of the penis. In the brain the cerebellum is completely
exposed ; the hemispheres are furrowed, "but the corpus callosum
is rudimentary. An allantoic placenta is rarely present.

Structurally the Marsupials are somewhat intermediate be-
tween the Prototheria and the more typical Eutheria, with a
greater resemblance to the latter.

The name Marsupial indicates what is perhaps the most
salient character of this order. The pouch in which the young
are carried is almost universally present. It is less developed

1 Works dealing exclusively with the Marsupials are : Lydekker, in Allen's
Naturalists' Library, 1894 ; Aflalo, Natural History of Australia, Macmillan and
Co. 1896 ; Waterhouse, Natural History of Mammalia, i. London, 1848 ; Old-
field Thomas, British Museum Catalogue of Marsupialia arnd Monotremata, 1888.

MARSUPIALIA POUCH

123

on the whole in the Polyprotodont forms, such as the Thylacine,
Dasyures, etc., but is found in so many of them that the
two divisions of the Marsupials, the Diprotodonts and the Polv-
protodonts, cannot be raised to distinct orders on this and other
grounds. The marsupial pouch of the Marsupials must not, as
has been already pointed out, be confounded with the pouch of the

Fro. 59. — Rock Wallaby (Petrogale xaiithopus), with young in pouch.
(After Vogt and Specht.)

xf.

Monotreme mammals. Distinct teats are found in the marsupium
of the Marsupials, while there are none in the mammary pouch of
the Monotreme, the pouch itself indeed representing an un-
differentiated teat, of which the walls have not closed up. The
pouch opens forward in the Kangaroos, and backwards in the
Phalangers and in the Polyprotodonts. Its walls are supported by
a pair of bones diverging from each other in a V-shaped manner ;
these are cartilaginous and vestigial in the Thylacine. They

124

MARSUPIAL " LARVAE "

are the precise equivalents of similar bones in the Monotremata.
It has been held, but apparently erroneously, that these bones
are mere ossifications in the tendons of the external oblique
muscle of the abdomen, or of the pyramidalis of the same region ;
and vestiges have been asserted to exist in the Dog. Such
bonelets are undoubtedly present in the
Dog ; but it seems clear from their develop-
ment in Marsupials, as structures actually
continuous with the median unossified por-
tion of the symphysis pubis, that the
" marsupial bones " belong to that part of
the skeleton, and that they correspond with
the epipubis of certain amphibians and
reptiles. The pouch, it may be remarked,
exists in a rudimentary form in the males
of many Marsupials.

The most salient feature in the life-
history of the Marsupials is the imperfect
condition in which the young are born.
The egg is no longer
laid, as in the Mono-
tremes; but curiously

FIG. 60.— Ventral surface of enough the OVU111,

innominate bone of Kan- wMch hag the gmall
garoo (Macropus major),

x £. «, Acetabulum ; ab, size of that of the

supial " bone ; pb, pubic completely at the

border ; nt, pectineal tu-r- i* • • / ~\ir

bercle; *s symphysis; si, nrst division (as Mr.

supra -iliac border ; ss, Caldwell has shown),
sacral surface ; thf. thy- j , , •

and this

(From Parker and Has-
well's Zoology.)

roid foramen : ti, tuber-

osity of ischium. (From mental feature may FIG. 61. — Mammary foetus

Flower's Osteology.) , , , , of Kangaroo attached to

perhaps be looked the te|L (Nat size>)
upon as a reminiscence of a former large-
yolked condition. The young when born
are small and nude : the newly born young of a large Kangaroo
is perhaps as large as the little finger. The young are trans-
ferred by the lips of the mother to the pouch, where they are
placed upon a teat. It is an interesting fact that they are
not merely imperfect foetuses, but that they are actual larvae.
They possess in fact at any rate one larval organ in the shape of

THE PLACENTA 125

a special sucking mouth. This sucking mouth is an extra-uterine
production, and is of course an adaptation to the particular needs
of the young, just as are other larval organs, such as the chin-
suckers of the tadpole, or the regular ciliated bands of the
larvae of various marine invertebrate organisms.

There are a number of other features which distinguish the
Marsupials from other mammals.

The cloaca of the Marsupials is somewhat reduced, but is still
recognisable. Its margins in Tarsipes are even raised into a
wall, which projects from the body.

The tooth series of the Marsupials was once held to consist of
one dentition only, with the exception of the last premolar, which
has a forerunner. The interpretation of the teeth of Marsupials
are various. Perhaps most authorities regard the teeth as being
of the milk dentition, with the exception of course of the single
tooth that has an obvious forerunner. But there are some who
hold that the teeth are of the permanent dentition. In any case
it is proved that a set of rudimentary teeth are developed before
those which persist. Those who believe in the persisting milk
dentition describe these as prelacteal. Another matter of im-
portance about the teeth of this order of mammals is that their
numbers are sometimes in excess of the typical Eutherian 44.
This, however, holds good of the Polyprotodonts only.

It was for a long time held that the Marsupials differed
from all other mammals in having no allantoic placenta. But
quite recently this supposed difference has been proved to be
not universal by the discovery in Perameles of a true allantoic
placenta. The Marsupials have been sometimes called the Di-
delphia. This is on account of the fact that the uterus and the
vagina are double. Very frequently the two uteri fuse above,
and from the point of junction an unpaired descending passage
is formed (see Fig. 48 on p. 74).

A character of the brain of Marsupials has been the subject
of some controversy. Sir Kichard Owen stated many years ago
that they were to be distinguished from the higher mammals by
the absence of the corpus callosum. Later still it was urged
that a true corpus callosum, though a small one, was present ;
while, finally, Professor Symington l seems to have shown that

1 "The Cerebral Commissures in the Marsupialia aiid Monotremata," Journ.
Anat. Phys. xxvii. 1893, p. 69.

126

THE SKULL OF MARSUPIALS

.p.maac

the original statement of Owen was correct, at least in part. It

is at most feebly developed (see Fig. 58, p. 118).

As to skeletal characters, the Marsupial skull has on the

whole a tendency towards a permanent separation of bones

usually firmly ankylosed. Thus the orbitosphenoids remain

distinct from the pre-
sphenoid. The palate is
largely fenestrated, a return
as it were — says Professor
Parker — to the Schizogna-
thous palate of the bird.
The mandible is inflected ;
this familiar character of
the Marsupials goes back
to the earliest representa-
tives of the order in Meso-
zoic times (see p. 96); but
it is not absolutely uni-
versal, being absent from
the much weakened skull
of Tarsipes. On the other
hand, the inflection is nearly
as great in certain Insecti-
vores, in Otocyon, etc. The
malar always extends back
to form part of the glenoid

FIG. 62. -Skull _of Rock Wallaby ' (P^ogale cayit The shoulder girdle
penicillata). (Ventral view.) all, Alisphenoid; *

bas.oc, basi-occipital ; bas.sph, basi- sphenoid ; has lost the large COraCoid

2Z-£2&?%,%SZ£££?^ of Monotremes; this boue

premaxilla ; pr.sph, presphenoid ; pt, ptery- has the vestigial character
eoid ; sq. squamosal ; ty. tympanic. (From . i •. ,\

Parlor and Haswell's /oology.) that lt; POSSCSSeS 111 other

Eutheria. The clavicle is

present except in the Peranielidae. A third trochanter upon
the femur seems to be never present.

The Marsupials cannot be regarded as an intermediate stage
in the origin of the Eutheria for a number of reasons. In the
first place, the nature of their teeth shows them to be degenerate
animals ; one set, whether we regard it as the milk or permanent
dentition, has become vestigial. The recent discovery of a true
allantoic placenta in Perameles removes one reason for regarding

bas.oc

vii GEOGRAPHICAL RANGE 1 27

the Marsupials as primitive creatures. It implies on the whole
that the Marsupials have sprung from a stock with an allantoic
placenta. The alternative is to assume the independent develop-
ment of an allantoic placenta in both groups of the Mammalia ;
unless indeed the genus Perameles is to be held to be the most
primitive race of Marsupials living, a hypothesis which does not
appear on the face of it likely. So long as it was believed that
the mammary pouch of the Monotremes was the equivalent of
the marsupium of the Marsupials, the persistence of this structure
seemed to be a bond of union between the groups. But it is now
known that the marsupium is a special organ confined to the
Marsupials, an argument which is rather in favour of their being
a lateral development of the mammalian stem. It is to be re-
marked also that the marsupium is feeblest in the Polyproto-
donts, which may perhaps be looked upon as the most primitive
of the Marsupials, owing to their more numerous teeth and other
points to be referred to immediately.

Not only are the Marsupials interesting from the point of
view of their structure ; their present and past distribution is of
equal interest. During the Mesozoic epoch they occurred in
Europe and North America ; but not, so far as negative evidence
means anything, in Australia, which is now their headquarters.
In Europe Marsupials lingered on into the Tertiary period, when
they finally became extinct. In America, of course, the group
has persisted to the present day. Now it is important to notice
that the two main subdivisions of the Marsupials, the Poly-
protodontia and the Diprotodontia, exist to-day in both Australia
and South America. These two divisions, it should be explained,
differ principally in that one has numerous, the other rarely
more than two,1 incisors in the lower jaw. It is perhaps the
more widely distributed opinion that the Polyprotodontia are the
more archaic group ; this opinion rests upon one or two facts in
addition to the absence of specialisation in the incisor teeth.
Among the Polyprotodontia the total number of teeth is greater —
a clearly primitive character; secondly, the general form of the
body of these animals, with four subequal limbs and carnivorous
or omnivorous diet, contrasts with the purely vegetarian and much
specialised Kangaroos at any rate. Finally — and sufficient stress

1 When there are more than two, two are especially developed. See Figs. 76, 77
(pp. 149, 150).

128 THE ANTARCTIC CONTINENT CHAP.

has perhaps not been laid upon this matter — the brain among the
Polyprotodonts is less convoluted than among the genera of the
other division. This statement is of course made with due regard
to parallelism in size (see p. 77). It is well known that the
complexity of a brain bears a distinct relation to the size of its
possessor within the group. Now the most ancient Marsupials
are decidedly more Polyprotodont-like. No European form from
the earlier periods is distinctly to be referred to the Diprotodonts.
But both divisions now exist in America and Australia.

We must assume, therefore, one of three hypotheses. Either
the differentiation into the two great divisions occurred in Jurassic
or Cretaceous times before the migration of the order southwards ; or
the Diprotodont type is only a type, and not a natural group, i.e. it
has been separately evolved in America and Australia ; or, finally,
there was formerly a land-connexion in the Antarctic hemisphere,
along which 'the Diprotodonts of Australia wandered into South
America. The middle hypothesis has this to commend it, that
syndactylism occurs in both divisions, and that in some Dipro-
todonts the pouch opens backwards as it does in the Polyprotodonts.
So great are the resemblances that but little difference is really left
— of great importance that is to say. Hence it is not difficult to
imagine the reduction of the incisors having taken place twice.
In favour of the first hypothesis there are no positive facts.
Finally, in favour of the last, which is so strongly supported by
the facts of distribution derived from the study of other groups
of animals,1 there is at least this striking fact or rather series of
facts : that some of the South American fossil Polyprotodonts
have a " strictly Dasyurine relationship." - If there has not been
a direct migration, then the Dasyurine type has been twice evolved,
an improbability that few will attempt to explain away. In any
case we shall adopt here the usual division of the Marsupials into
Diprotodontia and Polyprotodontia.

SUB-ORDER 1. DIPROTODONTIA.

This group includes the herbivorous Marsupials. The incisors
are as a rule three above, but one only in the Wombats. Below

1 See for a further discussion of this subject the zoogeographical handbooks of
Mr. Lydekker and myself, quoted on p. 78 (footnote).

2 To this may be added Mr. Thomas' observation that the family of American
Opossums is " very closely allied to the Dasyuridae, from which, were it not for its
isolated geographical position, it would be very doubtfully separable."

4.

AMERICAN DIPROTODONTS

129

is one strong pair, with occasionally one or two rudimentary incisors.

tor ,/ti.

FIG. 63. — Skull of Wombat (Phascolomys wombat}. (Lateral view.) ang, Angular pro-
cess ; cond, condyle of mandible ; ext.atid, opening of bony auditory meatus ;
ex.oc, exoccipital ; ju, jugal ; Icr, lachrymal ; max, maxilla ; nas, nasal ; p.max,
premaxilla ; sq, squamosal ; ty, tympanic. (From Parker and Harwell's Zoology.)

The upper canines, if present, are not large. The molars are

tuberculate or ridged. All Marsupials (ex- ^

cept the Wombats) to some extent, and the

Macropods especially, are characterised by

the prolongation of the tubes of the dentine

into the clear enamel. The significance of

this fact is, however, lessened by the fact

that the same penetration of the enamel

by dentinal tubes occurs in the Jerboa, the

Hyrax, and some Shrews. The feet have

two syndactylous toes,1 less marked in the

Wombats than in the Kangaroos and

Phalangers.

This order is mainly Australian at the
present day, using the term of course in
the " regional " sense (see p. 84) ; the only
exception indeed to this statement is the
occurrence of the genus Caenolestes in South FlG 64._Bones of right
America. But it is now known that Dipro- foot of Kangaroo (Macro-

L-II •••*•"* i • j. J • .4.-U I™8 bennetti). a, Astra -

todont Marsupials formerly existed in the gaius . c> caicaneum ;
same part of the world. <*• cuboid : *> *nt°-cu-

. neiform ; n, navicular;

Fam. 1. Macropodidae. — This family //. v, second to fifth toes,
contains the Kangaroos, Wallabies, Eat- (rrom Flower's Osteology.)
Kangaroos, and Tree- Kangaroos. With the exception of Dendro-

1 Except in the South American Diprotodonts.
VOL. X K

I

130

KANGAROOS

lagus the family is terrestrial, and its numerous species
progress by leaps effected by the long hind -limbs, which are

orb

cbd

FIG. 65. — Skeleton of Wallaby ( Mao-opus ualabatus). The scapula is raised some-
what higher than in nature. The end of the tail is omitted. The head of
the femur has been separated from the acetabulum. acet, Acetabulum ; acr,
acromion process ; ast, astragalus ; cal, calcaneum ; cbd, cuboid ; chev, chevron-
bones ; cl, clavicle ; cun, cuneiform of carpus ; epi, epipubis ; fb, fibula ; fern,
femur ; hd, head of femur ; hu, humerus ; il, ilium ; isch, ischium ; obt, obturator-
foramen ; orb, orbit ; pis, pisiform ; pub, pubis ; rad, radius ; rbl, first rib ; r61:i,
last rib ; sc, scapula ; st, sternum ; tb, tibia ; troch, great trochanter of femur ; idn,
ulna ; unc, unciform; IV, fourth toe. (From Parker and Harwell's Zoology.)

decidedly, often greatly, longer than the fore -limbs. In the
hind -limb the fourth toe is very long and strong; the fifth
moderately so ; the second and third are slender and united by
skin. The tail is always long, but differs in its characters from

vii STOMACH OF KANGAROOS 131

genus to genus. The stomach is much sacculated. The dental
formula is I f C J^p P f M f . The atlas is often open below,
forming thus an incomplete ring.

Though the number of the incisor teeth in the adult
Diprotodonts is never more than three on each side in each jaw,
more numerous rudiments are present. Mr. M. Woodward l has
lately investigated the subject with interesting results. He
finds that many species present decided traces of two additional
incisors, raising the total to that which characterises the Poly-
protodontia ; but in two cases, viz. Macropus giganteus and Petro-
gale penicillata, a sixth is present, the total number being thus
in excess of that found in any other Marsupial. This, as the
author himself admits, proves too much. No mammal is known
which in the adult condition has so many incisors ; nor do the
fossil Mammalia help us to get over the difficulty ; even among
reptiles it is not usual for so many teeth to occur upon the
premaxillaries.

It is a curious fact that the two long lower incisors can be
used after the fashion of a pair of scissors, or rather a pair of
shears. Their inner edges are sharpened, and they are capable
of some motion towards and away from each other ; by their
means grass is cropped.

The stomach of Macropus (and of other allied genera) is
peculiar by reason of its long and sacculated character ; the
oesophagus enters it very near the cardiac end, which is bifid.
Messrs. Schafer and Williams 2 have shown that the squamous,
non- glandular epithelium of the oesophagus extends over the
greater part of the stomach, only the pyloric extremity and one
of the two cardiac caeca being lined with columnar epithelium.

The Macropodidae are clearly divisible into three sub-families,
which are distinguished by marked anatomical characters.

In the sub -family MACROPODINAE (including the genera
Macropus, Petrogale, Lagorchestes, Dorcopsis, Dendrolagvs, Onyclio-
gale, and Lagostrophus) there is no hallux, and the tail is hairy.
The oesophagus enters the stomach near the cardiac end. The
caecum when short has no longitudinal bands ; the liver has a
Spigelian lobe.

The second sub-family, POTOROINAE or HYPSIPHYMNINAE (in-
cluding the genera Potorous, Aepyprymnus, Bettongia, and Calo-
1 Proe. Zool. Soc. 1893, p. 450. 2 Ibid. 1876, p. 165.

132 KANGAROOS AND WALLABIES CHAP.

prymnus), consists of smaller animals than the Macropodinae,
which, however, resemble them in having no hallux, but a hairy
tail. The oesophagus enters the stomach near the pyloric end
of that organ. The caecum, though short, has lateral longitudinal
bands. The liver has no special Spigelian lobe. The canines
are always present, being rarely so in Macropodinae, and are
usually well developed.

The third sub-family, that of the HYPSIPRYMNODONTIDAE, is
doubtfully referable to the family ; it consists of but one genus
Hypsiprymnodon, which is in many points more like a Phalanger
than a Kangaroo. It has an opposable hallux and a non- hairy,
but scaly, tail. It has canines in the upper jaw.

Sub-Fam. 1. Macropodinae. — The genus Macropus includes
not only the Kangaroos but also the Wallabies, which are really

FIG. 66. — Red Kangaroo. Macropus rufus. x ^.

indistinguishable, though they have sometimes been placed in a
separate genus Halmaturus. The genus thus enlarged contains
twenty-three species. It may be thus characterised : the ears
are long, the rhinarium is usually naked, but in M. giganteus
and others a band of hairs descends to the upper lip ; a
naked band extends from the ankle to the pads on the digits,
which is interrupted in M. rufus by a band of hairs just in front
of the digits. The mammae are four. The tail is not bushv,

vii BRUYN AND SIR JOSEPH BANKS 133

but is crested in M. irma. They are for the most part found on
the Australian continent, but some species are found in the
islands to the north which belong to the Australian region.
Thus M. brunii, which is of interest as the first Kangaroo seen
by a European, is a native of the Aru islands. A specimen of
this animal, which was then living in the garden of the Dutch
governor of Batavia, was described by Bruyn in the year 1711.
M. rufus, the largest member of the group, is remarkable for the
red secretion which adorns the neck of the male. It is caused
by particles which have the appearance and colour of carmine.
M. giganteus is not, as its specific name might imply, the " giant "
of the race ; its dimensions are given as 5 feet, while M. rufus
is said to attain a length of 5 feet 5 inches, exclusive (in both
cases) of the tail.

The account which Sir Joseph Banks gives1 in his diary
of the Kangaroo is interesting, since he was one of the first
naturalists to see that creature. In July 1*770 it was reported
to him that an " animal as large as a greyhound, of a mouse
colour, and very swift " had been seen by his people. A little
later he was surprised to observe that the animal " went only
upon two legs, making vast bounds just as the jerboa does."
The second lieutenant killed one of these Kangaroos, of which
Sir Joseph Banks wrote that " to compare it to any European
animal would be impossible, as it has not the least resemblance
to any one I have seen. Its fore-limbs are extremely short and
of no use to it in walking ; its hind, again, as disproportionately
long ; with these it hops seven or eight feet at a time, in the
same manner as the jerboa, to which animal indeed it bears much
resemblance, except in size, this being in weight 38 Ibs., and the
jerboa no larger than a common rat." The beast was killed and
eaten, and proved excellent meat. Sir Joseph Banks' observations
upon the leaping of the Kangaroo are of interest, because it is
often asserted that the tail is largely made use of as a third foot
or as a support. Mr. Aflalo declares in the most positive way
that after repeatedly examining the tracks upon soft sand imme-
diately after the animal had passed, not the very faintest trace
of the impression of the tail could be discovered. The leaps of
a big Kangaroo seem to be somewhat greater than is recorded

1 Journal of the Rt. Hon. Sir Joseph Banks, Bart., K.B., P.R.S., edited by Sir
Joseph Hooker, London, 1896.

134 NAIL-TAILED WALLABY CHAP.

by Banks. It is said that 15 or even 20 feet are covered at
a bound, and in bound after bound. But in walking slowly it
can be readily seen from an inspection of Kangaroos at the
Zoological Society's Gardens that the animal does rest upon its
tail, which with the hind-legs forms a tripod.

Petrogcde with six species comes next to Macropus, and is
indeed only to be differentiated from it by the thickly-haired
and more slender tail, which is not used, as it is sometimes in
the Kangaroos, as an extra hind-limb. The Kock-Kangaroos live
among rocks, which they climb, and from which they leap ; and
the tail acts rather as a balancing pole. The most elaborate
account of the anatomy of Petrogale known to me is by Mr.
Parsons.1 The dentition as given by Mr. Thomas is I ^ C {}
Pm ^ M ^ — that of Macropus without the occasionally occurring
canine of the upper jaw. The osteological characters which
separate it from Macropus are quite insignificant. Mr. Parsons
mentions a wormian bone, " os epilepticum," at the junction of
the coronal and sagittal sutures. It was found to occur in two
out of five skulls examined, and appears not to occur in other
Kangaroos. The palatine foramina of Petrogale are so large that
the posterior part of the bone is only a narrow thickened ridge.
The small intestine of P. xanthopus is 102 inches long, the
large intestine 44 inches. The caecum has a length of 6 inches,
and is not sacculated, differing in this from the caecum of
Macropus major. The best known species are P. xanthopus and
P. penicillata. The genus is confined to Australia itself, and
does not enter Tasmania.

Onychogale includes the so-called "Nail-tailed Wallabies," which
have a thorn at the end of the tail, reminding one of the Lion and
the Leopard, whose tails have a similar armature. The muffle is
hairy. Three species are allowed by Mr. Thomas.

Lagorchestes has, like the last genus, the rhinarium, i.e. that
part of the nose immediately surrounding the nostrils, hairy
instead of smooth as in the Kangaroos proper. It is distinguished
from Onychogale by the absence of the terminal callosity to the
tail, which is rather short. The name Hare-Kangaroo is given
to the members of this genus (three species) on account of
their exceeding fleetness. This genus is limited to Australia
itself. L. conspicillatus is said to present " a remarkable resem-
1 Proc. Zool. Soc. 1896, p. 683.

TREE-KANGAROOS I 3 5

blance to the English hare," and L. leporoides was so called by
Gould on account of general appearance as well as face.

Dorcopsis has shorter hind-legs than Macropus, and a naked
muffle. The ears are small. The structure of D. luctuosa has
been studied by Garrod,1 who pointed out the existence of four
enlarged hair follicles on the neck near the rnandibular sym-
physis. These are, however, represented in the next genus
Dendrolagus, and occur also in Petrogale. The limbs are not so
disproportionate as in Macropus, and the tail is naked at the tip.

Dorcopsis and the next genus to be described, Dendrolagus,
differ from Macropus and its immediate allies, Petrogale and Lagor-
chestes, in a number of anatomical points. In the first place, the
premolars are twice the size of those of Macropus, and they have
a characteristic pattern not observable in the Kangaroos. This
consists of a median ridge (the whole tooth being rather prismatic
in shape), with lateral ridges at right angles to it. The upper
canines are developed, but are minute.

The stomach is not quite like that of Macropus, though built
upon a similar plan. The blind cardiac extremity is a single, not
a double cul-de-sac ; in this it is like that of Petrogale. The dis-
tribution of the squamous, white, oesophageal epithelium is very
much like that of Dendrolagus. In both genera the orifice of the
oesophagus into the stomach is guarded by two strong longi-
tudinal folds, which run for some distance towards the pylorus.
In Dendrolagus, at any rate, this tract is bordered on each side
by glandular patches. In Dendrolagus, moreover, the squamous
.epithelium does not extend into the cardiac cul-de-sac. This
latter is separated from the rest of the stomach by two slightly
diverging folds, which are faintly represented in Petrogale and in
Halmaturus. In the last two genera the folds surrounding the
oesophageal orifice are but slightly represented ; better in Halma-
turus than in Petrogale. But there are not the patches of glands
already referred to. The small intestine of Dorcopsis is 97
inches in length, the large being 32, i.e. proportionately long, as
in Marsupials generally. The small caecum (2|- inches) is not
sacculated.

The spleen is Macropodine, being T-shaped or Y-shaped. The
differences between Dorcopsis and the evidently closely allied
Dendrolagus will be further considered under the description of
1 Proc. Zool. Soc. 1875, p. 48.

136

MUFFLE OF DENDROLAGUS

CHAP.

the latter. Dorcopsis is confined to New Guinea, and contains
three species, viz. D. muelleri, D. luctuosa, and D. macleani.
D. muelleri has a striking resemblance to Macropus brunii, with
which it has been confounded. Though intermediate between
Macropus and Dendrolagus, these Kangaroos are not arboreal.

The genus Dendrolagus is remarkable for its un-kangaroo-like
habit of living in trees. In accordance with this change of
habit is a relative shortening of the hind-limbs, a feature which

FIG. 67. — Tree-Kangaroo. Dendrolagus bennetti. x ,15.

begins to be observable in Dorcopsis. " The general build," writes
Mr. Thomas, " is of the ordinary mammalian proportions, not
macropodiform at all." The muffle is not naked for the greater
part, though the shortness of the hairs gives that effect. As in
Dorcopsis, but not as in Macropus, the bulla tympani is not
swollen. There are altogether five species, the fifth, D. bennetti,
having been lately described from specimens living in the Zoo-
logical Society's Gardens.

The anatomy of this genus has been described by Owen for
D. inustus,1 and by myself for D. bennetti. The stomach, which

1 Proc. Zool. Soc. 1852, p. 103.

KANGAROO-RATS 137

has a single, not bifid, cul-de-sac, is sacculated by two principal
bands and other subsidiary ones. Its internal structure has
already been to some extent described. The spleen of D. bennetti
is remarkable for the fact that it is not T-shaped, whereas
D. inustus agrees with other Macropodines in the form of this
organ. The small intestine of D. bennetti is 95 inches long, the
large 38. The caecum appears to differ in the two species; it
is smaller in D. bennetti, where it is only 2 inches in length.
The most remarkable feature of the liver is the large size of the
left lateral lobe and the bilobed condition of the Spigelian lobe ;
this at least was the case with D. bennetti. A recently-described
species l has been attentively studied in its native haunts by Dr.
Lumholtz.2 It lives in the highest parts of the mountainous
scrubs of Queensland, where it moves quickly on the ground as
well as among the trees. It is hunted with Dingos by the
" blacks," and is eaten by them.3

Lagostroplius is a generic name that has been proposed by
Mr. Thomas for a small Wallaby 18 inches in length, which is
distinguished by the fact that the long claws of the hind-limbs
are entirely hidden by long and bristly hairs ; the muffle is
naked ; there is no canine. The bullae are swollen. There is but
one species of the genus, L. fasciatus, a native of West
Australia.

Sub-Fam. 2. Potoroinae. — Aepyprymnus and the other
genera placed in this sub-family are known by the vernacular
name of Eat -Kangaroos, or sometimes Kangaroo -Eats. The
latter term has been called " incorrect," though it is just as
good as the former, both of them in fact being inaccurate as
implying some likeness to or relation with a Eat. The present
genus has a partially hairy rhinarium ; the auditory bullae are
not swollen. It contains but one species, Ae. rufescens, a native
of Eastern Australia, which is distinguished by its very long
hind-feet.

Bettongia has long hind -feet as in Aepyprymnus, but the
rhinarium is entirely naked instead of being partially hairy,
while the ears are much shorter. The genus, which contains
four species, is remarkable as being the only ground -living
mammal with a prehensile tail, which it uses to carry grass, etc.

1 Proc. Zool. Soc. 1895, p. 131. 2 Ibid. 1884, p. 387.

3 Ibid. 1884, p. 407.

138 MUSK- KANGAROO

B. lesueuri burrows in the ground, often to so great a depth as 1 0
feet. The genus occurs in Tasmania as well as in Australia.

Caloprymnus, with one species, is a genus instituted by Mr.
Thomas in his Catalogue of Marsupials for a form (C. campestris)
which combines in a remarkable way the characters of Aepyprym-
nus, Bettongia, and Potorous. The external characters and the
general shape of the skull are as in Bettongia, while the molars
have the structure of those of Aepyprymnus. The last premolar
is as in Potorous.

Of the genus Potorous there are three species, which are
Tasmanian as well as Australian. Unlike the other Rat-
Kangaroos, the hind-feet are comparatively short, and the animal
is therefore less addicted . to jumping than its relatives. The
rhinarium is naked, and the ears are of fair length.

Sub-Fam. 3. Hypsiprymnodontinae. — The Musk-Kangaroo,
Hypsiprymnodon, is the last genus of the present family, and the
only genus of this sub-family. It is intermediate between the
Macropodidae and the Phalangeridae, the annectant character
being mainly the hind-feet, which though they have the same
long fourth digit as the Kangaroos, have it more feebly developed,
and possess also an opposable hallux, which is one of the salient
features in the structure of the Phalangeridae. The tail is naked
and scaly ; the rhinarium is entirely naked. The ears are large
and not furry. The single species, H. moschatus, appears to feed
upon insects as well as vegetables.

" Its habits are chiefly diurnal, and its actions when not dis-
turbed by no means ungraceful. It progresses in much the same
manner as the Kangaroo - Rats (Potorous\ to which it is
closely allied, but procures its food by turning over the de'bris in
the scrubs in search of insects, worms, and tuberous roots,
frequently eating the palm berries, which it holds in its fore-
paws after the manner of the Phalangers, sitting up on its
haunches, or sometimes digging like the bandicoots." This is
Mr. Ramsay's description of the animal, which he was the first
to discover.1

Fam. 2. Phalangeridae. — The genus Hypsiprymnodon bridges

over the not very wide gap which separates the Kangaroos from

the Phalangers. The Phalangers are Marsupials with five fingers

and toes ; the second and third toes are bound together by a

1 Proc. Linn. Soc. N.S. Wales, i. 1877. p. 34.

CHARACTERS OF PHALANGERS

139

common integument as in the Macropodidae. The hallux is oppos-
able and nailless. The tail is nearly always long and prehensile.
The pouch is well developed ; the stomach not sacculated ; a
caecum is present (except
in Tarsipes}. These are
really the principal dis-
tinctions between the two
families. In addition, it
may be mentioned that the
lower incisors have not a
scissor-like action as in the
Kangaroos.

The Phalangers may be
divided into four sub-
families.

The first of these, that
of the PHALANGERINAE,
contains the genera Phal-
anger (including Cuscus),
Acrobates, Distaechurus,
Dromicia, Gymnobelideus,
Petaurus, Petauroides,
Daetylopsila, Pseudocliirus,
and Tricliosurus.

These genera agree in
the following generalities: —
Tail well developed, often
very long ; three incisors
above, and at least two pre-
molars both above and
below ; caecum long and FIG
simple ; stomach without
a cardiac gland ; liver not
very complicated by second-
ary furrows, with a distinct
caudate lobe; the vaginal median culs-de-sac often coalesced;
lungs with an azygos lobe.

The second sub-family, PHASCOLARCTINAE (with the Koala only),
is thus characterised : — Tail rudimentary ; cheek-pouches present ;
superior incisors three, but only one premolar above and below ;

68. — Bones of leg and foot of Phalanger. ast,
Astragalus ; calc, calcaneum ; cub, cuboid ;
ect.cun, ecto-cuneiform ; ent.cun, ento-cunei-
form ; fb, fibula ; mes.cun, meso-cuneiform ;
nav, navicular ; tib, tibia ; 7- V, first to fifth
toes. (After Owen.)

140 CUSCUS AND TRUE PHALANGERS CHAP.

caecum extraordinarily long ; stomach with a cardiac gland ; liver
complicated by additional furrows, without a free caudate lobe ;
no azygos lobe to lungs : vaginal culs-de-sac free.

The third sub -family, PHASCOLOMYINAE, contrasts with the
others as follows : — Tail rudimentary ; cheek-pouches present, but
rudimentary; one incisor on each side above, but no additional
pre molars ; all the teeth rootless ; caecum not peculiar in shape ;
stomach with a cardiac gland ; liver complicated by secondary
furrows, without a free caudate lobe ; lung with an azygos lobe :
vaginal culs-de-sac free.

The last sub -family, TARSIPEDINAE, is thus defined: — Tail
long ; tongue extensile : only one premolar ; molars reduced ;
caecum absent.

Sub-Fam. 1. Phalangerinae. — The genus Phalanger embraces
five species, sometimes called by the generic name of Cuscus, They

FIG. 69. — Vulpine Fhalanger. Trichosurus mdpecula. x J.

are largish animals with short ears ; only the end of the tail
is naked. Of these animals only one species is found in Australia
itself, the rest inhabiting the islands lying to the north. The
Spotted Cuscus, Ph. maculatus, is in spite of its vegetarian diet.
and perhaps on account of its spots, spoken of as the " Tiger
Cat." Mr. Aflalo remarks of it that though provided with a
prehensile tail, it is little better as a climber than the tailless
Koala.

Trichosurus, including the " True Phalangers," includes largish
species, which can be distinguished from the last genus by a
chest-gland similar to that which occurs in Myrmecolius and
some other Marsupials of the present group. There are but
two species, which are purely Australian. The "Brush-tailed
Opossum," T. vulpecula (perhaps better known as Phalangista

vii MINUTE ARBOREAL FORMS 141

vulpina), like its American pseudo-namesake (a true Opossum,
genus Didelphys), " plays 'possum " on occasions. The dental
formula is I % C ^ Pm f M ^. The ears are shortish.

The Eing-tailed Phalangers, Pseudochirus, are more widely
distributed than the last two genera ; they range from Tasmania
in the south to New Guinea in the north. They are not, however,
ring-tailed, though the tip of the tail is generally white. As
in the last genera, which have prehensile tails, the end of this
appendage is naked. The mammae are four. The tooth formula
is I4C^ Pm ^ M ^. There are some ten species of the genus.

The Striped Phalanger, Dactylopsila trivirgata, is an animal
about a foot long, whose identity can be ascertained by its striped,
black and white skin. It is an arboreal creature that lives
apparently both on leaves and grubs like so many arboreal
creatures of quite different groups — Squirrels, for instance, and
New- World Monkeys. The tooth formula is I f C ^ Pm f M ^.

Gymnobelideus leadbeateri is a small creature with a body
6 inches in length. It is restricted to the colony of Victoria.
The general look is that of Petaurus ; the ears are naked.

Dromicia is a genus of Phalangers which although devoid of
a parachute, such as is possessed by certain genera that will be
considered immediately, is able to leap with great agility from
branch to branch. The ears are large and thin and almost
naked ; the tooth formula is I ^ C ^ Pm | M ^. They are minute
creatures, the longest measuring, with the -tail, but 10 inches.
Dormouse -Phalanger is a name sometimes given to them.
There are four species, ranging from Tasmania to New Guinea.
The name Dormouse as applied to the genus seems to be owing
to the way in which they hold a nut in the paws when feeding.
D. nana is 4 inches long, with a tail of nearly the same
length. It is thick at the base.

Distaecliurus is the last genus of non-flying Phalangers. Its
name refers to the arrangement of the hairs on the tail, which
are disposed on either side in a row like the vane of a feather.
The tooth formula is I -| C ^ Pm ^ M §, very nearly as in Aero-
bates. The ears are as in that genus.

Petaurus is the first genus of the Flying Phalangers, all of
which are provided with a parachute-like expansion of the skin
between the fore- and hind-limbs ; the ears are large and naked :
and the tooth formula is If C^Pmf M|. There are three

I42 STRUCTURE OF KOALA CHAP.

species of the genus, which extend through pretty well the
entire Australian region. The term " flying " as applied to these
and the other " flying " genera is of course an exaggeration. The
animals cannot fly upwards ; they can only descend in a skim-
ming fashion, the folds of skin breaking their fall. P. Ireviceps is
perhaps the best - known species. The body is 8, the tail 9
inches long.

Petauroides seems to be chiefly distinguished from Petaurus
by the fact that, as in its ally Dactylopsila, the tail is partly
naked terminally. In Petaurus and Gymnobelideus the tail is
bushy to the very end, including its extreme tip below.

A third genus of Flying Phalangers is the minute Acrobates,
which has a distichous tail like that of Distaechurus. It is not
more than 6 inches in length including the tail. As to these
Flying Phalangers it is exceedingly instructive to observe that
the same method of " flight " has been apparently evolved three
times ; for . the three genera are each of them specially related
to a separate type of non-flying Phalanger. The same observa-
tion can be made about the Flying Squirrels, Anomalurus and
Sciuropterus. The dental formula is I f C ^ Pm f M f. The
ears are thinly clad with hair. There are four teats.

Sub-Fam. 2. Phascolarctinae. — The Koala, or Native Bear,
Phascolarctos cinereus, is the only representative of its sub-family.
It is, like the Wombat, aberrant in the lack of an obvious tail.
The absence of this appendage is curious in an arboreal creature
whose near allies have a long and prehensile one. The structure
of the Koala .was investigated by the late Mr. W. A. Forbes.1
There are some unexpected points of likeness to the Wombat :
thus they agree in the absence of the tail, in the structure of the
stomach, and in the great subdivision of the lobes of the liver.
The brain, however, is smooth, and the caecum is exceedingly large
and complicated in structure, that of the Wombat being short.
That both animals have cheek-pouches is perhaps due to similar
habits of temporarily storing masses of food. This animal has
only eleven pairs of ribs. The tail has only seven or eight verte-
bra, and these have no chevron-bones.

A peculiarity of the skull is seen in the great size of the
alisphenoid bulla, which is comparable in size and appearance with
that of the Pig. As in the Kangaroos, the atlas is incomplete below.
1 "On some Points in the Anatomy of the Koala," Proc. Zool. Soc. 1881, p. 180.

HABITS OF KOALA

143

The tooth formula of the genus is I ^ C ^ Pin -£ M •£-*-•§-. The
additional lower molar seems to be exceptional, and has been found
in one specimen only.

In the alimentary tract the most remarkable structure is the
large intestine, which is very capacious for the first 28 inches or
so of its course. This section of the colon is lined with rugae
precisely like those which are found in the caecum. These folds,
whigh at first are some twelve in number, fuse lower down, and
by the time that the colon approaches the external orifice are
reduced to five. Similar folds, as already stated, occur in the
caecum, but do not extend as far as its blind end. The caecum
is proportionately and actually larger than in any other Marsupial.
The gall-bladder is unusually elongated.

The Koala is mainly crepuscular or nocturnal in its habits. It
feeds so exclusively upon the leaves of the gum-tree (Eucalyptus}

FIG. 70. — Koala. Phascolarctos cinereus.

that it is impossible to keep the creature long in captivity in
lands where that particular kind of food is not available.

The female, though she seems to bear but a single young one,
which is carried on the back after the fashion of some Opossums,
has two nipples. The animal's slow habits seem to require a
nocturnal and retired life. It is about as lethargic as the
Sloth, and it is said to further resemble that animal in clinging
firmly to a branch even after it is shot.

144 DESCRIPTION OF WOMBAT

Sub-Fam. 3. Phascolomyinae. — Pkascolomys, the Wombat, is
the only genus of this sub-family. This animal has the appearance
of a heavily-built Marmot, like which it has a mere stump for a
tail, and a pair of strong chisel-shaped and Rodent-like incisors,
which, however, differ from those of Rodents in having a complete
coating of cement. All the teeth of the animal are rootless, and
there are no canines. The incisors have enamel on the front and
lateral faces only. The dental formula is I ^ C $ Pm \ M -|. The
affinities with other Diprotodont Marsupials are shown by the
commencing syndactyly of the second and third toes. The

FIG. 71. — Wombat. Phascolomys toombat. x TV.

rhinarium is naked or hairy. There is a rudimentary cheek-
pouch, as in Phascolarctos. The Wombat has, like the
Koala, and also the Beaver — which does away with some of the
value of the comparison — a peculiar gland -patch in the
stomach, a raised area of collected glands. In no other Marsupial
is such a structure found, " whilst in the two forms under con-
sideration its identity is almost precise. That such a unique
structure should have been independently developed in two forms
unrelated to each other, appears to me to be in the highest degree
improbable." This is Mr. Forbes' opinion. It might be strength-
ened by adding the observation that, as there are other points of
likeness between the Wombat and the Koala, it seems more unlikely
that a structure so nearly identical should have been twice

VII

TARSIPES

145

developed in two not very distant forms. As in the Kangaroos,
the atlas is open below. Ph. ursinus has 15 ribs; the other species
the normal (for Marsupials) 13. Other points of likeness will be
mentioned under the description of the Koala. These animals

Fm. 72. — Skull of Wombat. Phascolomys wombat. (Lateral view. ) ang, Angular pro-
cess ; cond, coudyle of mandible ; ex.oc, exoccipital ; ext.aud, opening of bony
auditory meatus ; ju, jugal ; Icr, lachrymal ; max, maxilla ; nas, nasal ; p.max,
premaxilla ; sq, squamosal ; ty, tympanic. (From Parker and Haswell's Zoology. )

mainly feed upon roots ; they live in companies in burrows. There
are three species — Ph. ursinus, Ph. latifrons, and Ph. mitchelli.
Ph. ursinus is Tasmanian in range, the other two species South
Australian,

Sub-Fam. 4. Tarsipedinae. — The genus Tarsipes ought per-
haps to be removed from the present family. There is but a
single species, which is a small creature of 7 inches in total
length, of which the tail measures 4 inches. The teeth are
much dwindled, the formula being I f C £ Pm £ M f = 22. The
lower incisors are procumbent. The lower jaw, moreover, has
not the characteristic Marsupial inflection. The intestinal canal
is without the caecum present in the remaining Phalangeridae.
It is a curious fact that this aberrant little Phalanger should
come from Western Australia, like the even more aberrant
Myrmecobius. Like the latter also, Tarsipes has a long exsertile
tongue, with which, however, it extracts honey from flowers.
Probably it also catches minute insects in the corollas of the
flowers. It has been proved, in fact, that in captivity at any
rate the animal is insectivorous ; for it has been known to
eat moths.

Fam. 3. Epanorthidae. — The extinct Epanorthidae of Pata-

VOL. X L

•*.- •

146 GIANT EXTINCT FORMS

gonia are represented to-day by a small Marsupial which has
been rediscovered within the last two or three years. This little
animal, formerly called Hyracodon (a pre- occupied name), is now
termed Caenolestes, and is a native of Colombia and Ecuador.
There are two species, and of these C. obscurus is called by
the inhabitants " Eaton runcho," which means opossum-rat. It
lives apparently upon bird's eggs and small birds, though it
belongs to the Diprotodont division of the Marsupials. Caenolestes,
however, although diprotodont, has not the syndactylous character
of the digits of the feet already referred to in the Kangaroos and
their allies. The pouch is small and rudimentary. The denti-
tion is I | C^ Pm -|M| = 46, and the teeth are said by Mr.
Thomas to be much like those of the Australian Dromicia.1

In the skull a peculiarity which does not bear upon its
affinities to other Marsupials, but is still interesting, is mentioned
by Mr. Thomas. The nasals are not sufficiently prolonged to
meet the upper edge of the maxillae, and so a vacuity is left, as
in the skulls of many Euminants (e.g. the Sable Antelope). The
palate is very imperfect ; the foramina, which render it so, reach
as far forward as the last premolar. The lower jaw has quite
the appearance of that of a Macropus or Phalanyer, with long
and forwardly projecting incisors.

Extinct Diprotodonts. — The great Diprotodon is a creature
with a skull a yard long, which must have been of the size of a
large Ehinoceros. Though closely allied to Macropus, it seems
that this great beast did not hop after the fashion of a Kangaroo,
its limbs being of a more equal size than in the Kangaroo.
Eecently some further remains of Diprotodon have been discovered
in a lake known as Lake Mulligan, where they had apparently
been bogged. Professor Stirling has contributed an account of
these remains, which fills up a considerable gap in our knowledge.
He has been able to state the structure of the fore- and hind-limbs.
Both limbs are pentadactyle, the fingers of the fore-lirnb being
approximately equal in length and general development. In the
hind-limb the hallux is small, and consists of the metatarsal only.
This bone is fixed in the position of " extreme abduction," and is
suggestive of an arboreal limb. Digits two and three may have

1 Thomas, "On Caenolestes, a still existing survivor of the Epanorthidae of
Ameghino, and the representative of a new family of recent Marsupials," P.Z.S.
1895, p. 870.

THYLACOLEO

147

been syndactylous, and the authors of the account 1 of these bones
think that the fourth toe may have shared in this syndactyly.
The metatarsal of the fifth digit is enormously expanded at its

FIG. 73. — Diprotodon australis. (After Owen.)

edge, and seems to have furnished a strong support to the
creature ; this is also seen in the metacarpal of the fore-limb.
Probably, therefore, Diprotodon was quadrupedal in its mode of
progression, with the em-
phasis laid upon the
little finger and the
little toe instead of, as
in ourselves, the first
toe The hind-foot of
the Diprotodon could
not be more unlike that
of a Kangaroo than it
actually is.

Another giant among
these Marsupials was the
genus Thylacoleo, whose name was given to it by Sir Eichard
Owen on the view that it was a Marsupial Tiger. Sir W. Flower
has, however, controverted this opinion, and the genus is in fact,
in spite of its large size, closely allied to the Phalangers and

FIG. 74. — Thylacoleo carnifex. Side view of skull.
(After Flower.)

1 Stirling and Zietz, Mem. Roy. Soc. South Australia, i. ; see also a notice in
Nature, January 18, 1900.

148

NOTOTHERWM

Cuscuses.1 The dental formula is I ^ C ^ Pm ^ M \ ; the last
premolar is a great blade-shaped tooth like that of Potorous.

Nototherium was a creature smaller than Diprotodon, but still
of large size ; it is believed to have been a burrowing creature,
and to connect the Wombats with Diprotodon. More certainly
allied to the existing Wombat was Phascolonus, a Wombat as
big as a Tapir.

Of extinct American Diprotodonts the Epanorthidae, already
referred to in connexion with the living Caenolestes, were the most

FIG. 75. — Nototherium mitchelli. Side view of skull, x £. (After Owen.)

prominent forms. The genus Epanorthus occurs in the Santa Cruz
formation of Patagonia, which is believed to be Miocene. The
incisors are three in the upper jaw ; and the single incisor of each
ramus of the lower jaw is a great chisel-shaped, cutting instrument.
Abderites is also typically Diprotodont by reason of the large
projecting incisors of the lower jaw. It has a large cutting
tooth in the lower jaw, which appears to be the last premolar,
and is thus comparable to the great cutting tooth of the lower
jaw and of the upper jaw of the extinct Phalanger, Thylacoleo.

1 Quite recently (Proc. Linn. Soc. N.S. W. 1898, p. 1) the carnivorous character
of Thylacoleo has been reasserted by Mr. Broom.

vii CARNIVOROUS MARSUPIALS 149

It may also be comparable to the great premolar of such Multi-
tuberculata as Ptilodus and Plagiavlcuc. It is, moreover, marked
with vertical grooves.

An interesting form, which is unfortunately but little known,
is the Australian and Pleistocene genus Triclis, with one species,
T. oscilhlns. In having a minute canine tooth in the lower jaw
it agrees with some Phalangeridae, and being otherwise closely
allied to Hypsiprymnodon, it unites the Macropodidae with the
Phalangeridae.

SUB-ORDER 2. POLYPPtOTODONTIA.

In this mainly carnivorous or insectivorous division of the
Marsupials the incisors are four or five on each side of the upper

FIG. 76. — Front view of the skull of Tasmanian Devil (SarcophUus -ursinus), showing
Polyprotodont and carnivorous dentition. (After Flower.)

jaw, and one or two fewer in the lower jaw. Figs. 76 and 77
illustrate the Polyprotodont and Diprotodont dentitions. The
canines are those of flesh-eaters and so are the molars, being as a
rule sharply cuspidate. As a rule, which has an exception in the
Peramelidae, there is no syndactylism of toes in the hind-foot.
This sub-order is at the present day Australian and American in
its range.

Fam. 1. Dasyuridae. — This family consists of Marsupials
which are generally pentadactylous, but with occasionally the
hallux missing The tail is long but not prehensile. The pouch
is present or absent. The teeth vary in the different genera, but

TASMANIAN WOLF

the upper incisors are never less than three, and may be as many

as five in the upper jaw and
six in the lower. The canines
trenchant. There

are

is no

caecum.

The genus Tliylacinus con-
tains but a single species,
which is now limited to Tas-
mania, and is generally known
as the Tasmanian Wolf. It has
the build of an ordinary Wolf,
and is of about the same size.
The hinder part of the body is
marked with a series of black
transverse bands. The hallux
is entirely wanting; the pouch

FIG. 77.— Front view of skull of Koala (Phas- ,J an.

colarctos cinereus), illustrating Diprotodout Opens backwards. Ihe mar-
and herbivorous dentition. (From Flower.) SUpial bones are minute and

unossified. The dental formula is I f C l Pm f M f = 46. There

T>a

xo

FIG. 78. — Longitudinal section of the skull of the Thylacine (Thylacinus cynocephalus).
x \. a, Angular process of mandible ; A f!, alisphenoid ; BO, basioccipital ; SS,
basisphenoid ; cd, condyle of mandible ; ET, ethmoturbinal ; Ex. 0, exoccipital ;
•Fr, frontal ; ME, ossified portion of mesethmoid ; MT, maxilloturbinal ; MX,
maxilla ; No,, nasal ; OS, orbitosphenoid ; Pa, parietal ; Per, periotic ; PI, pala-
tine ; PMx, premaxilla ; PS, presphenoid ; Pt, pterygoid ; SO, supraoccipital ;
Sq, squamosal ; Vo, vomer. (From Flower's Osteology.)

are four mammae. This animal, now confined to Tasmania,

DASYURES

is getting rarer on account of its sheep -killing propensities,
and the consequent war of extermination declared upon it by the
colonists. It will, however, feed upon other animals; and it is
related that the first specimen ever captured had in its stomach
the remains of an Echidna ! Mr. Thomas thinks that the persist-
ence of this and of some of the other larger carnivorous Marsu-
pials in Tasmania after their extinction in Australia is not uncon-
nected with the advent of the Dingo. But it is stated that the
Thylacine is quite capable of keeping even a pack of dogs at bay.
The genus Sarcophilus has been frequently confounded with
the next, but it is kept apart by Mr. Thomas, who follows

FIG. 79. — Tasmanian Devil. Sarcophilus ursinus. xTV

Cuvier in this. An alternative generic name is Diabolus,
which, like the first name, refers to the habits and character
of the single species which this genus contains. The genus is
more like Thylacinus than is Dasyurus. The hallux is wanting,
and the teeth, though fewer in number (42), resemble those of
the Thylacine more closely than do those of the Dasyure. The
species is called S. ursinus, the popular name being Tasmanian
Devil. It is black with a variable number of white patches on
the body. It is of about the size of a Badger, and is, like the
Thylacine, a nocturnal animal. The Tasmanian Devil is said to
be one of the most ferocious of animals, and to express its
ferocity by a " yelling growl."

The next genus of this family, Dasyurus, comprises five
species, which range over the whole of the Papuan and Australian
sub-regions. The general form is Viverrine, and the hallux is
sometimes present though small. The dental formula is as in the

152

FOOD OF DASVURES

CHAP.

last genus, but the teeth " are more insectivorous in their
character." There are six or eight mammae. The members of
this genus are grey or brown, and spotted with white ; they are

te

p.maoc

s.oc

FIG. 80. — Skull of Dasyurus. (Lateral view.) al.sph, Alisphenoid ; ang, angular process
of mandible ; fr, frontal ; ju, jugal ; Icr, lachrymal ; max, maxilla ; nas, nasal ;
oc.cond, occipital condyle ; par, parietal ; par.oc, paroccipital process ; p.max,
premaxilla ; s.oc, supraoccipital ; sq, squamosal ; sq', zygomatic process of squa-
mosal. (From Parker and Haswell's Zoology. )

all arboreal, and feed largely upon birds and their eggs. Mr.
Thomas has pointed out that in two species, D. viverrinus and
D. geoffroyi, the striae upon the foot-pads are absent, and that

FIG. 81. — Dasyure. Dasyurus riverriniis. x \. (After Vogt and Specht.)

therefore these at least are probably not so purely arboreal as
the rest. The animals are not diurnal, and during the day hide
themselves in the hollow trunks of trees. They are spoken of
as " Native Cats," but have the general habits of Martens. D.
maculatus is common in Tasmania, but is rare in Australia, thus
" approaching the condition now exhibited by the Thylacine and

POUCrtED JERBOA I 5 3

Tasmanian Devil, namely, complete extermination in Australia,
where both once lived." D. hallucatus shows an approach to
Phascologale in its five-toed hind-feet and slender build.

Pliascologale is a gemis which, like the last, is usually arboreal
(although not P. virginiae of North Queensland), but is of much
smaller size, the species not exceeding the dimensions of a rat.
They have no spots, but there is sometimes a stripe down the back.
There are thirteen species, which have the same range as the last
genus. The hallux is present though small, but the pouch is
" practically obsolete," though there is a small fold of skin behind
the teats. The rhiuarium is naked ; the tail is long, " bushy,
crested, or nearly naked." The mammae are four to ten in
number. The dental formula is as in Dasyurus, and the teeth
are not very different in form ; sometimes the last premolar is
wanting. " The members of this genus," remarks Mr. Thomas,
" evidently take the place in the Australian region filled in the
Oriental by the Tupaiae, and in the Neotropical by the smaller
Opossums."

The genus Sminthopsis comprises not more than four species,-
even smaller than the last. The largest species, S. mrginiae, is
only 125 mm. in length. The hallux is present, and there is
a well -developed pouch. There are forty -six teeth, as in the
Dasyures. The feet are narrow with granulated or hairy soles,
whereas in Phascologale they are broad with smooth soles. The
mammae are eight or ten. The genus ranges through Australia
and Tasmania.

The genus Antechinomys has but a single species, which is a
native of Queensland and New South Wales. The build is
Jerboa-like, and the animal is, as might be inferred, terrestrial.
The ears are very long, and the limbs elongated ; the hallux is
absent ; the teeth are exactly as in Sminthopsis.

Antechinomys has thirteen dorsal and seven lumbar verte-
brae ; three sacrals and twenty-five caudals, the latter number
being in excess of that of its allies. The stomach is. nearly
globular, with approximated orifices ; the intestine measured
6 '8 inches, a little more than twice the length of the animal
itself. A. lanigera is a native of East Central Australia, and
appears to be entirely terrestrial in habit, and to progress by a
series of leaps — at any rate when going at full speed.

Professor Spencer, who found examples of this rare species, gives

154 MYRMECOBWS

an interesting description of its habits. Antcchinomys has much
the look of the Australian Rat, Hapalotis mitclielli ; uiid as the
two animals lead a similar kind of life, the resemblance is not
unexpected. Professor Spencer wonders why these creatures are
saltatory in habit. The country which they inhabit is arid, but
with patches of grass and shrubs. For a big kangaroo the
advantage of the power of leaping over such obstacles may be
obvious, but not for the small and slender Antechinomys. The
chief foes of this rare Marsupial appear to be predatory birds ;
and Professor Spencer thinks that the saltatory mode of pro-
gression may be more baffling to such pursuers than even a
rapid run.

The genus Dasyuroides has been lately instituted by Professor
Spencer for a Marsupial from Central Australia somewhat in-
termediate between Sminthopsis and Phascologale. As there is
but one species, the generic will be considered with the specific
characters. D. hyrnei is an animal of about the size of the
Common Eat. The hallux is absent. The tail is fairly thick,
but not " incrassated." There are six mammae, and the pouch is
but slightly developed, with two low lateral folds. The dentition
is I ^ C ^ Pm |^ M |. This Marsupial is nocturnal, and burrow-
ing in habit. Its food consists of insects.1

Myrmecolnus is so different from the last- described genera (!)ASY-
URINAE) that it is usually separated from them as a sub-family
MYRMECOBIINAE. The animal is of a bright rufous colour, banded
posteriorly with white. There is no hallux, though the metatarsal
belonging to that digit is present. There are four mammae.2
On the chest is a naked patch of some extent, upon which open
the ducts of a complex gland, which has been described and
figured by myself.3 There is no pouch, but a tract of skin
shows indications of a pouch - like structure. The teeth are
extraordinarily numerous, fifty to fifty-four ; the formula being
I g^f) C ^ Pm § M ^. Their resemblance to those of certain Jur-
assic Marsupials is dealt with on p. 100.4 In this matter lies of

1 Horn Scientific Expedition, pt. ii. Zoology, 1896, p. 36.

2 Leche found five, and Waterhonse stated eight to be the number.

3 Proc. Zool. Soc. 1887, p. 527. See also Leche, Biol. Foren. Forhandl. 1891, p.
136, and literature quoted.

1 Traces of horny pads, like those of the Duck-bill, have been asserted to exist
in this animal. This is exceedingly interesting when regarded in conjunction
with its multituberculate molars.

AMERICAN OrOSSUMS

155

course the chief interest of the genus, which may be " an un-
modified survivor from Mesozoic times, and therefore from a
time long before the Didelphyidae, Peramelidae, and Dasyuridae
were differentiated one from the other." Another ancient feature

Fig. 82. — Banded Aiistraliaii Anteater. Myrmecobius faaciatus. x i.

(found in Jurassic mammals) is a mylo-hyoid groove upon the
lower jaw, which, however, is not always present, and its exist-
ence has therefore been denied. The single species, M.fasciatus,
is partly arboreal and partly terrestrial in habit, and feeds upon
ants. It is a Western and Southern Australian form.

Fam. 2. Didelphyidae. — All the members of this family are
pentadactylous. The
teeth are fifty in number,
arranged thus : I % C 1
Pm | M A. The caecum
is small ; the pouch is
generally absent ; the
tail generally long and
prehensile.

The genus Didelphys
contains most of the
forms belonging to this
family, including as it
does some twenty -three
species. The Opossums
are mainly arboreal ani-
mals, insectivorous in FIG. 83. — Virginian Opossum. Didelphys virginiana.
, i • /• i • i , , i i XT- (After Vogt and Specht.)

their food ; but the larger

species eat reptiles, birds, and their eggs,
species carry their young, when able to

Several of the small
leave the teats, on

156 BANDICOOTS

their back, the tails of the young being wrapped round that
of the mother. It is not only the pouched species which
carry their young in something of this fashion. Azara's
Opossum, an animal as big as a cat, is said to carry its eleven
young ones (themselves as large as rats) on the back, though
their foothold does not appear to be strengthened by inter-
twining the tails. Even with this huge family on her back, the
mother can climb trees with considerable alacrity. The mammae

FIG. 84. — Thick tailed Opossum. Diddphys crassicaiulata.

are seven to twenty-five in number. The genus has been lately
split up into a number of genera, Marmoset,, Dromiciops,
Peramys, etc.

Chironectes is hardly different from Didelphys. It has
webbed hind-feet, and is aquatic in habit. The one species of
the genus is known as the Yapock, and is a Central and South
American form. It is of about the size of a large rat, and appears
to be an expert diver after the fish upon which it lives.

Fam. 3. Peramelidae. — The Bandicoots, although clearly be-
longing to the Polyprotodont Marsupials, yet agree with the
Diprotodonts in the fact that the second and third toes of the
feet are bound up in a common integument, which is not
the case with the Diprotodont Caenolestes. The hind-feet are
longer than the front ; of the former limb, two or three of
the fingers alone are long and functional ; the others are rudi-
mentary or absent. Tail long, hairy, and non-prehensile. Denti-
tion 1 1 - C ^ Pm j| M ^ = 48, or sometimes, owing to the absence
of a pair of upper incisors, 46. There is a caecum.

The genus Peragale, the Rabbit-Bandicoots, consists of two

RABBIT BANDICOOT

157

species entirely Australian in range. The enormous ears (whence

FIG. 85. — Bones of maims.
A, of Choerqpus cas-
tanotis. x 2. B, of
Bandicoot (Peramdes).
x l^. c, Cuneiform ;
I, lunar ; m, magnum ;
It, radius ; s, scaphoid ;
td, trapezoid ; tm,
trapezium ; u, unci-
form ; U, ulna ; I- V,
digits. (From Flower's
Osteology.)

" Babbit " Bandicoot) distinguish this genus from Perameles.

FIG. 86. — Rabbit Bandicoot. Perayale lagotis. x I.

The pouch opens backwards, and there are eight mammae. P.
lagotis, the only species about whose ways of life anything is

PIG-FOOTED BANDICOOT

CHAP.

known, burrows in tjie soil, whence it extracts grubs ; it is also
a grass-feeder, and it is said that its likeness to a Rabbit in
appearance is strengthened by its similarity in flavour !

Perameles is a genus consisting of twelve species, which are
found in Tasmania, Australia, and New Guinea. Like the last
genus, from which it does not widely differ in other points,
Perameles consists of species which combine insectivorous and
vegetarian habits. One species is said to become in captivity an
expert in catching mice. The pouch opens backwards, and there
are six or eight mammae.

The last genus of this family is Choeropus, containing but
one species, Oh. castanotis. It is confined to the Australian con-

FIG. 87. — Pig-footed Bandicoot. Choeropus castanotis. x J.

tinent. It is to be distinguished from the last two by the fact
that there are only two functional digits, the second and third,
in the fore-limb ; the fourth is rudimentary ; the other two are
absent. It burrows, and is omnivorous like its allies. The two
metacarpals that are developed are very long and closely apposed ;
they have hence a remarkably pig-like aspect, a*nd justify its
name. The pouch opens backwards, and there are eight mammae.
Fam. 4. Notoryctidae. — This family contains but a single
genus and species, the recently-discovered Notary ctes typhlops)

1 See for an account of this animal, Professor Stirling's Memoir in Trans. Roy.
Soc. S. Australia, 1891, p. 154, and Gadow, Proc. Zool. Soc. 1892, p. 361.

MARSUPIAL MOLE 159

We may regard as family-characters the pentadactyle limbs, the
existence of three pairs of incisors in the lower and four in the
upper jaw ; and the tritubercular nature of the upper molars.
Notoryctes typhlops, the " Marsupial Mole " as it has been termed,
was originally discovered by Professor Stirling in Central South
Australia. It is a burrowing creature, clothed in a silky fur of a
pale golden red, without external ears. It has been compared in
appearance with Chrysocliloris, the Cape Golden Mole, and the
eminent palaeontologist, Professor Cope, has even insisted upon a
real genetic affinity. Edentate affinities have also been suggested.
But Notoryctes has a small pouch opening backwards as in other
Polyprotodonts,1 and as it also possesses marsupial bones it must

*

FIG. 88. — Australian Marsupial Mole. Notoryctes typhlops. x \.

undoubtedly be referred to the Marsupialia. The animal shows
many curious adaptations to its underground mode of life.
Certain of the vertebrae in the neck and in the lumbar
region are firmly welded together, giving of course a strength
of push, and suggesting the Armadillos ; the claws of the third
and fourth front-toes are greatly enlarged, and must be efficient
digging organs. The track of the animal is like that of a rail-
way in mountainous country ; it burrows for a short distance,
emerges, and then descending beneath the surface re-emerges.
The red colour of the fur is said to be in harmony with the
arid soil in which it lives. The native name of the creature is
" Urquamata." It feeds upon ants and other insects.

Extinct Polyprotodonts. — Of extinct Polyprotodonts (apart
from those Mesozoic forms which are considered on p. 100)
extinct species of Thylacinus and Dasyurus are known from

• 1 The male, according to Professor Spencer, has a rudimentary pouch. •

160 EXTINCT POLYPROTODONTS CHAE- vn

Australia. The most interesting fact in connexion with the
Tertiary Polyprotodonts is the existence in South America of such
genera as Prothylacinus and Amphipromverra, which are not
merely Polyprotodonts but definitely Dasyures, and not referable
to the Didelphyidae.

These forms have been included in an order, SPARASSODONTA.
But it is not by any means certain whether these forms are
rightly placed in the neighbourhood of the carnivorous Marsupials ;
it is possible that they ought to be relegated to the Creodonta or
to their allies. Their structure is in fact somewhat intermediate
between those two groups. The teeth seem to be carnivorous and
Marsupial -like inform; but as already mentioned, in connexion
with the general structure of teeth, more than a single premolar
is replaced. These animals in fact, in so far as regards their
teeth, are midway between the Marsupials and the typical Eutheria.
The angle of the lower jaw is inflected, but the palate is not
marked by deficient ossification. At least this is not the case
with all the members of the group. Whether the small Micro-
biotherium, which is made the type of a family, is rightly referred
here is not certain. This animal had palatine vacuities as well
as an inflected angle to the lower jaw.

CHAPTER VIII

EDENTATA GANODONTA

Order II. EDENTATA

TERRESTRIAL, partly subterranean, or arboreal creatures of quite
small to gigantic size (some extinct genera), with frequently a
covering of scales or bony scutes. Limbs clawed. Teeth either
totally absent or, if present, imperfect in structure, being with-
out enamel, and not forming a complete series ; incisors and
canines being as a rule absent. Teats axillary, pectoral, or
inguinal.1 Eetia mirabilia very common in the extremities.

To this group the name of Bruta was given by Linnaeus,
but then it included not only the families which we now place
in the modern order Edentata, but also the Elephant and the
genus Trichechus. Mr. Thomas has proposed to change the
name into Paratheria, wThich name is suggestive of what he and
some others think concerning the systematic position of the
group, i.e. that it is not to be placed in the Eutherian group of
mammals at all, but represents a separate twig which has arisen
with the Eutheria from a low mammalian stock. This view can
hardly be accepted if the Ganodonta — which will be treated of
presently — be really ancestral Edentates, for they are not in any
way a Prototherian mammalian group, so far as their remains
enable us to judge.

The Edentata contain the Sloths, Ant-bears, Armadillos,
Manis and Orycteropus, among living forms. The great Ground-
Sloths, Megatherium, etc., and Armadillos, Glyptodon, etc., repre-
sent the extinct forms.

The name that has been applied to this group is inappropriate

1 Pectoral and abdominal in the Armadillo Tatusia.
VOL. X M

1 62 TEETH OF EDENTATES CHAP.

inasmuch as many Edentates have teeth. It is, however, by
a number of small tooth -characters that the order can be
defined. Thus if teeth are present they are simple in struc-
ture, without enamel in the adult condition, though a rudi-
mentary enamel -organ has been discovered in an Armadillo.
The teeth, moreover, are not found in the anterior part of the
mouth, and they grow from persistent pulps ; neither is there
much differentiation among them. It is not possible, however,
to speak of the Edentates as quite homodont, since in Orycteropus
there are large cheek-teeth ; but there is at any rate not a marked
heterodonty in that or in any other Edentate. It used to be said
that the Edentates were monophyodont. But the Armadillo
Tatusia was subsequently found to possess a second suppressed
dentition, and after this discovery Mr. Thomas proved that
Orycteropus is also diphyodont. Since then other Armadillos
have been shown to be diphyodont ; and the whole group there-
fore, so far as concerns those members that have teeth, may in all
probability be regarded as typically mammalian in this respect.

These characters are slender enough, but there seem to be
no others by means of which the members of this order can be
satisfactorily linked together. The fact is, that we have here
a polymorphic order which contains in all probability repre-
sentatives of at least two separate orders. We have at present
a very few, and these perhaps highly modified, descendants of a
large and diverse group of mammals. For convenience' sake they
will be all treated of under the head of Edentata.

Although for the probable reasons already stated it is a
hard matter to frame such a definition as will include all
existing Edentates, it is easy enough to define two groups in this
heterogeneous order; to define one group we should say, rather,
and then to regard the leavings as forming another not so easily
definable a group.

The perfectly - definable group is that which includes the
American Anteaters, the Armadillos, and the Sloths. In all
these creatures, which may certainly be regarded as representing
on their own account as many family types, there are a number
of important and highly-characteristic anatomical features which
they share in common. So exceedingly different are these three
types in general appearance and (correlated with that) in way of
life that these common characters acquire increased importance.

VERTEBRAE OF ANTEATERS

The first of these characters is the series of additional
zygapophyses on the posterior dorsal and lumbar vertebrae ;
these are very clear in the Anteaters and Armadillos ; less clear,
but still obviously represented, in the Sloths. In the second
place, they all possess a clavicle, rudimentary, it is true, in the

tc

FIG. 89. — Great Anteater (Myrmecophaga jubata). A, Side view of twelfth and thirteenth
thoracic vertebrae. B, Posterior surface of second lumbar vertebra. C, Anterior
surface of third lumbar vertebra, x |. az, Anterior zygapophysis ; az1, az'2, az:i,
additional anterior articular facets ; cc, facet for capitulum of rib ; in, ineta-
pophysis ; pz, posterior zygapophysis ; pzl, pz2, pz3, additional posterior articular
facets ; t, transverse process ; tc, facet for articulation of tubercle of rib. (From
Flower's Osteology.)

Great Ant-bear, but still present. Thirdly, the testes are
abdominal throughout life, a character which they share with
such lowly -organised animals as the Monotremata and the
Whales. Finally, and this is by no means a matter to be over-
looked, not only are all the existing members of this group
American in range, but there is no evidence to prove that they
have ever existed elsewhere. No European or Old- World repre-

164

AMERICAN EDENTATES

CHAP.

sentatives have as yet been discovered which can be referred to
the Anteater, Armadillo, or Sloth type with certainty.1

Of these American forms, which will be treated of first, the
Armadillos are further apart from either Sloths or Anteaters
than the last two are from each other. The name XENARTHKA
has been suggested for the American Edentates with " abnormal "
vertebral articulations ; the corresponding NOMAETHRA includes
the Old-World forms.

Between the Sloths and Anteaters the extinct Megatherium
and some of its allies are to a certain
extent intermediate. But it may be
pointed out in the first place that there
are certain important resemblances be-
tween the living forms. In both, retia
mirabilia are developed in the tail (in
spite of its reduction in the Sloths) and
in the limbs. But, as is well known,
retia are also found in other mammals
far removed in the series from these
under consideration. The reproductive
organs generally are very similar, and
they have both a dome -shaped and
deciduate placenta. The latter char-

FIG 90 '.-RigM scapula and clavi- acter they share with the Armadillos
cte or Two-todd Biota (CAogogHM . J

a, Aero- and with the Aard Vark ; Manis hav-

inS a *on-d<*iduate Placenta which is,
like that of the Carnivora, zonary in
form- The Edentates, at any rate the
American forms, have a double vena
cava posterior and no azygos vein. This condition is also met
with among Whales.

Osteologically the Sloths and Anteaters are united by the
fact that the coracoid becomes fused with the coracoid border
of the scapula, thus forming a foramen ; the importance of this
character is, however, discounted by its occurrence in three
genera of Cebidae.

The above facts embody the views of Sir William Flower.2

1 A rather problematical Armadillo, Necrodasypus, has been recorded from
French strata. It consists of a few scutes only.

2 Proc. Zool. Soc. 1882, p. 358.

x i

coraco- scapular foramen

vni BRAIN OF EDENTATES 165

A subsequent study of the brain and of the muscles of these
animals has led to results not entirely in harmony with these
views.

Dr. Elliot Smith is of opinion,1 after an exhaustive study of
the Edentate brain, that in this region of the body the present
group shows very decided points of likeness to the Carnivora ;
that is, so far as concerns the Anteaters. On the other hand,
Orycteropus is as distinctly comparable with a primitive Ungulate
type, such as is exemplified by Moschus. " If the brain of
Orycteropus" he remarks, " were given to an anatomist
acquainted with all the other variations of the mammalian
type of brain, there is probably only one feature which would
lead him to hesitate in describing it as an exceedingly simple
Ungulate brain. That one feature is the high degree of
macrosmatism.2 Manis, on the other hand, does not come especially
near to Orycteropus. The brain of Manis conforms to a simple
type of architecture, which agrees in many points with both those
of Orycteropus and the American Edentates ; there is not sufficient
evidence to show which type it really favours." Elliot Smith
would, in fact, agree with Max Weber that it is better, if a
division is to be made, to divide the group into three orders : —
the Xenarthra (Sloths, Anteaters, and Armadillos), Tubulidentata
(Orycteropus), and Squamata (Manis}, instead of into Xenarthra
and Nomarthra.

Messrs. Windle and Parsons3 are disposed to see in muscular
similarities reasons for uniting Manis with the American Edentates,
though they confess' to being unable to place Orycteropus; in
this animal, they say, " we are more struck by the generalised
mammalian arrangement of its muscles than by any special
Edentate characters. There are, however, two muscles in Oryctero-
pus which show peculiarities not found elsewhere than in the
Edentates " ; — the triceps, which has more than one scapular head,
and the tibialis posticus, which is double. They conclude that
Orycteropus "presents some feeble claims to be taken into the
order."

We shall here adopt the following divisions.

1 Trans. Linn. Soc. (2) vii. 1898, p. 277.

2 i.e. large olfactory lobes.

3 Proc. Zool. Soc. 1899, p. 1014.

I 66 AMERICAN ANTEATERS

SUB-ORDER 1. XENAKTHKA.

Fam. 1. Myrmecophagidae. — The family Myrmecophagidae
contains three genera, all South American in range. These
genera, Myrmecophaga, Tamandua, and Cycloturus, agree greatly
in their outward form. They are all without teeth, and have
long snouts and long protrusible tongues. The fur is thick, and
they have powerful claws wherewith to break down the strong
ant-hills upon whose inhabitants they feed. Tamandua and
Cycloturus are arboreal, Myrmecophaga is terrestrial in habit,

FIG. 91. — Great Anteater. Myrmecophaga jubutu. x -fa.

The claws of the arboreal forms are useful to destroy the bark,
and thus bring to light insects which lurk in such situations.

The genus Myrmecophaga contains but one species, the Great
Anteater, Myrmecophaga jubata. It is a large and handsome
animal, with long, shaggy, greyish-black hair and a broad white
stripe across the shoulder. The coloration is similar in the two sexes.
Including the long and bushy tail it reaches a length of over
7 feet. It is on account of its long tongue and greatly developed
salivary glands that this and the allied genera were originally
placed with Manis. It is the submaxillary glands which are so
enormous ; they extend back over the chest, and open by three
distinct ducts, of which two unite just before the external orifice.

vin TAMANDUA AND CYCLOTURUS 1 67

Along their course these ducts are provided with a sphincter
muscle, which squeezes the secretion towards the external orifice
into the mouth-cavity. The stomach is somewhat gizzard-like.
The intestine has no caecum.1

The Anteater's great claws are not only serviceable in tearing
up the ground to get at its food ; armed with them he does not
fear, as Mr. AVaterton remarked, " the fatal pressure of the
serpent's fold or the teeth of the famished jaguar." An Anteater,
too, is more than a match for a big dog, and will rip open its
belly with the claws while the dog is vainly trying to make an
impression with its teeth upon the shaggy hair.

Tamandua is a smaller animal than Myrmecophaga, and, as
has been stated, is arboreal ; associated with this habit is a pre-
hensile tail. Like the last genus, Tamandua has a rudimentary
clavicle, this bone being well developed in the little Cycloturus.

The skull of the Anteater 2 is very long and low ; the fore-
part is tubular, and there appear to be no traces of teeth. The

FIG. 92. — Skull of Anteater (Myrmecopliaga). Lateral view, cd.sph, Alisphenoid ; coiul,
condyle of mandible ; cor, coronoid process of mandible ; ex.oc, exoccipital ;
ext.aud, external auditory meatus ; fr, frontal ; jit, jugal ; Icr, lachrymal ; max,
maxilla ; nas, nasal ; occ.cond, occipital condyle ; pal, palatine ; par, parietal ;
p.iiiax, premaxilla ; s.oc, supraoccipital ; sq, squamosal ; ty, tympanic. (From
Parker and Harwell's Zoology).

premaxilla is very small ; the zygomatic arch is imperfect, and
does not reach the squamosal behind. A curious feature of this
genus, which it shares with some Dolphins and other Whales, is
that the pterygoid bones develop palatine plates which meet each
other in the middle line, and thus shift the opening of the

1 See for anatomy Owen, Trans. Zool. Soc. iv. 1862, p. 117, and Forbes, Proc.
Zool. Soc. 1882, p. 287.

2 For the skull of Edentates generally see Parker, Phil. Trans, clxxvi. 1885,
pt. i. p. 121.

i68

OSTEOLOGY OF ANTEATERS

posterior nares backwards. This is also, of course, a character
of various lower vertebrates. Another Whale -like character in
the skull is the weak character of the mandible, which does
not give off a marked coronoid process. But then in neither
group is there much mastication. The tympanic, periotic and
squamosal are ankylosed together. A peculiarity of the cervical

b.oc

msf

FIG. 94. — Side view of three
mesosternal segments of a
young Anteater (Tainan-
duo), showing the mode of
articulation of the sternal
rib (sr). mst. The upper
or inner surface of the
mesosternal segment ; sy,
the synovial articulation
FIG. 93.-Skull of Anteater (Myrme- ^^ ^> segments.

cophaga). Ventral view. Letters (From Flower s Osteology,

as in Fig. 92. In addition, b.oc,
basioccipital ; glen, glenoid sur-
face for mandible ; pter, ptery-
goid. (From Parker and Has-
well's Zoology.)

vertebrae is that (as in the Camels) the vertebrarterial canal of
several of the vertebrae perforates the pedicle obliquely. There
are fifteen or sixteen dorsal and three or two lumbar vertebrae.
The additional zygapophyses upon the former have been already
referred to. The mode of articulation of the ribs is highly
singular.

Each segment of the sternum (of which there are eight) is
separated from the next by a synovial membrane : and it has on
either side two facets for articulation with the ribs. The way in

169

which these latter bones are connected with the sternum is
curiously like their mode of connexion with the spinal column at
their other end. With this may be possibly compared the double
articulation of the single rib (which articulates with the sternum)
in the Eorquals. In Cydoturus this mode of articulation does
not occur.

The manus of Myrmecophaga is five -fingered. Of these
the third digit (as in Perissodactyles) is the most prominent ;

FIG. 95. — A, Mauus of Great Anteater (Myrmecophaga jubata). x J. B, Manus of
Little Anteater (Gycloturus didactylus). x 2. c, Cuneiform ; I, lunar ; m, magnum ;
p, pisiform ; s, scaphoid ; td, trapezoid ; tm, trapezium ; u, unciform ; I- V, digits.
(From Flower's Osteology.)

it is at least double the width of the second or third finger ; the
pollex is very slender. In the little Cydoturus this is carried to
a greater extent : the third digit is relatively enormous ; the first
and the fourth have become quite rudimentary ; while the fifth is
only just recognisable as a minute ossification.

The chevron-bones in the tail surround a well-developed rete
mirabile, a rete being found in precisely the same position in
the Eastern Manis. Tamandua has also retia, which are also
found in the Spider-monkeys.

Cydoturus is by far the smallest of the Anteaters. It has

FOSSIL ANTEATERS

CHAP.

only two toes on the fore -feet. It is to be distinguished,
anatomically, from its larger relatives by the complete clavicle,
and by the fact that the pterygoids do not meet in the middle
line of the skull. The ribs, too, are unusually wide, as in the
Whale Neobalaena, and form a bony encasement for the body.
It has two small caeca. Of fossil Anteaters but little is known.
The most interesting form is Scotaeops, interesting because it has
two small back teeth, which are totally lost in its living allies.
The huge Patagonian extinct bird Phororhacos, first known by a
lower jaw, was at one time regarded as a member of this group on
account of the form and edentulous character of the jaw.

Fam. 2. Bradypodidae. — The Sloths, genera Brady pus and

FIG. 96. — Unau, or Two-toed Sloth. Choloepus didactylus. x I.
(After Vogt and Specht.)

Choloepus, come, as already stated, very near to the Anteaters, in
spite of their striking difference in appearance. The Sloths are
purely arboreal creatures, with strong recurved claws, which serve

vni PROTECTIVE COLOUR OF SLOTH 171

as hooks to keep them suspended from the lower side of a
branch. The three -toed sloth, Bradypus (or "Ai"), has the
exceptional number of nine cervical vertebrae ; the two-toed sloth,
Choloepus lioffmanni (or " Unau "), has the equally exceptional
number of six. The hair is long and shaggy, and gets an
adventitious green colour from the presence of minute algae.1 This
gives to the animal the appearance of a lichen-covered bough, a
resemblance which is increased in one species by an oval mark
upon the back, which suggests forcibly a broken end of such a
branch. The likeness of a Sloth to its surroundings is pointed

Lcr

FIG. 97. — Skull of Three-toed Sloth. Bradypus tridoctylus. Lateral view, fr, Frontal ;
ju, jugal ; Icr, lachrymal ; may, maxilla ; nas, nasal ; par, parietal ; s.oc, snpra-
occipital ; ty, tympanic. (From Parker and Haswell's Zoology.)

out by Dr. Siemann,2 who observed that a species occurring in
Nicaragua " has almost exactly the same greyish-green colour as
Tillandsia usneoides, the so-called ' Vegetable Horsehair ' common
in the district. ... If it could be shown that it frequented trees
covered with that plant . . . there would be a curious case of
mimicry between the sloth's hair and the Tillandsia, and a good
reason why so few of these Sloths are seen." The stomach in the
Sloths is complicated in structure, with several chambers ; one of
these gives off a long crescent-shaped caecum. The skull of the
Sloths agrees in a number of particulars with that of the Anteaters.

1 The colour fades in captivity owing to the disappearance of the algae.
- In a letter addressed to Dr. Gray, quoted by the latter in a revision of the
Sloths, Proc. Zool. Soc. 1871, p. 428.

OSTEOLOGY OF SLOTH

The zygoma is incomplete, though the part connected with the
frontal has a strong downward process like that found in .Diprotodon

and some other mammals. There is, moreover, a process from the
squamosal, though it does not reach the anterior part and thus

TEETH OF ARMADILLOS

complete the arcade. The premaxillaries are very small, and are
usually lost in dried skulls. Coupled with these points of likeness
are some differences. The lower jaw, for instance, has a well-
marked coronoid process. The pterygoids do not meet in the
middle line. The teeth are five or four in each half of each jaw.
There is no trace of a second set.

A peculiarity of the Sloths is the enormous number of dorsal
vertebrae. There are twenty -three of these in Clioloepus hoffmanni,
but only fifteen to seventeen in the Three-toed Sloth, Bradypus.
As in other American Edentates, the acromion joins the coracoid.
This connexion occurs in both the Two-toed and the Three-toed
species. The limbs of these creatures are very long, a concomitant
of an arboreal life. The femur has no third trochanter. The
genus Bradypus, which by reason of the fact that it has not lost
the third toe on the manus seems to be more primitive than
Clioloepus, shows another structural feature which does not bear
out this conclusion. The trapezoid and the os magnum of the carpus
are united, while in Clioloepus they are perfectly distinct bones.

The intestine has no caecum.

There are several species of Sloths. Eminently perfect though
the organisation of the Sloth in relation to its particular sur-
roundings appears to us, Buffon selected the animal as the very
type of imperfection in nature. " One more defect," he wrote,
" they could not have existed."

Fam. 3. Dasypodidae. — The family Dasypodidae or Arma-
dillos contains a considerable number of genera. Tatusia, Toly-
peutes, Dasypus, Xenurus, Priodon} and Clilamydopliorus. They
have all a more or less rigid covering of bony plates imbedded in
the skin, which are not in the least comparable with the scales of
the Manis. Save the Whales, in one or two genera of which
traces of a dermal armature exist, the Armadillos are unique
among existing mammals in this particular. The term " Edentate "
is especially inapplicable to the Armadillos ; the genus Priodon
may have more than forty teeth in each jaw ; a total of ninety
was found in one specimen examined by Professor Kiikenthal.
In the tendency of the teeth to multiply, we have another
example of a state of affairs which characterises so many Whales.
Generally, however, seven to nine is the number of teeth in each

1 This name is written " Prionodos " by Gray, which might lead to a confusion
with the Carnivore Prionodon.

174 SKULL OF ARMADILLOS CHAP.

half jaw, of which one is often implanted in the premaxilla.
The Armadillos show their alliance with the other American
Edentates in the points enumerated above. Their teeth specially
ally them to the Sloths, while the salivary and digestive organs
generally are on the Anteater plan, but present a less extreme
development. There are, however, caeca, paired as in birds, in
the genera Dasypus and Chlamydophorus. The others have none.
But there is a dilatation at the commencement of the large
intestine, which is not very different from the slightly-developed
caeca of Dasypus.

There are certain peculiarities in the skeleton, which dis-
tinguish this family.

The skull in the Armadillos presents a number of likenesses
to the other American Edentates.1 The premaxillaries are

*&

FIG. 99. — Skull of Armadillo. Dasypus seed itctus. x f . ex.oc, Exoccipital ; //•,
frontal ; max, maxilla ; 'IMS, nasal ; par, parietal ; peri, periotic ; p.max, pre-
maxilla : s.oc, supraoccipital ; sq, squamosal ; ty, tympanic. (From Parker and
Has well's Zoology.)

small, but are larger in Dasypus than in Tatusia. On the other
hand the lachrymals are larger in the latter. The zygomatic arch
is complete, but there is no downward process as in the Sloths.
In Tatusia (but not in Dasypus} the " short thick pterygoids add
somewhat to the hard palate." This is clearly a beginning or a
remnant of the quite crocodilian character of the palate of
Myraucophaga. In the cervical vertebrae we see the Whale-like
character of fusion between individual vertebrae ; and also, as
in the Whales, the degree to which this fusion is carried out

1 For the anatomy of several forms, see Garrod, Proc. Zool. Soc. 1878, p. 222,
who quotes other memoirs.

VERTEBRAE AND RIBS

175

varies ; two to four may be thus united. The additional
articular facets upon the dorsal vertebrae have been already
commented upon as a point of important likeness to other
American Edentates. The dorsal vertebrae are commonly eleven
in number, the lumbar being three. But in Priodon the numbers
are twelve and two respectively. There are traces to be observed
of the double-headed attachment of the ribs to the sternum. The

FIG. 100. — Bones of the right
maims of the Hairy Armadillo.
Dasypus villosus. x •§. c,
Cuneiform ; I, lunar ; m, mag-
num ; p, pisiform ; R, radius ;
s, scaphoid ; id, trapezoid ; tm,
trapezium ; «, unciform ; U,
ulna; I - V, digits. (From
Flower's Osteology.)

FIG. 101. — Bones of the manus of
the Great Armadillo. Priodon
giganteus. x £. a. An acces-
sory carpal ossicle in front of
the pisiform, which is not seen
in the figure. Other letters as
in Fig. 100. (From Flower's
Osteology. .)

shoulder girdle of the Armadillos is somewhat diverse in form in
different genera ; the acromion is always large, and is remark-
able in Priodon for the fact that the humerus also articulates
with it, its extremity being recurved, and forming a socket for
this purpose. As in some other Edentates there is a second
spine on the scapula behind the first. The clavicle is strong.
There is some variation in the form of the manus. It is five-
fingered in Dasypus ; in TolypeuUs the first digit has vanished ;
on the other hand, in Priodon, the fifth has* become rudimentary

76

PICHY-CIEGO AND APAR

ac

and the third enormously enlarged. This latter fact recalls
the arrangement characteristic of Myrmecopliaga. The pelvis is

greatly attached by the
ischiuin to the verte-
bral column. The
femur has a third tro-
chanter.

The various forms
of Armadillos are
largely distinguished
by the number of mov-
able thin bands of
scutes lying between
the large anterior and
posterior shields. Thus
we have Dasypus sex-
cinctus, Tolypeutes tri-
cinctus, etc.

The little Pichi-
ob&for , ,

•^ chago (or, more cor-

FIG. 102. — Pelvis and sacrum of Armadillo. Dasypus rec^y Pichv - cie^o)
sexcinctus. ac, Acetabulurn ; il, ilium ; isch, ischium ; * ' •

obtfor, obturator-foramen ; pect.t'itb, pectineal tuber- ChlamydopllOTUS,^i\\\C\\
cle ; pfA, pubis. (From Parker and Haswell's , t fe fc 5

Zvoloyy.) * &

inches in length, has

no movable bands at all. It is covered with a uniform series
of plates, which, moreover, are not discontinuous at the neck.
It differs, too, from the prevailing Armadillo- type by the absence
of conspicuous external ears. In the anterior part of the body
the armature consists of little more than the horny plates, which
in other Armadillos overlie the bony dermal plates. In the
hinder region the bony plates are strong. In this animal, there-
fore, we have the dermal armature reduced to a minimum ; but
it must be noticed that, like the extinct Glyptodons, the armn-
ture is continuous and nowhere ringed.

The genus Tolypeutes, of which the best-known species is T.
tricinctus, the Apar (there are two other species in the genus), can
roll itself up into a ball like the Pill-Millipede (Glomeris), and,
protected by its armour, roll away from its enemies like the
Arthropod under similar circumstances. This mode of protection,
be it observed, is also adopted by the Pangolin and by the Hedge-

PELUDO ARMADILLO

177

hog. The genus has only three movable bands. The tail is short,
and is covered with large tubercles. This genus is very markedly
digitigrade when running.

FIG. 103. — Three-baiided Armadillo or Apar. Tolypeutes tricinctus. x £.

The Peludo, Dasypus sexcinctus, is, like other Armadillos, an

FIG. 104.— Peludo Armadillo. Dasypus sexcinctus. x £. (After Vogt ami Speclit.)

omnivorous creature, and appears to be particularly fond of carrion.
It will burrow up to a decaying carcase like the ground-beetles.
VOL. x N

178 ARMADILLOS AND SNAKES CHAP.

Mr. W. H. Hudson has described the way in which this Armadillo
will kill a snake by holding it down and literally sawing the
reptile in half by help of the sharp and serrated edges of the
carapace. Dasypus has a very short tail, which is shielded by
distinct rings near the base.

Tatusia novemcincta is a species with nine movable bands.
The genus has four teats ; the ears are near together. There are
no caeca and no azygos lobe to the lung. A species apparently
belonging to this genus, but described under the generic names of
Cryptophractus and Praopus, is remarkable for the thick covering
of hair, not entirely wanting but usually thin in other Armadillos.
In this particular species the coat of hair is so thick as to
conceal the underlying plates of the carapace. The individual
hairs are stiff, and one inch and a half in length.1

The genus Xenurus contains several species, the best known of
which is inaptly named X. unicinctus. As a matter of fact the
characteristic feature of the genus is the existence of twelve
or thirteen movable plates between the two ends of the body.
X. unicinctus has twelve dorsal and three lumbar vertebrae. This
Armadillo, known by the vernacular name of the Cabassou, has one
of the most modified hands that are found in the family. The first
two digits are slender and elongated ; but are quite normal in the.
number of their phalanges. In the remaining three digits the
metacarpal is short and broad, while the proximal phalanx is
either suppressed altogether or fused with the metacarpal, the
middle phalanx is present but short, while the third phalanx is very
large indeed. As in Dasypus, but not as in Tatusia, which is in
so many other respects divergent from these genera, the lungs
have an azygos lobe. As a small point of difference, tending to
show an alliance between the genera Xenurus and Dasypus
and their difference from Tatusia, is the deeply-imbedded gall-
bladder ; this sac is not nearly so deeply plunged into the hepatic
tissue in Tatusia. Xenurus has no caecal dilatations. The
brain " is intermediate in its form and surface markings between
Dasyjms and Tulypeutes." The small intestine is nearly eighteen
times the length of the large. But these intestinal measurements
are not of much avail in this group as marks of affinity, since in
three species of Dasypus Garrod gives the following widely-
divergent lengths: — D. villosus, 11-5 feet and 1'25 ; D. mimttus

1 Flower, Proc. Zool. Soc. 1886, p. 419.

GIANT ARMADILLO 179

5 • 1 , with a large intestine of no less than 7 feet ; D. vellerosus
4-3 and -66.

Priodon is the giant of its race. This Armadillo may
reach a length of 3 feet to the base of the tail. The tail is
some 20 inches long. The large number of teeth has been already
noticed. There are twelve or thirteen bands. Other points
in the structure of this genus have already been mentioned, and
need not be recapitulated. This Armadillo feeds upon termites
and carrion.

Scleropleura is unfortunately but imperfectly known. The
single species, named by Milne-Edwards 1 S. fyruneti, is apparently
a very rare inhabitant of Brazil. It is known by a single skin,
which was tanned by the hunter who obtained it. Thus the
hair, if any, has dropped out. The plates in the skin are
deficient along the back and even upon the top of the head, and
are barely represented upon the tail posteriorly. The ears are
small and distant from each other. The tail is longish, about
one-third of the length of the body. The total length of the
creature including the tail is rather more than a foot and a half.
The hunter who obtained it regarded it as a hybrid between an
Armadillo and an Anteater.

Extinct Xenarthra. — There are a good many extinct forms
of Armadillo, apart of course from the Glyptodons. Feltephilus
is referred to later (p. 186). Dasypus was represented by a large
form, 6 feet long, with a skull of one foot in length. The genus
Eutatus was also large. The carapace was formed of thirty-three
distinct bands, of which the last twelve are soldered together, but
not fused into a shield as in Dasypus, etc.

An extinct group of American Edentates, termed the GRAVI-
GRADA,2 are somewhat intermediate between the Sloths and the
Anteaters. A number of the genera are well known from com-
plete skeletons.

One of the typical forms of this group is Mylodon, which,
together with its immediate allies, is often placed in a separate
family, Mylodontidae.

Mylodon itself was a large creature, as big as a Rhinoceros.
It was covered externally by armour in the skin, which did not
form a massive armature as in the Glyptodonts, but was in the

1 Milne-Edwards, Nouv. Arch. Mas. vii. 1871, p. 177.
2 See especially Lydckker, An. Mus. La Plata, Pal. Art/, iii. 1894.

i8o

EXTINCT XENARTHKA

CHAP.

form of scattered plates, small and not fused together. The
general aspect of the skull is decidedly Sloth-like. As in that
animal, the malar bone is bifid posteriorly, and between the
bifurcation is embraced the process of the squamosal. This latter

is thus more de-
veloped than in the
Sloth, but there is
no actual union be-
tween it and the
malar. The pre-
maxilla is small.
The lower jaw has
both coronoid and
ascending processes,
and is massive.
There are five teeth
on each side above,
and four on each
side below, as in the
Sloths. There are
the normal seven

FIG. 105. — Mylodon robiistus. (Restoration, after Owen.

cervical vertebrae and sixteen dorsals. The limbs are not long and
slender, but short and strong, the animal having been terrestrial. The
fore-feet were five-toed, of which the three inner toes had claws. The
hind-feet were only four-toed, and the two inner only were clawed.

Scelidotherium is a genus which is a trifle smaller than the
last. It has only four properly-developed toes in the fore-foot,
the thumb being rudimentary ; of these, the first two bear claws.
The hind-feet are also four-toed. Like Mylodon, Scelidotherium
is a Pleistocene genus.

Glossotherium has a skull very much like the last two
genera ; but it is remarkable for the fact that the nostrils instead
of being unprotected with bone anteriorly are there closed by a
plate of bone formed by the well -developed premaxillae, the
nostrils appearing at the sides, and giving the skull a curious
likeness to that of a Chelonian. From a series of recent and
most important observations it appears to be clear that this genus
has survived into quite modern times.1

1 Dr. Moreno and Mr. A. Smith Woodward in Proc. ZooL Soc. 1899, p. 144 ;
Wiss. Ergeb. Schwed. Exped. Magellansldnd. ii. 1899, p. 149.

vin NEOMYLODON OR GLOSSOTHER1UM l8l

The well-known naturalist of La Plata. Senor Moreno, engaged

3 * O O

in studies connected with the political boundary line between
Chili and the Argentine, had occasion to visit Consuelo Cove
on Last Hope Inlet in Patagonia. Hanging from a tree he
noticed a piece of dried skin, which at once struck him as looking
more like the remains of a Mylodon than of any living animal.
The inhabitants regarded this piece of skin as a great curiosity,
but were of opinion that it was the hide of a cow encrusted with
peebles ! This fragment from a bygone age was originally
described by Professor Ameghino, who had apparently seen some of
the bonelets imbedded in it, as Neomylodon listai, " a living
representative of the ancient Gravigrade Edentates of Argentina."
That this piece of skin is of quite recent date seems to be proved
by a number of considerations. In the first place it is covered
by long hair of a light yellowish-brown colour ; it does not seem
likely that hair would preserve its character for geological epochs.
The nearest corresponding case is that of the remains of Moas in
New Zealand, whose feathers, dried skin, and tendons are known.
Now the Moa was unquestionably contemporaneous with man, as
abundant surviving legends prove, and indeed it cannot have been
long extinct. Still, hair is a resisting structure, and in a dry cave,
with no possibility of irruptions of floods, might retain its characters
for long periods. The evidence, however, of more recent date is
stronger than this. The skin shows patches of reddish colour, sugges-
tive of course of blood-stains. A small piece of the outside of the
skin at the cut edge, which presented the appearance of freshly
or comparatively freshly dried fluid, was submitted to a chemical
examination and shown to be serum ! Dr. Lb'nnberg examined
chemically a bit of the skin itself and found in it, after boiling,
glue, " which proves that the collagen and gelatinous substances
are perfectly preserved." After this it seems impossible to suppose
that the skin can be of any very great age ; for bacteria would
have finished their work upon the serum and gelatine long ago.
Combined with the fresh appearance of the skin is the very
fresh appearance of the skull. In fact it is impossible to believe
that the animal was not alive quite a few years since, relatively
speaking. It is admitted that this animal was contemporaneous
with man. There are actually legends of a creature which may
have been this Glossotherium. "Ancient chroniclers inform us
that the indigenous inhabitants recorded the existence of a

I 82 HAIR OF NEOMYLODON

strange, huge, ugly monster, which had its abode in the Cordillera
to the south of latitude 37. The Tehuelches and the Gennakens
have mentioned similar animals to me, of whose existence their
ancestors had transmitted the remembrance; and in the neigh-
bourhood of Eio Negro, the aged Cacique Sinchel, in 1875,
pointed out to me a cave, the supposed lair of one of these
monsters, called ' Ellengassen ' ; but I must add that none of the
many Indians with whom I have conversed in Patagonia have
ever referred to the actual existence of animals to which we can
attribute the skin in question."

A rude painting in a cavern, in red ochre, seems to Dr.
Moreno (whose words we have just quoted) to be somewhat
suggestive of a Glyptodon. There are some reasons for believing
that this quadruped was kept by man as a domestic creature.
In the cave are two walls of rough pieces of stone which seem
to have dropped down owing to the wearing away of the roof;
they also seem to have been loosely piled together to form two
walls, within which enclosure an imperfect skull of the animal was
found. This skull shows clearly that the so-called " Neomylodon "
must be referred to Glossotherium or Grypotherium, as it is
sometimes termed. This skull is perforated on the roof in such
a way as could only have been effected (in the opinion of
experts) by a weapon in the hand of a man. A hole in the skin
has been even compared to a bullet-wound. But this it is per-
haps unnecessary to discuss. The skin of Glossotherium is, like
that of other extinct " Ground-sloths " (e.g. Mylodon), filled with
small and irregular ossicles. But in Mylodon, the sculptured
appearance of the dermal ossicles appears to indicate that they
reached the surface of the body and were covered by epidermis
alone, which is not the case with the animal now under con-
sideration. The microscopic characters of the ossicles, too, show
differences in the two. Glossotherium being " precisely inter-
mediate between Mylodon and the existing Armadillo (Dasypus}."
Now Glossotherium and Mylodon are regarded as forms which lie
between the existing Anteaters and the Sloths of the same part
of the world. We have already pointed out the facts of structure
which lead to this conclusion. It might therefore be reasonably
surmised that the hair of Glossotherium would be also inter-
mediate, or at least like that of one of the two genera Myrme-
cophaga and Bradypus. But microscopical investigation has

ME GA THERIUM I 8 3

negatived this supposition. It has shown that the Armadillos
are in this matter the nearest relatives of Glossotherium. This
result is important as tending further to confirm the close inter-
relationship of all the American Edentates as contrasted with the
Old- World forms — a matter which has already been emphasised.
It is suggested, however, that the absence of under fur, which is
so well developed in the Sloth, and the difference shown in trans-
verse sections from the hair of Myrmecophaga, may be explained
by difference in habitat. Glossotherium lived under conditions
similar to those under which the Armadillos live to-day. Thus
the outer covering of the body became alike in the two cases, the
same needs supervening in both genera.

Lestodon is another allied genus, which seems to possess
canines. At any rate, in front of the four molars, and separated
from them by a diastema, is a smallish, somewhat canine-like
tooth, in both jaws.

Megalonyx and its allies are sometimes placed in a distinct
family, Megalonychidae. Megalonyx itself had a skull very
like that of Bradypus, being shorter and not so elongated as
in the Mylodontidae. There is a strong tusk anteriorly, which
is separated by a considerable space from the three molars lying
behind it. Both pairs of limbs seem to have possessed five toes.
This is a North American genus. It differs from the bulk of
the American Edentates in having a complete jugal arch.

Megatherium is the type of yet a third family, Megatheriidae,
of the Gravigrade Edentates. This creature is familiar from
the many restorations which have been built up, and from its
huge bulk, little short of that of an elephant. The skull, which
is small for the size of the creature, has a complete jugal
arch, from the middle of which depends a downward process as
in other allied forms. The teeth grow to an extraordinary
depth, and there are five of them in the upper and four in the
lower jaw — on each side of course. The fore-limbs of the
Megatherium are very much more slender than the enormously
bulky hind-limbs, upon which and the equally massive tail the
animal seems to have supported itself while tearing down
branches of trees, upon whose leaves it fed. In the scapula
the acromion joins the coracoid as in Bradypus ; the clavicle is
large. The fore-limb is four-toed, and the hind-limb three-toed.
The latter has but one clawed digit (the third, i.e. the inner).

1 84

ARMOURED EDENTATES

On the manus, the three inner digits have powerful claws. This
animal, too, was Pleistocene in time. The Megatheriidae had,
however, small as well as gigantic forms.

The genus Zamicrus had a skull no bigger than that of a
Sloth, while Nothrotherium was also a comparatively small
creature ; the teeth of the latter genus are reduced to £.

The extinct group of the Glyptodontidae comprises large
creatures with a dense covering of bony scutes which are arranged
in a tesselated fashion, and thus form an immobile armature of
immense strength. In correspondence with this massive carapace
the dorsal vertebrae have fused together, and the lumbar vertebrae
form a series ankylosed to each other and to the following sacrals.
These creatures are all South American.

Glyptodon, the genus which gives its name to the family, is
known from numerous remains in South America, and also from

t -

FIG. 106. — Glyptodon davipes. x jV. (After Owen.)

so far north as Texas and Mexico. It grew to be as long as 16
or 17 feet. In the skull there is an exceedingly long
downward process of the zygomatic arch, as in Sloths, the arch
itself being complete. The process extends so far down as to
reach a point about on a level with the middle of the lower jaw.
The nasals are short or rudimentary. As in Myrmecophaga, the
pterygoids enter into the formation of the bony palate. The
lower jaw has a spout-shaped extremity, and, behind, it rises into
an enormous vertical branch as high as the front part of the jaw
is long. There are eight teeth in each half of each jaw. As in

viit PANOCHTHUS AND GLYPTODON 185

some Armadillos, the cervical vertebrae are at least partly fused.
The atlas is free, but the rest, or at any rate five of them, are
united. The last cervical is sometimes fused with the succeeding
dorsals ; the latter are twelve in number, and are fused together

' ' O

so far as concerns their centra and neural processes. The
succeeding region of the vertebral column includes seven to nine
lumbars, which are fused with the eight sacrals ; in this region the
neural processes are high, and there is thus produced a strong and
lofty ridge along the back, which forms a powerful support for
the carapace. The fore-limbs are shorter than the hind-limbs,
which latter are attached to an unusally massive pelvis. The
claws of the limbs are blunt and almost hoof-like.

The heavy carapace consists of sculptured, five or six-sided
plates, which have no particular arrangement in the middle, but
towards the margins show indications of an arrangement in trans-
verse rows. The moderately long tail is also encircled by bony
skin-plates which are thorny above, or at least provided each with
a blunt upstanding process. It appears that outside this bony
system of scutes were horny epidermic scales, corresponding
exactly with the tesserae which they cover. There are apparently
a good many species of Glyptodon.

In the allied genus Panochthus the tail is rather longer, and
the bony rings which surround it, instead of being all movable
as in Glyptodon, are at first so, but later, i.e. towards the end of
the tail, become welded into a single and massive piece. Both
feet are here four-toed, while in Glyptodon the hind-feet are five-
toed and the fore-feet four-toed.

Daedicurus shows a further specialisation, in that the feet have
three and four digits respectively. The orbit too shows a
specialisation in being separated from the temporal fossa. The
descending process of the zygomatic arch is not so extraordinarily
exaggerated as it is in Glyptodon. It has the same terminal
tube of osseous scutes upon the tail. This creature seems to have
reached a length of about twelve feet.

Propalaeokoplophorus is, unlike the great Armadillos that we
have hitherto dealt with, a small animal, not exceeding 2 feet
or so in length of carapace. A small alveolus on each side of the
premaxillae seems to suggest the former presence of an incisor
tooth ; and it seems that the animal possesses both true molars
and premolars; for the first four of the eight teeth are much

I 8 6 PEL TEPHIL US

simpler in structure than those which follow. The dorsal
vertebrae again are not fused together ; the hind-limbs are five-
toed. All the plates of the carapace are arranged in definite
transverse rows ; it has been observed, too, that some of the
anterior scutes overlap like those of the Armadillos, to which this
animal possesses further likenesses in the exclusion of the
maxillae from the border of the nostril (a Glyptodont character),
and the comparative feebleness of the scutes.

A primitive genus also appears to be Peltephilus, which is
perhaps rather an Armadillo than a Glyptodon. However, it
comes somewhat between the two, like Propalaeohoplophorus, with
which it may therefore be treated. A most singular feature
of this genus has been mentioned on p. 27 in connexion with
the skull in the Mammalia generally. That is the fact that
a portion of the squamosal surrounding the articular facet for
the lower jaw is separated by a suture from the rest of that
bone, and is therefore obviously suggestive of the quadrate in
the lower Vertebrates. As in certain Armadillos and Glyptodons,
etc., the pterygoids appear in this genus to have taken, a share in
the formation of the hard palate. The plates of the carapace were
movable, as is shown by the fact that they sometimes slightly
overlap. In view of the possible origin of the Edentates from
lowly-organised Mammalia, it is noteworthy that the humerus
has been especially compared to that of the Monotreme. Pelte-
philus differs from other Armadillos in having teeth in the front
of the jaws. The total number of teeth is twenty-eight, i.e. seven
in each half of each jaw.

SUB-ORDER 2. NOMARTHEA.

As already explained, the Old -World Edentates differ from
the New-World forms in having normal dorsal vertebrae, that is
to say, without additional zygapophyses. That negative feature,
however, though combined with the positive fact that both the
Old- World forms feed upon ants, is hardly sufficient to outweigh
the many structural differences which distinguish the Oryctero-
podidae from the Manidae ; which will be placed therefore in
different groups. To that containing the Aard Vark, the name
TUBULIDENTATA may be applied.

AARD VARK i 87

This group contains but one family, the Orycteropodidae, of
which there is but a single genus.

The Aard Yark (earth-pig), genus Orycteropus, is characterised
by its heavy build, the body being covered by rather coarse and
not very abundant hair ; the snout is long and pig-like, with round
nostrils at its end ; the ears are long, erect, and pointed ; the
tail is very thick at first, so that it has been aptly described as
" a tapering of the body to a point." The fore-limbs are four-toed,
the hind five-toed.

FIG. 107. — Aard Vark, or Cape Anteater. Oryrferopus capensis. xTV

In the skull there is a complete though slender zygoma ; the
premaxillaries, though small, are not so rudimentary as in the
American Edentates. The annular tympanic is not ankylosed to
the surrounding bones, a character found in other low mammals.
Contrary to what is found in Manis, Orycteropus has a huge
lachrymal. There are thirteen dorsal and seven lumbar vertebrae.
The clavicle is well developed. Orycteropus is peculiar among
Edentates in that the ischia do not unite with the vertebral
column. The femur has a third trochanter.

As mentioned on p. 1 02, the Aard Yark is diphyodont like normal
mammals. The permanent teeth consist of five molars and pre-
molars on each side of each jaw ; the first two of these are pre-
molars, and are simpler in their form than the succeeding two
teeth, which are partly divided by a median furrow into two
halves. These teeth are also peculiar in that they consist entirely
of vaso-dentine. They have been compared in minute structure
to those of the Eay Myliolates. According to Mr. Oldfield

TEETH OF AARD VARK

Thomas1 there are seven milk teeth on each side of the upper
jaw (limited to the maxillae, and thus not incisors). An eighth
tooth was discovered on one side of one of the specimens examined
by Thomas. In the lower jaw there are only four milk teeth on
each side. It is interesting to note that the histological structure

FIG. 108.— Section of lower jaw with the teeth of Orycteropus. x 2. (After Owen.)

of these milk teeth agrees with that of the permanent teeth.
There are two species of this genus found in Africa : the southern,
0. capensis, is more hairy than the northern, 0. aethiopicus.
0. gaudryi is a Pliocene species from the Island of Samos and
from Persia, described by Dr. Forsyth Major and Dr. Andrews.2
It closely resembles the existing 0. aethiopicus.

Of the Scaly Anteaters, Group SQUAMATA or Manidae, there
is really but one genus, though Phatagin, Pholidotus, Smutsia,
and Pangolin have been used to distinguish various forms. The
genus Manis is African and Oriental in range. Dr. Jentink, who
has lately revised the species, allows seven.3 The external form
of these animals is fairly well known, the remarkable scales dis-
tinguishing the Pangolins from other animals. Between the
scales lie hairs, which seem to be absent in the adults of the
African species, though present in the young, thus affording a
convenient method of distinguishing the Ethiopian from the
Oriental forms. The scales have been compared to agglutinated
hairs. That they are not " merely mimetic of the Lizards' scales "
is held by Weber,4 who compares them directly with those struc-

1 Proc. Roy. Soc. xlvii. 1890, p. 246.

2 Proc. Zool. Soc. 1893, p. 239, and 1896, p. 296.

" Revision of the Manidae in the Leyden Museum," Notes Leyd. Mus. iv. 1882,
p. 193.

4 Weber, Zool. Ergebnisse einer Seise in Niederl. Ost Indien, 1892. See also
Romer, in Jen. Zeitschr. xxxi. 1896, p. 604, and Reh, ibid. xxx. 1895, p. 137.

THE PANGOLINS 189

tures, as he does the scales of other mammals, such as those upon
the tail of Anomalurus, etc. This, however, is not a universal
opinion. It is true that these scales occur chiefly in the lower
forms of mammals such as those under consideration, Marsupials,
Rodents, and Insectivores ; but the fact that the hairs are developed
before the scales shows, or seems to show, that the former are the
older structures, and to lead to the inference that the scales of
mammals are new structures. The scattered hairs of the Pangolin
have no sebaceous glands excepting on the snout. This, again,
looks as if they were degenerate structures, and emphasises the non-
archaic character of the scales. These animals have no trace of
teeth except possibly some slight epithelial thickenings which have
been interpreted as a last remnant ; the tongue is suited for the
capture of ants, and is therefore much like that of the not nearly-
related American Anteaters. The stomach is of simple form ;
it is characterised by a large gland, which suggests that of the
Koala (see p. 144); the intestine has no caecum. Eetia mirabilia
occur on the limb arteries. The placenta is non-deciduate and
diffuse ; it is specially compared by Weber with that of the Horse.
Considering the many adaptive resemblances between this genus
and the American Anteaters, especially in the mouth cavity, it is
remarkable that in Manis the pterygoids are not joined as they
are in Myrmecophaga. In spite of statements to the contrary,
it appears that there is sometimes a distinct lachrymal.

A remarkable feature in the skeleton of Manis is the
singular sternum. The xiphoid cartilage is extraordinarily
elongated into thin strips, which reach the pelvis and return.
This state of affairs is to be found in the African species only.
This structure is not comparable, as it has been said to be, with
abdominal ribs such as those of the reptile Hatteria.

These animals are mainly anteaters. The Japanese have a
curious legend as to the method adopted for the capture of ants,
which is related by Dr. Jentink in his monograph of the genus.
The Manis " erects his scales and feigns to be dead ; the ants
creep between the erected scales, after which the anteater again
closes its scales and enters the water ; he now again erects the
scales, the ants are set floating, and are then swallowed by the
anteaters " ! The same story is related by Mr. Stanley Flower
on the authority of the Malays.

Though it seems clear that the likenesses which Manis shows

190

ANCIENT EDENTATES

to the Anteaters of the New World are chiefly adaptive and have
nothing to do with real affinity, being merely an expression of a
similar mode of life, it is curious to note that here and there we
do find certain resemblances which do not seem to be susceptible
of the latter explanation. The jugal bone, absent in Manis, is
small in Myrmecophaga ; the clavicle is absent and again small or
rudimentary in the Anteaters; it is large in other Edentates.

Fio. 109. — Manis. Manis gig antea.

The third trochanter is absent, as in Myrmecophaga (and the
Sloths). There are many scales on the body ; in Myrmecophaga
there are traces of these structures on the tail, as also in
Tamandua. In the features mentioned, the Myrmecophagidae
differ from either or from both of the two other American
families (i.e. Dasypodidae, Bradypodidae) and agree with Manis.
The facts are not a little remarkable.

Order III. GANODONTA.1

Allied to the Edentata, and apparently representing the
ancestral forms from which they, at any rate the Xenarthra
were derived, is the order of the Ganodonta. Of this order a
number of genera are now known, which can be ranged in a
series which more and more approaches the Edentata as we pass
from the older to the newer forms. This interesting and transi-
tional series will be made manifest by a description of the
characters of the various genera taken in their proper chrono-

1 See Wortman, "The Ganodonta and their Relationship to the Edentata,"
Bull. Am. Mus. Nat. Hist. ix. 1897, p. 59.

vni PS1TTACOTHER1UM 19 1

logical order. The following genera are included by Wortinan in
his family Stylinodontidae.

The earliest type of the Ganodonta is the genus Hemiganus,
with but one species, H. otariidens. This animal lived during
the deposition of the lowest Eocene strata, the Puerco beds of
North America. It was about as big as a fair-sized Dog, and had
powerful jaws. There were at least two pairs of incisors in the
upper jaw, together with powerful canines and the full premolar
and molar formula. In the lower jaw the canines were also
strong, but the incisors are not certainly known to be more than
two pairs. The enamel upon the posterior surface of the canine
'is thin, and in the case of the incisors the enamel seems to be
limited to the anterior face. The lower molars are quadrituber-
cular. It is believed from the presence of a suture on the upper
surface of the premaxillary that the snout of the creature was
tubular. The cervical vertebrae, only known by their centra, are
like those of the Armadillos (and for the matter of that of the
Whales) in the great transverse as opposed to the antero-posterior
diameter. The feet are especially compared with those of the
Ground Sloths. The single ungual phalanx is marked by a large
subungual process, which is pierced by a considerable foramen.
The tibia again is to be compared with that of the Armadillos.

In the Upper Puerco (Torrejon) beds the remains of Psitta-
cotherium are found. This genus, when first discovered, was
referred to the Tillodontia by some and to the Ungulates, the
latter being a refuge for indeterminate Eocene mammals, just as
the " Multituberculata " is for similarly-placed Secondary mammals.
It is now known to be clearly a member of the order Ganodonta.
Wortman thinks that there is but one species, P. multifragum.
It seems to have had a general aspect much like that of Hemi-
f/anus — that is judging from the skull — and was not very greatly
different in size. The facial portion of the skull is short, and the
zygoma is deep. The infra-orbital canal is double, a feature which
crops up in the Sloth, and has been mentioned in the later form of
Ground Sloth, Megalonyx (but it must be remembered that the
same characteristic is not unknown in Kodents). The dentition
is reduced as compared with that of Hemiganus, that is to say, as
far as concerns the molars and the incisors. There is but a single
pair of incisors in each jaw ; the canines are strong ; the premolar
and molar series seem to have been complete in the lower jaw,

I 92 GANODONTA — CALAMODON CHAP.

but reduced by one premolar at least in the upper jaw. It is very
important to notice that the incisors have enamel only on their
anterior faces, and that the same is the case with the canines,
the slender layer present behind the tooth in Hemigamts having
vanished in this later form. The tooth pattern of the molars is
like that of Hemiganvs. The fore-limb is decidedly Edentate-
like ; but it is the foot which presents the strongest likenesses to
that order. " If an anatomist," remarks Dr. Wortman, " had no
other part of the skeleton than that of the foot to guide his judg-
ment, and he should fail to detect a most striking similarity
between it and that of the Edentata, especially the Ground Sloths,
he would not only lay himself open to the criticism of being
lacking in the ordinary powers of observation and comparison, but
would be suspected of placing the matter upon a basis other than
that established by such a method." It is not certain how many
toes upon the fore-limbs were possessed by Psittacotherium, but
the close resemblance to Mylodon is indeed striking, the third
digit being in both forms the most pronounced. Some vertebrae
of this Ganodont have been discovered which do not show the
complex articular arrangements of later American Edentates. The
sacrum, on the other hand, is very like that of the Sloth, and there
is a foreshadowing of the attachment of the ilia to the sacrum by
co-ossification which is met with in later Edentates. A still later
type is the genus Calamodon, which has been shown to occur in
Europe as well as in America. C. simplex was a larger beast
than either of the genera that have already been treated of, thus
affording another example of the increase in size of later as com-
pared with earlier members of the same group, so pronounced
among the Ungulata. The lower jaw has the same massive
structure that characterises that bone in Hemiganus and Psitta-
cotherium. There is but one incisor, but the premolar and molar
series are complete. The canine is Kodent-like in appearance,
being imbedded throughout the greater part of the lower jaw ; it
evidently grew from a persistent pulp. It is enamelled upon the
anterior face only. The premolar and molar teeth are in this
genus commencing to lose their enamel, which is distributed in the
form of vertical bands, leaving interspaces which are not covered
by enamel. These teeth, moreover, are decidedly hypselodont,
more decidedly so than in Psittacotherium ; they are, when unworn,
quadricuspidate, with accessory cusps ; when more worn, the teeth

STYLINODON 193

are double-ridged, and that transversely to the long axis of the
jaw; finally, the much-worn teeth have flattish crowns more or
less surrounded by a ring of enamel.

A still later form, coming from the Lower and Middle Eocene
strata, is the genus Stylinodon. S. cylindrifer, which is the more
archaic of the two described species, is only known from a single
molar, fragments of a canine, and " some inconsiderable pieces of
the skull." The molar is interesting on account of the fact that
the enamel is still further reduced ; it is represented only by
narrow vertical strips, which are much narrower than those of
older forms of Ganodonts. It is also hypselodont, and has a
persistent pulp. So, too, the canine which had a thick anterior
facing of enamel. The later species, S. mirus, is more fully
known. The teeth seem to have been much the same as in the
last-described species ; the premolars and molars were seven in all
in the lower jaw, and the canine was imbedded in the bone for a
long distance, as in Calamodon. The cervical vertebrae have
short centra as in Hemiganus. The clavicles were well developed.
The humerus possessed an entepicondylar foramen, and its head
displays the pyriform pattern so characteristic of later Edentates.
The foot is clearly like that of Psittacoiheriwn.

In reviewing the series, therefore, we see a gradual diminution
of the incisors, a gradual loss of enamel on the teeth generally,
and the production of hypselodont teeth growing from per-
sistent pulps ; all of which are features of the later Edentates.
The progression is so gradual that the forms enumerated and
described seem to have been part of a continuous series cul-
minating in the Ground Sloths of later times. The other points
of similarity will be gathered from the facts given in the fore-
going pages.

There is another family belonging to the Ganodonta whose
position with regard to the Edentata is not so clear. This is
the family Conoryctidae, of which two genera are known. The
earliest of these, from the Lower Puerco, is Onychodectes. In
0. tissonensis the skull is long and narrow, thus contrasting with
that of the last family. The facial part is also long. The lower
jaw is much more slender. The molar formula was complete,
but there is some doubt as to the incisors. The molars are
tritubercular.

The other known genus is Conoryctes. Its skull has a shorter

VOL. x o

194 GANODONTA — ONYCHODECTES CHAP, vni

facial portion, and is thus more like that of Stylinodontidae
than that of Onychodectes. The dental formula is known, and is
complete save for the loss of one incisor above and below, and
one premolar above. The relationship of these Ganodonts to any
later forms is uncertain ; but their skeletal structure is as yet by
no means fully known.

CHAPTER IX

UNGULATA CONDYLARTHKA AMBLYPODA ANCYLOPODA

TYPOTHERIA TOXODONTIA PROBOSCIDEA HYRACOIDEA

Order IV. UNGULATA

THE existing members of this order can be readily grouped
into the Hyracoidea, Proboscidea, Perissodactyla, and Artiodactyla,
each of which divisions has quite the value of an order, and all of
which are sharply marked off from each other. But as the dis-
covery of so many fossil forms has to a great extent rendered
these demarcations less sharp, it is better to regard all these
groups as not more than sub-orders of a larger " Order " Ungulata.
Even when this conclusion has been necessarily arrived at from
a consideration of the more ancient groups of Ungulate animals,
the definition of such an order remains a difficult matter for
the systematist. For the earliest of these forms, more particu-
larly the Ancylopoda, the Amblypoda, and the Condylarthra,
whose peculiarities will be dealt with at length subsequently,
are not by any means easily differentiated from "the primi-
tive Carnivorous mammals of that date, the Creodonta ; these
latter, moreover, fade into the Marsupials through the so-
called Sparassodonta of Professor Ameghino. To confine our-
selves to the Ungulates, we may perhaps define them as terres-
trial animals with hoofs rather than claws or nails, and chiefly,
if not entirely, vegetarian in habit. The teeth are bunodont or
lophodont, the tendency to the production of the latter type being
always marked. The walk, although plantigrade in the older types,
becomes more and more digitigrade, except in such survivals from
antiquity as Hyrax. There is, too, as we pass from the ancient
types to the modern, a gradual perfection of the limbs as running

196

PHENACODUS

and not climbing or grasping organs ; the number of toes be-
comes reduced, and culminates twice (in the horse and in the

Litopterna) in one toe
on each foot ; at the
same time the ulna be-
comes rudimentary and
fuses with the radius,
and the fibula in the
hind -limb undergoes a
like reduction. The
clavicle is absent even
in some of the oldest
types ; its presence in
^ Typotherium l is highly
I remarkable. The tail
•2 too, an organ which is
« long in some of the
| early forms, gets short
| in their modern deriva-
^ tives.

» Coupled with the in-
•3 creasing perfection of the
£ foot as an organ used
%? merely for the support
* of the body, certain in-
1 teresting changes have
<^ taken place in the
"^ arrangement with re-
& gard to each other of
the several bonelets of
the wrist and ankle. It
has been held by Cope
and others that the truly
primitive disposition of
these bones was that pre-
sented to us by certain

early types, such as Meniscotherium or the existing elephant or
Hyrax. In these animals there is (see Fig. 112) a serial arrange-

1 This creature is, however, sometimes referred to the neighbourhood of the
Rodents.

BONES OF WRIST AND ANKLE

197

ment of these bones, the distal bones only, or very nearly only,
articulating with the corresponding bones in the upper series. In
the modern types (cf. Fig. 113) there is, on the other hand, an
interlocking, so that the bones of the distal series articulate with

H

H

H

FIG. 111. — Series of metacarpals and nietatarsals of Camelidae, to show secular and
progressive increase in size. From left to right the species are Protylopus
petersoni, Poebrotherium labiatum, Gomphotherium sternbergi, Procamdus occi-
dentalis. F, Fore-foot ; H, hind-foot ; III, IV, third and fourth nietapodials.
(After Wortmau.)

two of those of the proximal series. By this is produced, as it
would appear, a much firmer foot, less liable to " give " under
pressure, and thus more fitted for an animal that runs. It is the
same principle as that adopted in the laying of bricks. The actual
stress and strain of impact has been held responsible for those
changes. An equally ingenious and possibly truer explanation of
the undoubted facts has lately been advanced by Mr. W. D.

198

SERIAL AND INTERLOCKING CARPUS

Matthew.1 He has pointed out that in some ancient Ungulates
the carpus is not serial but interlocking, even in forms which
belong to the earliest Eocene groups,' such as the genus Protolambda
among the Amblypoda. Now in the fore-foot of Meniscotherium
and the living Hyrax there is a separate centrale which is wanting
in the greater number of Ungulates. The absorption, that is the
practical dropping out of this bone, would restore to an interlocking
carpus the serial arrangement ; while on the other hand, by the

B

IV

IV

FIG. 112. — Bones of the maun* A, of the Indian Elephant, Elephas indicus. x J.
B, of the Cape Hyrax, Hyrax capensis. x 1. c, Cuneiform ; cc, centrale ; I,
lunar ; in, magnum ; p, pisiform ; R, radius ; td, trapezoid ; tin, trapezium ; s,
scaphoid ; u, unciform ; U, ulna. (From Flower's Osteology.)

fusion of this bone with the scaphoid, the interlocking disposition
would be maintained.

The gradual perfecting of the fore- and hind-limbs as running
organs has been put down to the advent of the grasses, and the
formation of large plains covered with this herbage. The same
reason would also be in harmony with the equally gradual change
in the shape of the molar teeth, from a tubercular form calculated
for a mixed or even a carnivorous diet, to the natter crushing sur-
faces exhibited by the lophodont teeth of later Ungulates. Strong

1 Bull. Amer. Mus. Nat. Hist. ix. 1897, p. 321.

ORIGIN OF PERISSODACTYLES

199

canines would in the same way cease to be useful, and even
become encumbrances to such grazing creatures ; and their dis-
appearance is one of the salient features in the history of the
Ungulata, that is of the modern representatives of the order. The
extraordinary hypertrophy of these teeth in such a line as that of
the Amblypoda, which has left no descendants, was one of the
reasons perhaps for the decay of those great pachyderms of mid-
Tertiary times ; their excessive armature became an encumbrance,
since it was not accompanied by improvements in other necessary

FIG. 113. — Bones of the manus A, of Rhinoceros, Rhinoceros sumatrensis. x £. B, of
Pig, Sus scrofa. x *s. Letters as in Fig. 112. (From Flower's Osteology.)

directions. Some of the features of the Tertiary Ungulates have,
however, been dealt with in our general sketch of the mammalian
life during that epoch, and need not be again referred to here. Of
existing Ungulates there are no clear indications of the descent of
the Elephants or of the Hyracoidea. Their structure proclaims
these two divisions to be of ancient descent, and not to be modern
twigs of the Ungulate stem. As to the Perissodactyla and the
Artiodactyla we cannot bring them together nearer than in quite
early Tertiary times. The order Condylarthra seems to be the
starting-point of both these sub-divisions. Euprotogonia has been
considered to be an ancestor of the Perissodactyle branch, and
Protogonodon or Protoselene of the Artiodactyla. If this be true,

V£ * >^ -

I*

2OO

*

»r'
Jt

the likenesses which Titanotherium shows to the Artiodactyla
must be either purely superficial and secondary, or a cropping
out of ancient characters which had been dormant for many
generations.

Horns. — The Ungulata are the only order of mammals which
possess horns ; as they are on the whole a more defenceless group
than the Carnivora, it may be that the horns are a counterpoise
to the teeth and claws of the latter ; need for defence and for
armature in the combats with their own kind for the favours
of the does has led to a different kind of protective and
aggressive mechanism. Horns as weapons are, however, parti-
cularly effective in this group wherever they exist. A Ruminant
is most frequently a large and heavy animal without the agility
and litheness of the Carnivore. It is precisely to this sort of
animal, where weight is an important consideration, that horns are
the most suitable weapons. This is further shown by the fact
that although the general term horn is used to describe the
weapons of the Ungulate mammals, there is more than one kind
of structure included under this general term ; it is indeed prob-
able that the extreme terms in the series of horns have been
independently acquired by their possessors. There is but little
in common between the horns of a Giraffe and of a Rhinoceros.
In the Rhinoceros we have one or two horns, in the latter
case one placed behind the other, which are purely epidermic
growths ; they may indeed be regarded as matted masses of hair,
borne, it is true, upon a boss of bone, which however is not
a separate structure. The Giraffe supplies us with the simplest
term in that series of horns which are partly epidermal and partly
bony. The paired horns of this animal have often been contrasted
with those of the Deer, for example ; but there is no fundamental
difference between them. In the Giraffe a pair of bony out-
growths, originally separate from the skull which bears them, but
ultimately ankylosed to it, are covered by a layer of entirely un-
modified skin. A distinction of undoubtedly practical importance
is usually drawn between the Hollow-horned Ruminants, i.e. Oxen,
Goats and Antelopes, and the Deer tribe. There is nevertheless
no fundamental distinction. In the Antelopes there is a core of
bone, the " os cornu " as it has been termed, which is covered
by a horny layer, the horn proper, variously modified in shape and
size according to the genus or species. In the Deer there is the

HORNS AS A SEXUAL CHARACTER 2OI

same os cornu, which may however be branched, but which is in
the same way covered by a layer of modified integument ; this is
known as the " velvet " ; it only lasts for a certain period, and is
then torn off by the exertions of the animal itself, leaving behind
the bony core, which is popularly termed the horn. It will be clear
that here is only a difference of comparative unimportance ; the
same essential features are present in both groups of animals, but
the modification of the epidermis has progressed along different
lines. Both can be referred back to the primitive conditions seen
in the paired horns of the Giraffe. Even the difference, such as it
is, is bridged over by the Antelope Antilocapra, where the os cornu
is bifid and the horn is periodically shed, as is the velvet of the
stag ; but in the stag the bony part of the horn is also shed, a
state of affairs which has no parallel in the Hollow-horned
Ruminants. The great Sivatherium may conceivably be an
annectant form between the two types of compound horns, i.e.
those of the Antelope and those of the Deer. This creature had
two pairs of horns, of which, naturally, only the bony cores remain ;
the hinder pair of these were branched. But although so far they
resemble the Deer's horns rather than the Antelope's, Dr. Murie
has thought that they were covered by a horny sheath and not by
soft skin as in the Deer. In any case these horns were apparently
never shed, which is a point of likeness with the Antelope and of
difference from the Deer. Apart therefore from the nature of the
covering of the bony cores, there are good grounds for looking
upon them as intermediate between those of the Deer and those
of the Antelopes.

The horns of the Ruminants are frequently a secondary sexual
character ; this is especially the case with the Deer. The Rein-
deer is, however, an exception, both the stags and the does
having horns. That they are associated with the reproductive
function is shown by their being shed after the period of rut,
the destruction of the velvet at that period, and also by the effect
upon the horns which any injury to the reproductive glands
produces. Some useful facts upon this latter head have been
amassed by Dr. G. H. Fowler,1 who noticed in a series of stags,
horns showing various degrees of degeneration in the antlers pro--
duced by varying degrees and periods of gelding. From the facts

1 ' ' Notes on some Specimens of Antlers of the Fallow Deer, etc. , " Proc. Zool. Soc.
1894, p. 485.

202 CONDYLARTHRA

here collected it is clear that a direct effect is produced. If we
are to regard horns as secondary sexual appendages which have
been subsequently handed on to the female by heredity, we should
expect to meet with examples of animals now horned in both
sexes, of which the earlier representatives had the horns confined
to one sex. This is most interestingly shown by the extinct and
Miocene Giraffe, Samotherium, of which the male alone had a pair
of short horns, while the skull of the female was entirely hornless ;
the modern Giraffa, as is well known, has horns in both sexes.

It is interesting to note that the existing Perissodactyles and
Artiodactyles are to be distinguished by their unpaired or paired
horns. But while there are.no Artiodactyles with unpaired horns
(save occasional sports) the Perissodactyles have more than once
tried, so to speak, paired horns, which ultimately proved fatal
to them. The Rhinoceros Diceratherium apparently inherited and
improved upon the small paired horns of Aceratherium, but it has
left no descendant. The paired horned Titanotheria offer another
instance of the same apparent incompatibility between the Perisso-
dactyle structure and the persistence of paired horns.

SUB-ORDER 1. CONDYLARTHPvA.

This group is characterised by the following assemblage of
characters. Extinct, often plantigrade Ungulates, with five-toed
limbs. Bones of carpus and tarsus not always interlocking, but
sometimes lying above each other in corresponding positions.
The humerus has an entepicondylar foramen. Dental formula
quite complete ; the molars brachyodont and bunodont. The
premolars are simpler than the molars. The canines are small.
As with other early types, the zygapophyses are flat and do not
interlock. The astragalus is like that of the Creodonta. This
group was American and European in range, the remains of its
rather numerous genera being of Eocene time. The best-known
genus is Phenacodus, of which some account will be given before
discussing the, in many cases, more fragmentary remains of
other allied forms.

The genus Phenacodus was first described so long ago as 1872,
from a few scattered teeth. Since then several nearly complete
skeletons have been obtained, and we are in full possession of

ix PHENACODUS AND THE CREODONTS 2O3

the details of its osteology. It was not a large creature (see
Fig. 110, p. 196), about 6 feet in length, with a small head.
The feet were more or less plantigrade, and five-toed. The last
phalanges of the toes show that they carried hoofs and not
claws ; yet the fore-feet look a little as if they could be used as
grasping organs. The third digit of both hind- and fore-feet
exceeds the others, and thus a Perissodactyle-like foot characterised
this Eocene creature. The tail is exceedingly long, and must
have reached the ground as the animal walked. This is of course
by no means an Ungulate character. Still, in the totality of its
organisation the animal was decidedly Ungulate, though Professor
Cope spoke of Phenacodus as not merely an ancestral Ungulate
but as the parent form of Insectivores, Carnivores, Lemurs, Monkeys,
and Man himself ! The scapula indeed is from its breadth and oval
contour rather like that of a Carnivore. The clavicles as in other
Ungulates are absent. The femur is Perissodactyle rather than
Artiodactyle in the presence of a third trochanter. The creature
had fifteen pairs of ribs and five or six lumbar vertebrae. The
two bones of the leg which lie below the femur are perfectly
distinct and separate. A cast of the brain-case shows that the
cerebral hemispheres were smooth and small, the cerebellum of
course completely uncovered and nearly as large as the cerebrum.
The olfactory lobes were also large. The complete skeleton of
Phenacodus has lately been excavated more fully from the
enveloping matrix by Professor Osborn,1 and mounted in what
is regarded as the natural position of the beast. It appears
that though five-toed it went upon the three middle toes only,
and furthermore that of these the middle one was the more
prevailing, so that Phenacodus was distinctly " Perissodactyle,"
at least in habit. Moreover its " long hind-quarters, the long
powerful tail . . . are reminiscent of Creodont ancestry." The
genus was European and American in range.

Meniscotherium ( = Hyracops ~} comprises several forms of
about the size of a fox ; they are both European and American in
range. The teeth are more distinctly Ungulate in form than those
of Phenacodus, with a W-shaped outer wall. The skull is
described as possessing " indifferent, primitive characters," permit-
ting a comparison with those of Opossums, Insectivores, and

1 Bull. Amer. Mus. Nat. Hist. x. 1898, p. 159.
- Marsh, Amer. Journ. Sci. xliii. 1892, p. 447.

204 CONDYLARTHRA — PROTOGOKIA CHAP.

Creodonta. It has, as in Phenacodus, no orbital ring. The
humerus resembles that of a Carnivore rather than that of an
Ungulate. The carpus and tarsus are serial. The fibula articu-
lates with both the calcaneum and the astragalus, which is not
the case with Phenacodus. It is suggested that these animals
are ancestral forms of the Chalicotheres. In the brain the
hemispheres do not cover the cerebellum.

More primitive apparently than Phenacodus was the less-known
genus Euprotogonia, or Protogonia l as it has been called. The best-
known species is E. puercensis, so called from its occurrence in the
Puerco beds of the American Eocene. It was a slender, long-
limbed creature, smaller than Phenacodus, with a long and heavy
tail as in that animal. Like Phenacodus it was semiplantigrade,
and shows more likenesses to the Creodonta. The skull is only
known by a part of the lower jaw with teeth, and by the teeth of the
upper jaw. The vertebrae are not entirely preserved, but enough
remain to show that the animal had a tail of 1 6 or 1 7 inches, which is
a considerable length when compared to its height, about a foot at
the rump. In the fore-limb the most noteworthy point is that the
ulna has a convex posterior border as in the Creodonts, the same
border in Phenacodus being concave. The humerus is slender, with
less-marked tuberosities. The fifth digit seems to have been less
reduced. The phalanges seem to have borne horny sheaths some-
what intermediate between hoofs and claws. The pelvis is
described as being, as is also that of Phenacodus, rather like that of
the Creodonta. The right hind-limb is known in all its details.
It appears that the bones are not serial but interlocking ; this,
however, on the views with regard to the relations of these two
forms of tarsus mentioned on p. 198, does not militate against
regarding Euprotogonia as the ancestor of the genus Phenacodus.
The third toe is the pre-eminent one, the animal thus being
Perissodactyle. The lateral digits are larger than in Phenacodus,
and the metatarsals and the phalanges are slightly curved, which
is again a Creodont character as compared to the perfectly straight
corresponding bones of Phenacodus. It seems evident that this
animal is to be looked upon as a more ancient type than Phena-
codus, even if not as its actual ancestor.

Another group of the Condylarthra contains the genus
Pertipychus and some others. Periptychus has the full dentition

1 See W. D. Matthew, Bull. Amer. Mus. Nut. Hist. ix. 1897, p. 303.

ix TEETH OF EARLY UNGULATES 2O$

of forty-four teeth, the molars being of course bunodont, with the
three chief tubercles most developed. The bones of the tarsus
interlock and are not serial, as they are in many other members
of the Condylarthra. The astragalus has a shorter neck than in
Meniscotheriuin, for example. It has in this a likeness to the
same bone in the Amblypoda, to the primitive members of
which, such as Pantolambda, this animal bears much resemblance.
" Astragali and many skeletal bones of Periptychus rhabdodon and
Pantolambda bathmodon are almost indistinguishable," observes
Mr. Matthew. The fore-feet of this genus are unknown, but it
would seem that it was plantigrade from the evidence of the hind-
feet. There are several species of the genus.

Possibly, but not at all certainly, the Mioclaenidae, with the
genera Mioclaenus and Protoselene, are to be referred to this same
order of primitive Ungulates. It is only necessary to mention
them here, because they show very clearly the primitive form . of
dentition of these early Eocene mammals. The teeth are quite
complete and unbroken by a diastema. The canines are but little
pronounced. The molars are not strictly tritubercular, but have a
prevailing trituberculy. The nature of the feet is not known.
Since the genus Protoselene, as its name denotes, shows an indica-
tion of a commencing selenodonty, it has been suggested that this
group is the stock whence the Artiodactyles have been derived.

In any case, whether the particular comparisons that have been
made as to the relationship of various forms of Condylarthra are
valid or not, it seems to be plain that this group represents the
earliest Ungulate stock, but little differentiated from the con-
temporaneous Creodonts.

SUB-ORDER 2. AMBLYPODA.

This group of extinct mammals has the following principal
characteristics : —

They are large, semiplantigrade Ungulates, of heavy build and
apparently elephantine gait. The dentition is for the most part
complete as in other ancient groups, and the canines are in the
later forms big tusks. The back teeth are brachyodont and
ridged (lophodont). Both radius and ulna in the fore-limb, and
tibia and fibula in the hind-limb, are well developed. The bones

2O6

AMBLYPODA — DINOCERAS AND ITS ALLIES

in the carpus are alternating in position. The toes are five in
both feet, and are very short. There is a hint of commencing
" perissodactylism " in the fore -feet at any rate. The brain is
small and the hemispheres smooth.

The Amblypoda, or Amblydactyla, are so called on account of
their short and stumpy feet and toes. They were held by Pro-
fessor Cope to be on the direct line of ancestry of both Perisso-
dactyles and Artiodactyles, a view which is on the whole not
accepted at present.

As is the case with other groups, the Amblypoda commenced
existence as a sub -order with relatively small forms such as

FIG. 114. — Skull of Protolambda bathmodon. x f . e.a.m, External auditory meatus ;
m, mastoid ; m.f, mastoid foramen. (After Osboru.)

Pantolambda, the most ancient type known, which is in many
respects a transition between the later forms and other groups of
mammals such as the Creodonta.1 The race culminated and
ended in the giant Dinoceras and Coryphodon, and spread into the
Old World. In spite of their smooth and diminutive brain, these
mammals were able to hold their own and to multiply into many
species and genera ; in this they were perhaps aided by their
formidable tusks and by the horns which many of them possessed.
The teeth seem to imply an omnivorous diet, which was quite
possibly an additional advantage in the struggle for existence.
It does not seem to be necessary to divide off the Dinoceratidae
into a sub-order equivalent to the Coryphodontidae as was done

1 Or perhaps rather to the primitive Ungulates Condylarthra. It is especially
compared with Periptychus of that group.

CORYPHODON 2O?

by Professor Marsh ; the numerous points in common possessed
by the members of both families forbid their separation more
widely than as families.

The earliest types of Amblypoda belong to the genus
Pantolambda, of which the species P. bathmodon was about four
feet in length. As restored it seems to have had proportionately
short fore- and hind-limbs, and it had a long tail. It was
apparently plantigrade, and would have had not a little likeness
to a carnivorous type. The skull has no air cavities, such as are
developed in the later types from the Lower Eocene, e.g. Cory-
phodon ; Pantolambda is from the basal Eocene. The frontal
bones show no trace of the horns that are developed in subsequent
forms ; the nasals are comparatively long ; the zygomatic arch is
slender. The molar teeth are in the primitive form of trituberculy,
and the premolars, as is so often the case with primitive animals,
are unlike the molars in form, being less markedly selenodont.
As to the vertebral column, the dorsal vertebrae appear to have
had short spines, which argues, as it does also in the case of the
larger and heavier Corypliodon, a feebleness in the development of
ligaments and muscles supporting and moving the head. The
scapula seems to have the same peculiar leaf-like form that it has
in the later Coryphodon.1 This primitive type shows an entepi-
condylar foramen in the humerus. It is interesting to observe
that the posterior border of the ulna is convex, as in the Creodonts,
and in the early Condylarthrous form Euprotogonia. In the sub-
sequently-developed Amblypoda, as in the later Condylarthra,
that bone acquires a concave outer border. In the carpus the os
centrale is distinct. In the femur the third trochanter is well
formed ; it gradually dies out in later Amblypoda. The fibula
articulates with the calcaneum. This species, according to Osborn,
" typifies the hypothetical Protungulate, being more primitive
than either Euprotogonia or Phenacodus." *

The genus Coryphodon is known by a large number of species,
of which the first was discovered in this country, and was repre-
sented merely by a jaw with some teeth. This was named by Sir
E. Owen C. eocaenus, and was dredged up from the bottom of the
sea off the Essex coast. A second specimen consisted of a single

1 The scapula of P. bathmodon is unknown.

- For the structure of this genus and of Coryphodon, see Osborn, Bull. Am.tr.
Mus. Nat. Hist. x. 1898, p. 169.

COR YPHODON 2 OQ

canine tooth only, and was brought up from a depth of 160 feet
during the making of a well at Camberwell. More abundant
remains have since been found in North America.

This genus had a large head, and in some specimens traces of
the " horn cores," so marked in the related Dinoceras, are to be
noticed. The skull is broad behind and narrowed in front ; the
roofing bones show the cellular spaces so characteristic of the
Elephant. The jugal bone, however, is not, as it is in the
Elephant, placed in the middle of the somewhat massive zygomatic
arch. As in some other primitive Ungulates (e.g. Phenacodus)
there are twenty dorso-lumbar vertebrae, of which fifteen bore ribs.

The scapula seems to have possessed a peculiar leaf-like form,
swelling in the middle and ending almost in a point above. It
has a well-marked spine, and the acromion projects much. The
fore- as well as the hind-feet are in a state of transition between
plantigradism and digitigradism. It was at one time held that
the animal was digitigrade as to the fore-feet and plantigrade as to
the hind-feet. Though, as has been pointed out, it is a fact that
the hind-feet are often on a different plane of evolution from the
fore-feet, it seems that this amount of difference does not characterise
any Ungulate, not excepting the genus now under consideration.

The toes are very spreading. The pelvic girdle is of great
strength and broadness. The femur, as in the Perissodactyles,
has a well -developed third trochanter; but whereas in this
particular the hind-limb is Perissodactyle, it is Artiodactyle in
the fact that the tibia and the fibula articulate with the astragalus
and calcaneum. The ridged teeth have given the name to the genus.

A curious feature in the structure of the genus are the
slender spines of the dorsal vertebrae, which contrast with the
enormous ones of some other Ungulates — more curious in this
genus, which is of heavy build, than in the lighter Pantolamlda.
The back of the animal is short, and the limbs are very spreading,
so that the gait was doubtless shuffling. The large head, and
short and heavy limbs and limb girdles added probably to its
cumbrous walk or trot. The canines are great tusks, and spread
out on both sides of the mouth.1

The late Professor Cope, in 1874, described the probable appear-
ance of the Corypliodon in the following words : — " The general
appearance of the Coryphodons, as determined by the skeleton

1 Osborn, Bull. Amer. Mus. Nat. Hist. x. 1898, p. 81.
VOL. X P

2 I O DINOCERAS

probably resembled the bears more than any living animals, with
the important exception that in their feet they were much like the
elephant. To the general proportions of the bears must be added
the tail of medium length. Whether they were covered with bail-
or not is of course uncertain. Of their nearest living allies, the
elephants, some were hairy and others naked. . . . The movements
of the Coryphodons doubtless resembled those of the elephant in its
shuffling and ambling gait, and may have been even more awkward
from the inflexibility of the ankle."

The most recent members of this sub-order come from the
Middle Eocene beds, and are chiefly referable to the genus Dino-
ceras. with which Tinoceras and Uintatherium are at least very
nearly related, if not identical. These creatures were of great size,
larger than the earlier types which have been considered. They
show a certain superficial resemblance to the Titanotheriidae, on
account of the massive horn cores upon the skull. These horn
cores are large upon the maxillae and the parietals, and are
paired ; on the nasals are smaller horns. The bones of the
skull have air cavities. The incisors of the upper jaw are
absent; the canines are enormous tusks, and the lower jaws
are flanged downwards near the symphysis where these tusks
border them. Contrary to what is found in the older types,
where the position of the condyle of the lower jaw is normal,
this prominence faces backwards in the Dinocerata. The same
shortness of the spines of the dorsal vertebrae prevails in this
group as in the other Amblypoda, though it is perhaps hardly
so marked. The scapula has not the peculiar acuminate form
that exists in Coryphodon, but is triangular and broad above.
The limbs are elephantine, in that the angle between the
humerus and the femur respectively, and the bones which
follow, is not marked. The hind-limbs are especially straight.
The tail is short as compared with that of the primitive
Amblypoda. The Dinocerata are purely digitigrade. The
entepicondylar foramen has, as in the Coryphodonts, disappeared.
The os centrale of the carpus has become fused, and no longer
exists as a separate bone. The fibula no longer articulates
with the calcaneum, but both that bone and the ulna are well
developed. The genus Astrapotherium is placed among the
Amblypoda by some authorities.1

1 Gadow, A Classification of Vcrtebrata, Recent and Extinct, London, 1898.

THE " PANGOLIN GIGANTESQUE " 211

SUB-ORDER 3. ANCYLOPODA.

The history of the discovery of the members of this order
is very instructive as illustrating the dangers of laying too much
classificatory importance upon detached fragments of animals. So
long ago as 1825 terminal phalanges of a new creature were found
in the Miocene of Eppelsheim, and sent to Cuvier. Cuvier named
them " Pangolin gigantesque," deeming them, on account of their
general form and cleft terminations, to pertain to the Edentata.
In the same bed some seven years later were found certain teeth
clearly of an Ungulate character, to which the generic name of
Chalicotherium was applied. It was subsequently discovered
that the teeth and the claws belonged to the same animal, and,
later, further remains turned up which disclosed a creature
having the anomalous composition of an Ungulate with decisively
Ungulate teeth, but with the feet to a large extent like those of
an unguiculate animal. The same confusion of characters occurs
also, it will be remembered, in the distinctly Artiodactyle
Agriochoerus (see p. 331). Indeed the feet of the latter when
first discovered were erroneously, as it now appears, referred to
the present order of Ungulates under the name of Artionyx.
It is probable that the genus Moropus of Xorth America is a
member of this group, and that it is probably congeneric with a
somewhat different type of Ancylopod known as Macrotherium.
It is also clear that Anisodon, Schizotherium, and Ancylotherium,
if not congeneric with either of the two recognised genera, are
at least very close to them.

Chalicotherium has a skull which recalls that of some of the
earlier Ungulates ; it has, however, no incisors at all, and no
canines in the upper jaw ; this feature has led to the belief that
the animal is related to the Edentata, and that it is in fact a link
between them and the Ungulata. The molars, like those of the
Perissodactyla, are of the buno-selenodont type. It also agrees
with that group (to which it has been approximated by several
writers) in the tridactyl manus and pes, and in the characters
of the tarsus. But although tridactyl, the axis of the limb
passes through the fourth digit. Chalicotherium is not mes-
axonic, as are the Perissodactyles. Moreover, it has no third

212 COPE ON THE TYPOTHERIA CHAP.

trochanter, and the unguiculate claws have already been referred
to. As to the latter, which are short, it is not the end phalanx
but the first which is retracted ; thus Chalicotherium differs
markedly from both Carnivorous and Edentate types ; for in the
former it is the last phalanx which is retracted, while in the
Edentates the same phalanx is flexed downwards. The limbs of
Chalicotherium are nearly of the same size, and the animal
seems to have been stout and quadrupedal.1

Macrotherium, like the last genus, seems to have been
common to both New and Old Worlds. It is to be distinguished
by a number of characters. It is supposed to have been " semi-
arboreal and fossorial " ; the fore-limbs are much longer than
the hind, the relative proportions of the radius and tibia being
70 to 29. The ulna was distinct from the radius, whereas in
Chalicotherium the two are coalesced, or nearly so. Young
specimens appear to possess a full set of incisors ; whether this
is the case or not with Chalicotherium is not known.2

Homalodontotherium is sometimes placed in the group.

SUB-ORDER 4. TYPOTHEEIA.

It is a little difficult to be confident that the Typotheria are
rightly referred to the Ungulata, since they contradict two im-
portant Ungulate rules. They have clavicles, which are elsewhere
missing, and the thumb looks as if it were opposable.3 An
Ungulate is essentially a running animal, and has no need of a
grasping finger. Still Typotheria are placed by most within the
Ungulate series, though their undoubted likenesses to other
groups, especially to the Eodentia, are admitted, and indeed
emphasised. Cope places them definitely with the Toxodonts.

The Typotheria are an extinct group of smallish beasts,
confined, like the Toxodontia, to South America, a region which

O

during the Tertiary period, and into the Pleistocene, abounded
with strange and varied types of Ungulate animals.

The earlier forms of Typotheria may be exemplified by some

1 See Osborn, American Naturalist, February 1893, p. 118.

2 It is not absolutely clear whether both or only one genus ranged into
America. Different opinions have been expressed.

3 It must be remembered, however, that there is a suggestion of a prehensile
character in the hand of Phenacodus (see p. 203).

ix DENTITION OF TYPOTHERIA 213

account of the genus Protypotherimn. This animal was of
about the size of a Hyrax, which indeed it resembles in several
points of structure. The teeth have the primitive number of
forty-four, and they are close set, leaving no diastema ; the
molars are rootless and grow persistently ; they are simple and
Eodent-like in surface pattern. The shape of the lower jaw is
like that of Hyrax, being rounded in outline posteriorly ; there
is no projecting angle as in the Rodents, and this remark
applies to the Typotheria in general. The aspect of the Eodent
lower jaw is characteristically different from that of Hyrax and
the forms under consideration.

Some other characters of these early forms of Typotheria can
be gathered from an inspection of other genera. In Icocliilus
both hand and foot were five-toed, and, as in ancient Ungulates
generally, the bones of the wrist and of the ankle are serially
and not alternately arranged. Moreover, an os centrale is
present in the carpus. Both thumb and big toe were opposable.
The skull has a remarkably Eodent-like appearance, but the
palate is not so narrowed as in these animals.

In the more recent forms of Typotheria the dentition has
become reduced. The canines are lost, and as the incisors are
reduced also, to one on each side of the upper, and two on each
side of the lower jaw, the likeness to a Eodent skull is increased.
There is also evidence of a modification from the more primitive
forms in the loss of one premolar or even more, in the alternating
bones of the carpus, in the disappearance of the centrale, and in
the loss of a toe upon the hind -foot. In these more recent
forms the fibula articulates with the astragalus instead of with
the calcaiieum. Typotheria of these more recent forms may be
illustrated by the typical genus Typotherium. It has the re-
duced dental formula I ^ C % Pm f M f ; the molars are simple
in pattern, and much like those of Toxodon. The upper incisors
are powerful and curved, but are surrounded by a layer of enamel,
which is not limited to the anterior face, as it practically is in
Eodents. The sacrum is composed of a large number of verte-
brae— some seven — a state of affairs which recalls the Edentata.
The shoulder blade is not Ungulate in form. It has a strong
spine, with an acrornion and a well-developed metacromion. The
terminal phalanges are enlarged and hoof-like.

In the genus Pachyrucos there are three premolars, otherwise

214 "GIANTS' BONES" IN PATAGONIA CHAP.

the formula is the same as in Typotlierium. The animal seems
to have had nails rather than hoofs. The thumb was opposable.
The fibula is fused below with the tibia, whereas in the last
genus these two bones are quite separate from each other.

SUB-ORDER 5. TOXODONTIA

The group Toxodontia,1 like so many others, is exceedingly
hard to define. " Nor are its limits any easier to mark out than
many others of the groups of Ungulates. It will be best perhaps
to give an account of Toxodon, and of a few types which seem to
lie near it in the system, and then to indicate how far they
resemble or depart in structure from other Ungulates. Toxodon
itself is known from complete skeletons. It lived in Argentina
during the " Pampean " period, which seems to be of the Pleistocene
age. A large number of species, however, have been described, some
of which seem to go farther back in time, and to have existed
during the Miocene period further south in Patagonia.

The size of this creature was about that of a large Ehinoceros ;
it has a bulky body and a large head, which was borne low
down, on account of the bending downwards of the anterior
vertebrae ; in this aspect the figures of the skeletons recall
Glyptodon and similar Edentates. The beast was discovered by
Darwin, and originally described by Owen. " During his (Mr.
Darwin's) sojourn in Banda Oriental," writes the Eev. H.
Hutchinson, " having heard of some ' giants' bones ' at a farm-
house on the Sarandis, a small stream entering the Kio Negro,
he rode there, and purchased for the sum of eighteenpence the
skull which has been described by Sir E. Owen. The people at
the farm-house told Mr. Darwin that the remains were exposed
by a flood having washed down part of a bank of earth. When
found, the head was quite perfect, but the boys knocked the teeth
out with stones, and then set up the head as a mark to throw
at." The whole of the Pampean area is a valley of dry bones,
and the remains of Toxodon are abundant there. The skull of
Toxodon is not unlike that of a horse in general aspect ; but the
orbit is not separated from the temporal fossa. The premaxillae
are furnished above with a slight protuberance directed towards
1 Cope, American Naturalist, xxxi. 1897, p. 485.

ix TOXODON AND NESODON 2 I 5

the free end of the nasals, which may be related to the presence
of a short proboscis. The zygomatic arch is strong and broad ;
the mandibles are provided with a long symphysis. The dental
formula is I -| C ^-1 Pm s *:^ M |. The teeth are prismatic and
hypselodont, growing from .persistent pulps. The molar teeth are
slightly arched in form, whence the name of Toxodon, " bow teeth."
The strong chisel-shaped incisors suggest the Eodents and Hyrax.
The cheek teeth, moreover, are by no means unlike those of Eodents
in their pattern. They are at any rate not at all .like those of ex-
isting Ungulates. The small size of the canine and of the first pre-
inolar produces a diastema in the tooth series. The sacrum consists
of five vertebrae, and the ischium does not articulate with it.

The shoulder blade has a strong spine, but only a rudimentary
acromion ; nor is the coracoid well developed. The radius crosses
the ulna, as in the Elephant ; the whole fore-limb is shorter than
the hind-limb, which must have exaggerated the hang-dog ex-
pression of the creature when alive. The elements of the carpus
interlock in the modern fashion. Those of the tarsus, however,
are primitive in lying below each other without alternation. The
carpus has a centrale. The fibula articulates with the calcaneum.
The femur has no third trochanter. There are three toes to all
the limbs. It is clear that this assemblage of characters will not
allow the placing of Toxodon in any living Ungulate order. If
the middle toes appear by their slight pre-eminence to approach
the Perissodactyle form, the peculiar surface contour of the molar
teeth, letting alone the absence of a third trochanter on the femur,
will not permit this classification.

Allied to Toxodon is the genus Nesodon. It was so named
from an " island lobe " on the inner side of the upper molars.
This creature, smaller than Toxodon, also differs from it in the
fact that the dentition is complete, and in the pattern of the
molars, which is rather more complex. There is still the slight
projection upon the premaxillary bones, but the nostril is directed
more forwards than in Toxodon. The zygoma, too, is massive.
The second pair of incisors in the upper jaw and the outer (third)
pair in the lower jaw form biggish tusks in the adult. These and
the molar teeth are, however, finally rooted, and do not grow, as in
Toxodon, from persistent pulps. The genus is from the older
Tertiary of Patagonia. Five or six species have been described.
Some are as large as a Ehinoceros, others as small as a Sheep.

•
2l6 THE AFFINITIES OF TOXODONS CHAP.

There is no doubt about the close alliance of the two genera
just referred to. It is more doubtful whether Jfomalodonto-
therium and its allies should be placed, as they often are, in the
neighbourhood of the Toxodonts. Homalodontotlierium owes its

O

name to its even row of teeth without a diastema. It was a
creature of equally large size with Toxodon, and also carne from
the Tertiary strata of Patagonia. The teeth are the typical forty-
four, and the molars like those of a Ehinoceros ; they are, how-
ever, brachyodont and not hypselodout as in Toxodon. This genus,
however, shows an important difference from the Ehinocerotidae
and from the other Toxodontia in the fact that it was five-toed, and
that the bones of the carpus and tarsus are set in relation to each
other in the linear serial fashion.

Undoubtedly a near relative of Homalodontotlierium is Astrapo-
therium. This creature was of equal bulk, and was also Patagonian
in range. The teeth are reduced in number, but the animal was
provided, like a Wild Boar, with great tusks, which were, however,
formed by the incisors. This animal is very imperfectly known ;
it is the form of the molars and the large size of the incisors which
have led to its association with the Toxodontia. As to the resem-
blance of the teeth of this genus and of Homalodontotherium to
those of Rhinoceros, it is difficult to regard it as evidence of near
affinity. The likeness is probably to be looked upon as a case of
parallelism in development. Exactly the same explanation is
possibly to be given to the likeness which the teeth of Toxodon
and Nesodon show to Eodents, or even to Edentates. As to their
affinities Zittel observes : —

"The entirety of their osteological characters argues for the
Toxodon a separate position in the neighbourhood of the Perisso-
dactyla, Proboscidea, Typotheria, and Hyracoidea. The relations
to the Eodentia rest mainly upon the converging development of
the teeth, not upon true relationship."

SUB-ORDER 6. PEOBOSCIDEA.

I^arge vegetable-feeding animals, usually scantily covered with
hair, and with the nostrils and upper lip drawn out into a long
proboscis. Digits five on both limbs. Femur and humerus not
bent upon lower leg and fore-arm in a position of rest. Skull

ix CHARACTERS OF ELEPHANTIDAE 217

with abundant air cavities in the roofing and other bones. The
incisors are developed into long tusks, which exist in the upper
jaw alone, in the lower jaw alone, or in both jaws. There are
no canines. The molars are lophodont. The clavicle is absent.
The femur has no third trochanter. The bones of the carpus are
serially arranged and do not interlock. The stomach is simple.
The brain has much convoluted cerebral hemispheres, but the
cerebellum is completely uncovered by them. The intestine is
provided with a wide caecum. The testes are abdominal. The
teats are pectoral in position. The placenta is non-deciduate and
zonary. There are two venae cavae superiores.

The position of the limbs in the Elephant tribe is unique among
living animals : their straightness that is to say, and the absence
or very slight development of angulation at the joints of the
limb bones. This same feature has been observed in the extinct
Dinocerata and in the Titanotheria. It must not, however, be
assumed from the resemblance to these ancient forms that there is
much affinity between them and the Proboscidea, or that the latter
have retained an ancient feature of organisation. The oldest
Ungulates for the most part, and the Creodonts to which they are
undoubtedly related, have much bent limbs. It must be considered,
therefore, that the arrangement obtaining in the Elephants is purely
secondary. Professor Osborn has put forward the reasonable view l
that the vertical limbs of all these colossal creatures are due to
"an adaptation designed to transmit the increasing weight" of
these animals. The huge bulk of the body is better borne by
vertical pillars than by an angulated limb. Other points, however,
such as the exposure of the cerebellum, the two venae cavae, the
five digits, and the absence of a third trochanter, argue a low
position for the Proboscidea in the Eutherian group.

The group can be readily divided into two families, the
Elephantidae and the Dinotheriidae. We will commence with
the former.

The Elephants proper, Elephantidae, differ from the Dino-
theriidae in, and are characterised by, a number of anatomical
features. They possess long tusks (incisors) either in both jaws,
or, if only in one jaw, in the upper. The molar teeth are very
large — so large that only a few of them are simultaneously in use.
There are but three definable genera of Elephantidae, of which

1 American Nat. February 1900, p. 89.

2 I 8 SKULL OF ELEPHANT CHAP.

Eleplias alone survives. This genus also includes many extinct
forms, both American and European, as well as Asiatic and
African. The entirely extinct genera are Stegodon and Mastodon.
The group is clearly one dwindling towards extermination. From
the Middle Miocene downwards these great " pachyderms " have
existed ; and from the Miocene up to Pleistocene times they were
almost world-wide in range and numerous in species.

The genus Eleplias comprises usually large, but occasionally
(the pygmy Elephant of Malta) quite small forms. The external
features of the genus differ slightly in different species, and will
therefore be described in relation to those species which we shall
notice here. The vertebral formula is C 7, D 19-20, L 3-5,
Sa 4-5, Ca 24-30, or even more.

The bodies of the vertebrae are remarkable for their shortness
and for the very flattened articular surfaces.

The skull is large and massive. Its large and heavy character
is, as has been stated in the definition of the sub-order, due to the

Fiu. 116. — A section of the cranium of a full-grown African Elephant, taken to the left
of the middle line, and including the vomer ( Vo) and the mesethmoid (MK) ; an,
anterior, and pn, posterior narial aperture. x ^. (From Flower's Osteology.}

immense development of air cavities in the diploe ; the diameter
of the wall of the skull is actually greater than that of the
cranial cavity. These cavities are not obvious in the young
animal. They are most conspicuous in the roofing bones of
the skull, but are seen elsewhere, and thicken the basis cranii,

TUSKS AND MOLARS 2 19

the maxillae, and so forth. This state of affairs, together with
the presence of the huge tusks, has, as it were, pushed back the
nasal orifices to near the top of the skull in a very Whale-like
fashion. As in the Cetacea, the nasal bones are limited in size,
and the premaxillae send up processes to join the frontals and
the nasals. There is a straight and somewhat slender zygomatic
arch, but the orbit is not separated from the temporal fossa. The
malar bone is small, and, as in Eodents, forms the middle part
of the zygoma. This is not the case with most Ungulata. The
symphysis of the mandibles forms a spout-like rim. The scapula
has a narrow prescapular, but a very wide postscapular region.
The spine has a strong process projecting backwards from near
its middle ; this is a point of likeness to certain Eodents. No
Elephant has a clavicle. The most remarkable feature about
the fore-limb is the separation and crossing of the radius and
ulna. The arms of these animals are permanently fixed in the
position of pronation. The foot is short, and the bones of the
carpus are serially arranged. There are, however, traces of a
commencing interlocking of these bones in many forms. The
hind-feet are somewhat smaller than the fore-feet, and the tibia
and fibula are both developed.

As to the teeth, this genus is to be distinguished from allied
forms by the presence of tusks in the upper jaw only. These
tusks have no bands of enamel such as characterise those of
Mastodon. They are incisors. There is, however, a trace
of the former enamelling in the shape of a patch at the tip,
which soon wears away. The molar teeth of Elephas are so
large that the jaws cannot accommodate more than at the
most two and a part of a third at a time. These are gradu-
ally replaced by others to the number of three, the replace-
ment of teeth suggesting that of the Manatee. Each molar is
deeply ridged, the interstices between the ridges being filled up
with cement. As the tooth wears away, therefore, the surface
continues to be flat. Each ridge consists of a core of dentine
surrounded by a coat of enamel. The number of these ridges
varies greatly from species to species. The Indian Elephant is
one of those which have the greatest number of plates in a single
tooth, as many as twenty-seven.1 Of the six molars which eventu-

1 It must be borne in mind that the teeth increase in complexity, those first
pushed up having the feAvest plates. The first has only four transverse plates.

22O STOMACH AND BRAIN CHAP.

ally appear, the first three are considered to correspond to pre-
molars. But successional teeth are rare in the genus ; that is to
say as far as concerns the molars, for the tusks have their milk
forerunners. As to the molars it is apparently only E. planifrons
which certainly shows a milk dentition. In Mastodon and older
types a milk dentition is commoner.

The viscera of the Elephant have been examined by many
zoologists. The latest paper, dealing chiefly with the African
species, but containing facts about its Indian congener also, is quoted
below.1 The Elephant is remarkable in possessing, in addition
to the three usual pairs of salivary glands present in mammals, a
fourth, situated in the molar region, and opening on to the cheek
by many pores. This gland is especially well developed in Rodents.
There is a gland which may be mentioned in this connexion, though
it opens externally between the eye and ear, known as the temporal
gland ; its use does not seem clear. The thoracic cavity of the
Elephant, as may be inferred from the large number of ribs, is
very large as compared with the abdominal.

The stomach is simple in form, and the epithelium of the
oesophagus does not extend into it as is the case with the Horse
and Ehinoceros. A gland or a collection of smaller glands occurs
in the stomach, and recalls the " cardiac gland " of the Wombat
and the Beaver, also that of the Giraffe. The large intestine is
long, rather more than half the length of the small intestine.
The caecum is well developed in these animals. The liver has a
very simple form, being but slightly lobulated. It is actually
only bilobed, but it is important to notice that this division does
not correspond to the two halves of the liver. As shown by
the attachment of the suspensory ligament, one half consists of
the left lateral lobe alone, the other half embracing the remaining
primary lobes. The simplicity of the liver looks like an archaic
character. No Elephant has a gall-bladder. The lungs again are
simple in form through their slight lobulation. Each half in
fact is without subdivisions, and is of a triangular form. In
this the Elephants resemble the Whales, as in the simple liver.
In both cases probably the likeness is due to the permanence of
primitive features of organisation. The brain 2 of the Elephant

1 Forbes, Proc. Zool. Soc. 1879, p. 420.

2 See Krueg, Zeitschr. wiss. Zool. xxxiii. 1881, p. 652, and Beddard, Proc. Zool.
Soc. 1893, p. 311.

ix THE AFRICAN ELEPHANT 221

has hemispheres which are extremely well convoluted ; but they
leave the cerebellum entirely uncovered. This suggests a brain
which is a great specialisation of a low type. The brain has
been particularly compared with that of the Carnivora, with
which group the Elephants agree in the characters of the
placenta. It is, however, always a matter of the very greatest
difficulty to compare the brains of mammals belonging to different
orders.

There are but two living species of Elephant, of both of
which we shall now proceed to give some account. Only a few of
the rather numerous fossil forms can be touched upon here.

The African Elephant, E. africanus, has been sometimes re-
ferred to a distinct genus or sub-genus, Loxodon, by reason of the
lozenge-shaped areas on the worn grinding-teeth. It lives, as its
name denotes, in Africa. This species has a number of external
features which enable it to be distinguished from the Oriental
Elephant. The head slopes back more, and has not the two
rounded bosses which give so wise a countenance to the Indian
species. The ears are very much larger. The tip of the trunk
has a slight triangular projection on both the lower and the upper
part of the circumference of the aperture. There are four nails
on the fore-feet and three on the hind. As in the Indian form,
the toes are all bound together, and do not appear for any part as
free digits. A thick pad of fat, etc., makes the animal when
alive look as if plantigrade, whereas it is, as a matter of fact,
digitigrade. In internal features the most prominent difference
from E. indicus is in the molar teeth, which are ridged by much
fewer ridges. The outside number for a single tooth in the
present species is 10 or -11. In Eleplias indicus on the other
hand there are as many as 27.

The African Elephant, thinks Sir Samuel Baker, reaches a
height of about 12 feet, and it will be remembered that the
notorious "Jumbo" was found to be 11 feet high at the
shoulder. The tusks are found in both sexes, as in the Indian
beast, but are relatively larger in the female in the species now
under consideration. It is also a rather more active creature, and
is more savage ; * however it can be tamed, as is shown by several

1 So convinced are some persons of the untameable character of the African
Elephant, that it has even been suggested that the animals with which Hannibal
crossed the Alps were not E. africanus, but a now extinct species !

222

THE ELEPHANT AS AN EMBLEM

CHAP.

specimens which have been and are in the possession of the Zoologi-
cal Society, and other proprietors. It was apparently used in the
past. Certain Carthaginian coins are stamped with a figure of

the African Elephant ; but in Africa no attempts are now made
to utilise this creature except for food and ivory.

The meaning of an Elephant as an emblem upon a coin
appears to be eternity, and there is no question but that the

WEIGHT OF TUSKS 223

Elephant is a long-lived animal. It is said that it hardly reaches
proper maturity before forty, and that 150 years is not beyond
probability in the way of longevity. Even longer periods have
been assigned to it.

The tusks of the Elephant are by no means necessarily sexual
adornments, used for fighting purposes only. The African
Elephant is a most "industrious digger," and grubs up innumer-
able roots as food. It appears to be a fact that during these
operations the right tusk is mainly used, and in consequence that
tusk is shorter as well as thinner than the other. Two average
tusks would weigh respectively 75 and 65 Ibs., the latter of
course being the weight of the more worn right tusk. These
weights, it should be observed, by no means indicate the limits
to which finely-developed tusks can attain. The very heaviest
tusk known to Sir Samuel Baker l weighed 188 Ibs. This was
sold at an ivory sale in London in the year 1874. The pace of
the African Elephant, says the same authority, is at most at the
rate of fifteen miles an hour at first, and of course in a furious
rush. This pace cannot be kept up for more than two or three
hundred yards, after which ten miles an hour is a better ap-
proximation to the rate which can be kept up for long
distances.

The Indian Elephant, Eleplias indicus (or Euelephas indicus, if
the genus Loxodon is to be accepted), is better known and has
been longer known than the African. It occurs in India and
Ceylon, and in some of the Malayan islands, the Elephants of
which latter parts of the world have been regarded as a distinct
form, an apparently unnecessary procedure.

This species does not stand so high at the shoulder as the
African ; its back is more rounded in the middle. The trunk has
but one pointed tip ; there are five nails on the fore- and four on
the hind-feet: As this species comes from India and the East, it
has been longer as well as better known than the African form.
Thus many of the stories and legends that have congregated round
Elephants apply really to this form. As is well known, the Indian
Elephant is much used as a beast of burden, and for other purposes
where its huge strength renders it invaluable. But its great draw-
back as a servant of man is its great independability. On the
one hand we have furious, vicious, and generally unreliable
1 Wild Beasts and their Ways, London, 1890.

224

TIMIDITY OF INDIAN ELEPHANT

Elephants, and on the other perfectly docile creatures, who obey
the slightest hint from their driver. Huge though the Elephant

is, it is frequently a timid beast. Sir Samuel Baker relates how
one which he was riding fairly bolted at the sight of a Hare. To

ix ARISTOTLE'S OBSERVATIONS 225

be bolted with by an Elephant is far from pleasing, though a
rather exciting event. It makes for the nearest jungle at once,
being, much more than the African species, an inhabitant of forest.
And in rushing through the dense undergrowth, the occupiers of
the Elephant's back are apt to be swept off or cut to pieces by
innumerable thorns.

Elephants, no doubt of the Indian species, were used by the
Persians in battle, and from fifteen which were captured at the
battle of Arbela some notes were drawn up by Aristotle. In
stating that the animal reaches an age of 200 years, the
naturalist and philosopher was probably not very far out. The
mode of Elephant-catching as related by Aristotle is that pur-
sued at the present day. Then, as now, tame Elephants were
made use of as decoys. Pliny,1 who was apt to confound fact and
fiction in a somewhat inseparable tangle, had something to say
about Elephants, both Indian and African. Serpents, he thought,
were their chief enemies, which slew them by coiling round them
and thrusting their heads into the trunk, and so stopping respiration.
In Europe Elephants were first seen in the year B.C. 280. Pyrrhus
used them in his invasion, and copying his example the Romans
themselves learnt to use Elephants. The first Elephant seen in
England arrived in the year 1257, presented by the King of
France to Henry III. It was kept in the Tower (for long after-
wards a menagerie), and died at twelve years of age. Much use
of the Elephant has been made in symbols. We have spoken of
the African Elephant on Carthaginian coins as an emblem of
eternity. The Oriental Elephant resting on the back of a tortoise
and supporting the world is the same idea ; and it is instructive
to note that remains have been found in the Siwalik Hills of a
tortoise which would have been actually big enough to support
the creature, even " Jumbo," who weighed 6^- tons. Another
symbol is that of an Elephant upon whose back is a child with
arrows ; this occurs on a medal of the Emperor Philip. It can
perhaps hardly signify the eternity of a strong human feeling !

The intelligence of the Elephant has been both exaggerated and
minimised. Perhaps the most elaborate attempt to endow the
beast with unusual mental perceptions is that of Aelian, who
related that an Elephant carefully watching his keeper, wrote after
him with his trunk letters upon a board. That the animal does

1 See Natural History of the Ancients, by Rev. M. G. Watkins, London, 1896.

VOL. X Q

226 THE MAMMOTH AND THE FLOOD CHAP.

possess a good deal of brains, seems to be shown by the way in
which a well-trained animal will obey the slightest sign of the
mahout in India. According to Sir Samuel Baker, localities
which produce in abundance particular kinds of fruit are remem-
bered, as well as the time at which the fruit will be at its best.
Stories of revenge, which are numerous enough, attest, so far as
their data are to be accepted as accurate, the power of memory
possessed by the Elephant.

In spite of their longevity, however, Elephants, unlike Eome,
have not been built for eternity. We can only find two living
species ; but in past times Elephants were very numerous. They
commenced, so far as we know, in the Miocene.

The existing forms are known in a fossil, or at least sub-fossil
state, from diluvial deposits ; and it is interesting to note that the
African Elephant had formerly a wider range than now. Its bones
(described as E. prisons) have been met with in Spain and Sicily.

One of the best known of completely-extinct Elephants is the
Mammoth, E. primigenius. This great Elephant in most respects
more nearly approached the existing Indian Elephant. The teeth
have quite as numerous plates. The tusks were enormous, reach-
ing a maximum length of 1 5 feet ; they were much curved upwards
as well as outwards. A large tusk weighs as much as 250 Ibs.
The Mammoth was of exceedingly wide range. Not only was it
found in various parts of Europe, but it was especially abundant
in Siberia, as is exemplified by the fact that for the last two
hundred years as many or more than 100 pairs of tusks annually
have been sold from that region. It also occurred in America
together with forms at least not far removed from it, such as E.
columbianus. Mammoths have been more than once found as
entire carcases in the frozen soil of Siberia. The first was dis-
covered in the year 1799, and rescued some years later for the
St. Petersburg Museum. This example showed that the Mammoth,
unlike existing Elephants, was covered with thick wool mingled
with long and more bristly hairs of some 10 inches in length.
The softer wool formed a kind of mane beneath the neck, which
hung down as far as the knees. Another carcase was discovered
later by Lieut. Benkendorf, who did not save it, but was nearly
swept along with it into the sea by a flood. These creatures
died in the position in which they were found by being bogged
when in search of vegetation or water.

ix " CORNES DE LICORNE " 227

How primeval man, with his inferior weapons, slew the
Mammoth is not easy to understand ; but that they were con-
temporaneous is clearly shown by associated remains, and by the
notorious sketch of the Mammoth on a piece of its own ivory, in
which curved tusks and a forehead like that of an Indian Elephant
are plainly to be seen. Although it was only so recently as the
year 1799 that an example of this great creature was actually
studied on the spot, and removed to St. Petersburg, the existence
of Mammoths and of ivory is a matter of much more ancient
knowledge. M. Trouessart relates l that fossil ivory was known
to the Greeks. Theophrastus spoke of ivory imbedded in the
soil, and the tusks were recovered by the Chinese. It is a curious
fact that the Chinese described and figured the Mammoth as a
kind of gigantic Eat. The likeness between the elephantine molar
and that of Eodents has been commented upon ; but the existence
of its tusks below the level of the ground led the Chinese Natural
Historians to consider that the ways of life of the Mammoth were
those of the Mole. As to the carcases themselves, the Chinese
said that the flesh was cold, but very healthy to eat. This
expression can hardly be explained, except upon the view that
fresh carcases were known to that people long before they were
known to us of the Western world. The value of the Mammoth
ivory was known to antiquity ; the famous Haroun-al-Easchid
gave to King Charlemagne not only a pair of living Elephants,
but a " horn of Licorne," which seems undoubtedly to have been
a name for the tusks of the Mammoth. For in an account of the
sacred treasures of Saint Denis, published in the year 1646, the
author states this to be the fact.

The causes of the disappearance of the Mammoth are not easy
to understand. Some held that it was a naked animal like the
existing Elephants, and that the lowering of the temperature in
Siberia proved fatal; it is, of course, now certain that it was
clothed with dense woolly hair. Along with the bogged corpses
of the great pachyderm, numerous trunks of pine-trees have been
found, together with associated remains of other animals now
extinct in that neighbourhood. Thus it is plain that Siberia was
once covered by mighty forests, through which the Mammoth
roamed. The decay of these forests, upon whose branches the
Elephant fed, as is attested by the remains of pine leaves found

1 Bull. Soc. Nat. d'Acclimat. xlv. 1898, p. 41.

228 ELEPHANTS IN BRITAIN CHAP.

in the interstices of its teeth, was the signal for the disappearance
of their most colossal inhabitant.

The large number of remains of this and of other extinct
species of Elephas in this country gave rise to the supposi-
tion that they were Elephants brought over by Caesar to aid
in the subjugation of these islands. The Eev. J. Coleridge
(father of the poet) pointed out that though Caesar in his
Commentaries made no mention of any such importation of
Elephants, a passage in the Stratagems of Polyaenus expressly
mentions that Cassivelaunus was confronted by the Romans with
an Elephant clad in a coat of mail, by whose aid the crossing of
the Thames was effected. At the time that attention was called
to this (1757) it was not popular to hint at the possibility
of fossils. So that fact, conveniently historical, served to
explain away a difficulty. It is remarkable that the Elephant,
common enough of course in Asiatic monuments, actually occurs
in English architecture. Mr. "Watkins, from whose interesting
work (Natural History of the Ancients) a good many of the facts
detailed here are drawn, tells us that the church of Ottery St.
Mary has an Elephant's head sculptured on one of its pillars.
The same ornament appears in Gosberton Church, Lincolnshire.
Whether this has anything to do with a. reminiscence of formerly
existing Elephants is a hard question to answer. In this figure
of an Elephant the trunk has a spiral representation, and the
trunk of an Elephant is believed by some to be intended by the
common " so-called Pictish ornamentation " in Scotland ; this
spiral alone is to be seen constantly. If it is a reduction of an
Elephant to its simplest terms, it is highly interesting as an
almost undoubted survival of remembrance of Elephants. For at
such a period we cannot use the memories of Crusaders or others
who may have visited the East to explain the facts. The
sculptured Elephants' heads might conceivably be so explained.

The name Mammoth, thinks Mr. Watkins, may be derivable
from the Arabic word Behemoth. He quotes a writer, who first
described the beast in 1694, as using the two words indifferently.
The Arabs, moreover, were then as they are now great ivory traders ;
and in the ninth and the two succeeding centuries explored
the confines of Siberia, as they now do the forests of Africa, for
ivory. The " Behemoth " of Job " eateth grass as an ox. ... He
moveth his tail like a cedar " (the Hippopotamus has a much more

BEHEMOTH

229

stumpy appendage). " Behold, he drinketh up a river, and hasteth
not " is surely much more suggestive of the copious draughts of
an Elephant than the possibly equally copious hut not so visible
libations of a Hippopotamus.

The most ancient of the true Elephants (genus Elephas} is
E. meridionalis. It is of the African type, i.e. the plates of the
molar teeth are not abundant, and are not so many as in the
existing E, africanus. It seems to have been one of the largest of
Elephants, standing 4 metres high. Its remains are abundant in
Europe, and are known also from England. Like this species
E. antiquus is also of the African type. It was contemporary
with man. Certain dwarf or " pony " races found in caves in
Malta, and called Elephas melitensis or E.falconeri, are believed
to belong to this species. Mr. Leith Adams, who described these a
remains, placed them in two dwarf species called by the names
used above, and found associated with them a larger form, which
he referred to E. antiquus. The existence of these animals in
Malta seems to argue at least its former larger dimensions, and
the presence of more abundant fresh water. The remarkable
swimming capabilities of the Elephant do not necessarily imply
either a former absence of land connexion or, on the other hand,
its existence. Nor as a third possibility can it be suggested that
the dwarf size argues an island of limited dimensions, when we
bear in mind the huge tortoises of the Galapagos and some other
islands. It is important to notice that Elephants of the African
type (Loxodoii) were not formerly absent from India. E. planifrons
was one of these.

The genus Stegodon is so called from the fact that the molar
teeth, seen in longitudinal section, present a series of roof-shaped
folds, the interstices between which are not, or are, imperfectly
filled up with the cement which in Elephas reduces the surface of
the teeth to a level plane. This genus is exclusively Asiatic, and
is Miocene to Pleistocene in time range. The number of ridges
on the molars is small, not more than two. The incisors (tusks)
have no enamel ; the skeleton generally is like that of Elephas,
between which and Mastodon the present genus is intermediate.
Among the four or five species is S. ganesa (called after the Indian
Elephant-headed divinity), with tusks 10 feet long, to be seen at
the British Museum of Natural History.

1 Trans. Zool. Soc. ix. 1874, p. 1.

230 THE BONES OF TEUTOBOCHUS CHAP.

The last genus of the family Elephantidae is Mastodon, so called
from the structure of the molar teeth. These are provided with
but few transverse ridges, not more than five, so that their structure
is intermediate between those of Dinotherium and those of Stegodon.
Between the ridges are sometimes isolated, boss-like protuberances
(whence the name of Mastodon), produced by a subdivision of the
ridges. There is either but little or no cement between the ridges.
This genus differs from nearly all other Elephantidae by the posses-
sion of milk molars, which occasionally persist throughout life, the
permanent dentition in those cases being a mixture of milk
and permanent teeth, as has been (erroneously) stated of the
Hedgehog.1

The tusks (incisors) are sometimes present in both jaws, and
as they have, during youth at any rate, a coating of enamel, the
likeness to the chisel-shaped incisors of Eodents is patent. In
connexion with the implantation of incisors in the lower jaw,
many species have a prolongation of the bones of that part of
the skeleton. In the bones, generally, there is not very much
difference from Elephas, but the forehead is a little less pro-
nounced. The genus existed from the Miocene and became
extinct in the Pleistocene. It was nearly world -wide in range,
being known from all four continents. Naturally with this very
wide range was associated a large number of species. Zittel
enumerates no less than thirty-two..

This genus is the only one of the Elephantidae which extended
its range into South America, where the remains of two species
occur. The bones of these great Elephants have attracted attention
for some centuries. They were often held to be the bones of
giants (as they actually were !), and in one case were ascribed to a
deceased monarch, Teutobochus. The American Indians considered
that equally gigantic men lived who were able to combat these
great Proboscideans. There are legends of the Mastodons as living
animals, which is quite probable, considering their geological age.
There is a curious parallelism between the legends of two such
widely-separated localities as North America and Greece. Buffon
relates how among the Indians of Canada there was a belief that
the Great Being destroyed both Mastodons and men of equal
proportions, with thunderbolts. With this we may perhaps com-
pare the story of the destruction of Typhoeus by Zeus, who
1 See Busk in Trans. Zool. Soc. vi. 1868, p. 227.

DINO THERIUM 2 3 I

also used thunderbolts. One of the giants was not slain, but
was compelled to stand and bear up the heavens. Atlas holds
thus the position of the Elephant supporting the globe of Indian
mythology.

The genus Dinotherium, sole representative of the family
Dinotheriidae, differs in a number of important particulars from
the true Elephants. In the Elephants, if there is but a single
pair of incisors, these are found in the upper jaw ; in Dino-

Fifi. 119. — Dinotherium giganteiun. Side view of skull, -fgtii iiatural size.
Miocene, Germany. (After Kaup.)

therium there is apparently but a single pair, but these are
implanted in the lower jaw, the symphysis of which is much
prolonged and greatly bent downwards, so that the tusks emerge
at right angles to the long axis of the head, and are even bent
backwards. The molar teeth are five in number on each side of
each jaw and are bi- or tri-lophodont, not unlike those of the Tapir.
There is no cement in the valleys between the ridges of these
teeth, and there is a. regular succession, the premolars being two
and the molars three.1 All the teeth are in use at the same time,

1 There are, however, three milk forerunners of the premolars, of which one
has no successor.

232 POSITION OF D1NOTHERIUM CHAP.

their small size enabling them to be accommodated in the jaw
together. The skull of Dinotherium is lower than that of Elephas
or Mastodon. The bones of the skeleton generally are like those
of Elephas.

Though a suggestion of marsupial bones attached to the pelvis
has been discredited, there is no doubt that Dinotherium. occupies
the most primitive position among the Proboscidea ; but at the
same time it cannot be regarded as the ancestor of Elephants, as
it is so much specialised in various ways. The incisors for one
thing forbid this way of looking at the creature. It is an ancient
genus found in beds of Miocene age in Europe and Asia. It is
not known from America. The creature was larger than any
Elephant. Eighteen feet in length has been assigned to it. The
enormous weight of the lower jaw and tusks seems to argue that
it was at least partially aquatic in habit, and that it may have
used these tusks for grubbing up aquatic roots or for mooring
itself to the bank. At first there were naturalists who considered
it as an ally of the Manatee, and the skull is not unsuggestive of
that of the Sirenia.

Pyrotherium has been referred to the Proboscidea ; but our
knowledge of that form is limited to a few teeth from Patagonian
rocks of an uncertain age.1 They are simple bilophodont molars,
very like those of Dinotherium. A tusk has been found in the
neighbourhood of these teeth which may possibly belong to the
same animal ; but it is uncertain.

SUB-ORDER 7. HYEACOIDEA.

This group of small mammals contains only one well-marked
genus which is usually named Hyrax, although Procavia seems to
be the accurate term. Popularly these creatures are known as
Coneys. They have a singular resemblance to Kodents, the short
ears and much reduced tail, besides the squatting attitude adopted,
contributing to this merely skin-deep likeness. They agree with
other Ungulates in the structure of the molar teeth, which are
much like those of Rhinoceros ; in the absence of a clavicle ; in
the absence of an acromion ; in the reduction of the digits of the
limbs to four digits in the manus and three in the pes. On the
1 Lydekker, An. Mus. La Plata, Pal. Arg. iii. 1894.

THE CONEYS 233

other hand they differ from most Ungulates in the incisors grow-
ing from persistent pulps, a point in which they resemble the
Eodentia. The muffle also is split as in those animals. The
Hyracoidea are peculiar in the fact that in addition to the caecum
at the junction of the small and large intestines, there are a pair
of caeca (bird-like in being paired) some way down the large
intestine. The dorsal vertebrae are unusually numerous, 22. The
adult dentition according to Woodward,1 who has recently ex-
amined the matter, is I \ C (^) Pm ^ M ^, while the milk
dentition is I f C \ Pm J- .

The inclusion of the canine of the permanent set of teeth in
brackets signifies that it is the milk canine which occasionally

FIG. 120.— Cape Hyrax. Hyrax capensis. x J.

persists. It should further be remarked about the teeth that
they are both hypselodont and brachyodont, the extremes being
connected by intermediate forms. Another peculiarity of the
genus is the dorsal gland, which is covered with hair of a different
colour to that covering the body generally. This is present in all
species.

The genus Hyrax (the most recent authority on the subject,
Mr. Oldfield Thomas,2 only allows one genus) is limited in its
range to Ethiopian Africa and to Arabia, including Palestine,
It does not reach Madagascar. Mr. Thomas allows fourteen species
with two or three sub-species.

1 M. F. Woodward "On the Milk Dentition of Procavia (Hyrax) capensis,
etc," Proc. Zool. Koc. 1892, p. 38.

2 "On the Species of the Hyracoidea," Proc. Zool. Soc. 1892. p. 50.

234 HYRAX CHEWING THE CUD CHAP, ix

Some of the Coneys live in rocky ground, while others, formerly
placed in the genus Dendrokyrax, frequent trees, in holes in which
they sleep. The Coney of the Scriptures is familiar, who is " exceed-
ing wise," though a " feeble folk." But the further observation
that he " cheweth the cud but divided not the hoof," is obviously
entirely wrong. As to the wisdom, it is said that this beast is
too wary to be taken in traps ; while the suggestion of chewing
the cud is, according to Canon Tristram, to be interpreted in the
light of a habit of working and moving its jaws which the animal
has. The traveller Bruce kept one in captivity to see if it did
reallv chew the cud, and found that it did !

CHAPTER X

UNGULATA (continued} — PERISSODACTYLA (ODD-TOED

UNGULATES) L1TOPTERNA

SUB-ORDER 8. PEEISSODACTYLA
THESE Ungulates derive their name, which is that given by the late

FIG. 121. — Bones of the maims A, of Tapir (Tapirus indicus). x i. B, of Rhinoceros
(Rhinoceros sumatrensis). x £. C, of Horse (Equus caballus}. x J. c, Cunei-
form ; I, lunar ; m, magnum ; p, pisiform ; JR, radius ; s, scaphoid ; id, trapezoid :
tm, trapezium ; u, unciform ; U, ulna ; //- V, second to fifth digits ; V in B, and //
and /Fin C, represented by rudimentary 'metacarpals. (From Flower's Osteology.)

Sir Richard Owen, from the fact that the middle digit of the hand
and foot is pre-eminent. As will be seen from Fig. 121, the axis of

236

FOOT OF UNGULATES

the limb passes through the third finger, which is larger than any of
the others, and is symmetrical in itself. In this the present group
contrasts with the Artiodactyla, where the axis is not " mesaxonic,"
but where there are two digits, on either side of the axis, which are
symmetrical with each other. This arrangement of the limbs is
highly characteristic, but appears to be not quite universal. In
the Titanotheres, which form a group of the Perissodactyles, the
fore-limbs are not quite accurately mesax-
onic. Nor on the other hand can all Ungu-
lates which show the Perissodactyle condition
be safely included in the present group. The
ancient Condylarthra and the Litopterna
show precisely the same state of affairs.
But other features in their organisation lead
to their separation from the Perissodactyles,
of which, however, the Condylarthra are
probably ancestors. The Litopterna on the
other hand, which possess even one -toed
members like Equus, are believed to repre-
sent a case of parallelism in development.
The number of functional toes varies from
four to one. In the ankle joint the astra-
galus either does not, or does only to a
comparatively slight extent, articulate with
the cuboid as well as with the navicular
FIG. 122. — Bones of the bone. Moreover the fibula when present

manus of Camel (Camelus j i ,• -i , -.r ,1

bactrianus}. x £. c ^oes not as a ru^e articulate with the
Cuneiform ; i, lunar ; m, calcaneum. In the opposed group of

magnum ; R, radius ; s, . ,. , , ., . „ ,

scaphoid ; td, trapezoid ; Artiodactyles the precise reverse ot these
«, unciform. (From conditions obtains. It is usually stated

Flower s Osteology. ) . . .

as part ot the definition 01 this group

that they do not possess horns of the type of those met
with in the Cervicornia and Cavicornia. But the strong bony
bosses on the skull of. many Titanotheres, so curiously remin-
iscent of those of the not nearly related Dinoceras and Proto-
ceras, may well have supported horns of the Ox and Antelope
pattern.

The teeth of the Perissodactyles are lophodont, more rarely
bunodont. The selenodont Artiodactyle form of molar is not
met with, The dental formula, moreover, is at least near the

DIVISIONS OF PERISSODACTYLES

237

complete one, the more modern forms as usual being the more
deficient in numbers of teeth.

The dorso-lumbar vertebrae are as a rule twenty-three ; but
the extinct Titanotheres are again an exception ; for, at least in
Titanotherium, there are but twenty of these vertebrae — an Artio-
dactyle character. The femur has a third trochanter. There are
so few recent Perissodactyles that an enumeration of the dis-
tinguishing characters of the viscera may very probably be use-
less for purposes of classification. But
the living genera at any rate are to be
separated from the living Artiodactyles
by the invariable simplicity of the *
stomach coupled with a very large and
sacculated caecum. The liver is simple
and not much broken up into lobes, and
the gall-bladder is always absent. The
brain is well convoluted. The teats are
in the inguinal region. The placenta in
this group is of the diffused kind.

The living Perissodactyles belong to
three types only, indeed to three genera
only (in the estimation of most), which
are the Horses, Tapirs, and Rhinoceroses.
But taking into account the extinct

forms, they may be divided primarily Fia 123. -Anterior aspect of
(according to. Professor Osborn) into the
four following groups : — (1) Titano-
therioidea, including but one family,
Titanotheriidae ; (2) Hippoidea, includ-
ing the families Equidae and Palaeotheriidae ; (3) Tapiroidea,
with two families, Tapiridae and Lophiodontidae ; and (4) Ehino-
cerotoidea with families Hyracodontidae, Amynodontidae, and
Ehinocerotidae. It is conceivable, according to the same writer,
that the Chalicotheres (here treated of as a separate sub-order,
Ancylopoda) should be added to the Perissodactyle series.

Fam. 1. Equidae. — This family, which includes the living
Horse, Zebras, and Asses, as well as a number of extinct genera
agreeing with those types in structure, may be defined by the
possession of but one functional toe, the two lateral ones being mere
splints, or but little more. The molar teeth are hypselodont, and

right femur of Rhinoceros
(Rhinoceros indicus). x A.
h, Head ; t, great trochanter ;
t', third trochanter. (From
Flower's Osteology.)

238

TEETH OF HORSE

CHAP.

the premolars, with the exception of the first, resemble the molars
in their pattern. The orbit is completely surrounded by bone.
The incisors are chisel-shaped, with a pit on the free surface.
The canines are rudimentary if present. The radius and ulna
are fused, as are the tibia and fibula. Although for the sake of
uniformity a family, Equidae, is here separated from its allies, it
is quite impossible owing to the full state of our knowledge of
this group to draw a really hard-and-fast line between this family

FIG. 124. — Side view of skull of Horse with the bone removed so as to expose the whole
of the teeth, c, Canine ; Fr, frontal ; i1, i2, &, incisors ; Z, lachrymal ; m\ m2, m3,
molars ; Ala, malar or jugal ; MX, maxilla ; No, nasal ; oc, occipital condyle ; Pa,
parietal ; pml, situation of the vestigial first premolar, which has been lost in the
lower, but is present in the upper jaw ; pm?, prr<?, pm*, remaining premolars ; PMx,
premaxilla ; pp, paroccipital process ; Sq, squamosal. (After Flower and Lydekker.)

and the Palaeotheriidae. We shall deal presently with the con-
jectured pedigree of the Horse, which naturally involves that
family, and which presents an unbroken series from four -toed
Perissodactyles to the present one-toed Horse, the various bones
and teeth becoming modified in the course of the descent " with
the regularity of clockwork." We are compelled to draw the
line at functional second and third toes ; directly these are no
longer used the animal is a Horse in the strict sense ! This is
irrational and regrettable, but necessary for practical purposes, if

x DEFINITION OF GENUS EQUUS 239

we are to continue the plan of defining the various families of
Mammalia.

The genus Equus 1 contains not only the Horse, but the Asses
and Zebras. The genus is to be distinguished as regards external
characters by the following features : — The body is thickly
clothed with hair ; there is a more or less bushy tail and mane ;
the colours are apt to be disposed in stripes of black or blackish
upon a yellowish brown ground ; this is of course best seen in
the Zebras, but the wild Asses also have some traces of it, if only
in the single cross-bar of the African Wild Ass, and it is even
" reversionary " in the domestic Horse at times. There are no
horns upon the forehead or elsewhere ; the fore -limbs or both
pairs have a callous pad upon the inside, which is possibly to be
looked upon as an aborted gland, possibly originally 'of use as
secreting some odorous substance calculated to enable strayed
members of the herd to regain their companions. The terminal
phalanx of each of the (functionally) single digits is enclosed in
a large horny hoof.

The main internal features of structure which divide this
genus of Perissodactyles from the Ehinoceros or the Tapir, or
from both, are : the existence of strong incisors, three on each
side of each jaw ; there are canines, but these are small and do
not always persist in the full-grown mare. They are popularly
known as " tusks " or " tushes." The first of the four premolars
(the " wolf tooth ") is small and quite rudimentary ; it is often
absent. As there are three molars, the present genus has the
" typical " number of the Eutherian dentition, i.e. forty -four. In
the skull the orbit is — as it is not in Tapirs and Ehinoceroses —
completely encircled by bone. There is but one functional finger
and toe on each hand (Fig. 121 C) and foot ; the second and third
digits are represented by mere splints, one of which may as an
abnormality be enlarged, and reach nearly as far as the well-
developed digit. There are even occasionally traces of digit
number two.

The Horse, E. caballus, is to be distinguished from its con-
geners by the small callosities on the hind-limbs which it pos-
sesses in addition to the larger ones on the fore-limbs. The
hairy covering of the tail is more abundant, as is also the mane.
The head too is proportionately smaller, and the general contour

1 Sir W. H. Flower, The Horse, London, 1890.

•

240 STRIPES ON HORSES

more graceful. Though Zebra markings are not usual upon E.
caballus, there are plenty of examples of — what we may perhaps
in this case term — a " reversion " to a striped state. The cele-
brated " Lord Morton's mare," 1 whose portrait hangs in the Royal
College of Surgeons, is an interesting case of this. It was as a
matter of fact thought to be an example of that rather doubtfully-
occurring phenomenon, " telegony." Its history is briefly this.
The animal was the offspring of a mare that had previously pro-
duced to a male Quagga a hybrid foal. Afterwards a second foal
was produced by the same mare to an Arab sire. This foal, the
one in question, was striped, and hence was thought to be an
example of male prepotency. But instances are known of un-
questioned Horses which show the same stripes, such as a Norway
pony which had not even seen a Zebra !

A last remnant of the naked palm of the hand and sole of the
foot is left in the shape of a small bare area, smaller in the Horse
than in the Asses, known technically as the " ergot," the term being
that of the French veterinarians. As already mentioned, the Horse
differs from the Asses and Zebras in the fact that the hind-limbs
have callosities on the inner side. They are known as " chestnuts,"
and their nature has been much disputed. It has been suggested
that they are the last rudiment of a vanished toe ; but in all
probability they are, as already suggested, traces of glandular
structures, which are common upon the limbs in many animals
(see above, p. 12).

It is a singular fact that there are apparently no wild Horses
of this species. The case is curiously analogous to that of the
Camel, which also is only known as feral or domesticated. Why
the Horse should have become extinct as a wild animal, consider-
ing that when it does run wild it can thrive abundantly, is im-
possible to understand. Sir W. Flower thinks 2 that " the nearest
approach to truly wild horses existing at present are the so-called
Tarpans, which occur in the Steppe country north of the sea of
Azov between the river Dnieper and the Caspian. They are
described as being of small size, dun colour, with short mane and
rounded obtuse nose." But he adds that there is no evidence to
prove whether they are really wild. In favour, however, of their
possibly being wild and indigenous European Horses, may be

1 See Ewart, The Pcnicnik Experiments, Constable and Co., 1899.
2 The Horse, London, 1890.

DOMESTIC RACES 241

mentioned the fact that their general build and appearance is
highly suggestive of the wild Horses sketched by primitive man
upon ivory.

A really wild Horse, and possibly the ancestor of the European
domestic Horse, is E. przewcdskii of the sandy deserts of Central
Asia. This animal has been believed to be a mule between the
Wild Ass and a feral Horse ; but if a distinct form, and probability
seems to urge that view, it is interesting as breaking down the dis-
tinctions between Horses and Asses. The species possesses the four
callosities of the Horse, but has a poorer mane and an asinine tail.

There is no question that the Horse has been a domestic
animal for very many centuries. Hieroglyphics appear to show
that the Egyptians had not originally domesticated the Horse ; it
seems to have been first introduced among them by the Hyksos
or Shepherd Kings.1 Whatever the date may be, it is certain
that considerably anterior to the Egyptians the Assyrians and
Phoenicians possessed Horses. In Western Europe the date of
the introduction of the Horse seems to have been during the
bronze epoch. Lord Avebury2 has pointed out that out of
eighteen cases of graves in which the remains of Horse were
found, twelve contained metal implements, i.e. 6 6 per cent. This
does not of course prove that the Horse was domesticated at that
period, but it throws doubt upon the earlier occurrence of the
Horse in abundance. The Horse, however, does occur on the
Continent associated with the remains of man during the Quater-
nary period.3

Messrs. Cuyer and Alix enumerate between fifty and sixty
domesticated races of Horse, not counting the supposed wild
varieties which have been already referred to. These may be
further subdivided ; for instance, under the race " pony " we may
distinguish the Irish, Scotch, and Shetland varieties, all of which,
however, according to Sanson, have originated in Ireland. They
are used, remark the authors above quoted, " par les jeunes filles
des lords pour leurs promenades." The Arab, the Barb, the
Suffolk Punch, etc., are among the numerous races of domestic
Horses, into which to enter properly would require another
volume, and that of large size.

1 Cuyer and Alix, Le Cheval, Paris, 1886.

2 Lubbock, Prehistoric Times, London, 1865.

3 J. Geikie, Prehistoric Europe, London, 1881.

VOL. X R

242

THE ONAGER

The Asses and Zebras differ from the Horse in the characters
mentioned under the description of Equus caballus. In addition
to these may be pointed out a feature to which attention has
been directed by Mr. Tegetmeier.1 According to him the period
of gestation in the Horse is only eleven months ; in the others
more than twelve

Opinions as to the number of species of Asses differ. On
the most liberal estimate there are three Asiatic and two African

FIG. 125. — Asiatic Wild Ass. Equus onager, x -fa

species. The best known of the Asiatic Wild Asses is the Onager,
E. onager. It is of a uniform yellowish, " desert " colour, with
a dark stripe along the middle of the back, and is found in
Persia, the Punjab, and the country of Cutch. The creature is
of great swiftness ; it has been stated to be untameable, but Mr.
Tegetmeier makes the absolutely opposite statement that the
Ass occasionally " becomes so tame as to be troublesome " ! The
Syrian Wild Ass, E. hemippus, hardly, if at all, differs from this.
rlhe Kiang, E. hemionus, seems to have more claims to
distinctness. In the first place it has a more limited and a

1 Horses, Asses, and Zebras, London, 1895.

THIBETAN KIANG

243

different distribution ; it is confined to the high tablelands of
Thibet at an elevation of 15,000 feet and upwards. In correla-
tion with this habitat it has a thicker and more " furry " coat,
which is, moreover, of a darker shade than that of the Onager.
This coat is shed in the summer, and replaced by one which is
not so dark in hue. It is an interesting fact that the African
"Wild Asses approach to the zebra type in having at least traces of
stripings. There are apparently two species. The best known,

FIG. 126. — Nubian Wild Ass. Equus afncanus. x^.

the Nubian Ass, E. africanus, is probably the parent of the
domestic donkey. It has a dorsal longitudinal stripe, and
another across the shoulder — in legend the marks of the Saviour.
The matter of the name of this Ass seems difficult to decide. It
has been called also E. asinus and E. taeniopus. It has been
observed that this animal has a great aversion to water, and a
delight in rolling in the dust — both of which characteristics
argue a desert existence. But on the other hand the Kiang
will plunge boldly into streams, yet it would seem to be the
descendant of a purely desert form. The Ass is a longer-lived

244 GREVY'S ZEBRA

animal than the Horse. Mr. Tegetmeier calls attention to a
donkey living in 1893 which had been ridden fifty-five years
previously. The Horse, on the other hand, lives not much more
than twenty-five years.

A second species of African Wild Ass, E. somalicus,1 is distin-
guished by its greyer colour, by the absence of the shoulder stripe,
by the very faint development of the dorsal stripe, and by the
presence of numerous cross stripes upon the legs. It has, too,
smaller ears, and a longer and more flowing mane. Mr. Lort
Phillips, an experienced naturalist and traveller, saw a herd of
these Wild Asses in Somaliland, which he regarded as being of
quite a new species. A living example in the Zoological Society's
Gardens led Mr. Sclater to an identical conclusion, which was
supported, as he pointed out, by the fact that this Ass has a
different range to the African or Nubian Wild Ass.

Of the Zebras three species are usually allowed ; these are E.
zebra, the " Mountain " or " Common " Zebra, E. burchelli, E. grevyi,
as well as E. quagga. Professor Ewart thinks that the Common
Zebra, Burchell's, and the Quagga are not very distinctly marked
off from each other. No one, however, has any doubt of the
distinctness of E. grevyi. This latter differs from the rest in its
larger size, in the large head and ears, and in the marked hairiness
of the ears. It would seem to be a primitive type of Zebra, if
the fact that the occasional reversion of hybrids to a parent
form be allowed ; for Professor Ewart found a cross-bred Zebra to
present several characteristics in the face -marking of this, the
finest of the Zebra tribe. Only four specimens of E. grevyi have
been exhibited alive in Europe — two in Paris, and two in the
Zoological Society's Gardens in London. The latter were presented
to Queen Victoria by King Menelek of Abyssinia. The species was
named by Professor A. Milne-Edwards in honour of a late President
of the French Republic, from an example also sent by King
Menelek.

The Common Zebra has closer and darker stripes than Bur-
chell's, but not quite so close as in E. grevyi. It has also a very
characteristic arrangement of stripes on the withers in the form
of a gridiron. This latter is wanting in both the other species.
In E. grevyi, in fact, this part of the back is white. E. zebra
has also a dewlap in front. E. lurchelli has fewer and broader

1 Proc. Zool. Soc. 1884, p. 540.

BURCHELL'S ZEBRA

245

stripes, and between them lie in many cases shadow-stripes of a
taint brown.

All these animals, and the Quagga too, are absolutely
confined to Africa. Mr. E. Crawshay,1 in describing what he
considered to be a new variety, remarked upon the curiosity of
E. lurchelli. " They remain out in the sun on the plains all day
long, not retiring into covert at all. They are then an intoler-
able nuisance to any one in pursuit of other game ; indeed this
may be said of them at all times. If once they notice you, they

w

FIG. 127. — Burchell's Zebra. Kquus burchdli. x-jV-

draw in and mob you in their curiosity — only, however, when
one takes no interest in them, for when they fancy they are the
object of the intruder's attention, no animals are more watchful
and cunning in safeguarding themselves. If only their curiosity
were manifested in silence it would not so much matter, but it
vents itself in snorts and thundering stampedes, which puts every
beast within earshot on the qui vive."

Whether Burchell's Zebra 2 can be further subdivided into
species or sub-species appears to be doubtful. Dr. Matschie
considers that Equus boehmi may be regarded as a valid form,
and in addition to this two sub-species, JS. burchelli granti and

1 Proc. Zool. Soc. 1895, p. 688.
- See Pocock, Ann. Nat. Hist. (6) xx. 1897, p. 33.

246 THE QUAGGA

E. Imrchelli seloasi, have been proposed for what are at most
local races. But it is at present far from certain whether their
distribution favours this subdivision.

The Quagga was more striped than is sometimes represented in
illustrations. According to Dr. Noack, from whose paper * upon
the animal I quote here, the transverse stripes reached back as far
as the buttocks ; they were, however, completely absent from the
legs. The animal is, as every one knows, probably completely
extinct. In the year 1836 it was still abundant; in 1864 the
last specimen ever exhibited was received by the Zoological
Society. Mr. W. L. Sclater thinks that it may have survived in
the Orange Eiver Colony as late as 1878, but admits that any
certainty is difficult, as it was frequently confounded by the Boers
with Burchell's Zebra. Its rarity is emphasised by the fact that
it is not mentioned in the recent work of that most skilful of
hunters, Mr. F. Selous. Gaudry places the Quagga nearest of all
living Equidae to the Hipparion gracile of Pikermi.

Fossil Equidae. — The existing Equidae all belong to the
genus Equus, though there are some who would (quite unnecessarily)
divide off the Zebras as a genus Hippotigris. The genus Equus
itself goes back in time to the Pliocene, during which epoch there
lived in India E. sivalensis, the same species according to some
with the E. stenonis of Europe. None of these species, Old World
or New, are easily to be separated from E. cciballus. But many
names have been given to them. It is of course perfectly con-
ceivable that they may have differed among themselves as much
as do the existing Zebras and Asses, the separation of which would
be hardly possible did we know their bones only. There are,
however, extinct genera, undoubtedly related so closely to Equus
as to be placed in the same family, though clearly separable as
genera. Hipparion is one of these genera ; its remains are known
from Europe, Asia, and North Africa, from beds of Miocene and
Pliocene times. A large number of different species have been
described. It was a beast of about the size of a Zebra. The
principal characters are that each foot has three toes, of which,
however, the two side ones are smaller than the central toe.
There is a marked round fossa on the maxillary bone, a feature
shared by the South American Onohippidium.2 The pattern of

1 "Das Quagga," Zool. Garten, 1893, p. 289.

'* Of this Horse, remains have been lately discovered (see Lonnberg, Proc. Zool.

x EVOLUTION OF THE HORSE 247

the molar teeth is, too, a little different from that of Eqwus. Proto-
hirppus of the North American Pliocene is also three-toed, but the
two additionally-developed toes are smaller than in Hipparion.
Other forms are dealt with below in connexion with the ancestry
of Perissodactyles. It is a curious fact about Hipparion, which
is not now regarded as on the direct line of equine descent, that
the edges of the enamel plates of the molars may show a com-
plicated folding very like that presented by that clearly terminal
form of Perissodactyle life, the gigantic Elasmotlierium. This is
indicative of higli specialisation, which ended in extinction.

Ancestry of the Horses. — The Lophiodontidae and the
Palaeotheriidae are two of the most interesting extinct families
of Perissodactyles ; for among them we find what would appear
to be the ancestral forms of both the existing Tapirs and Horses.
The Ehinoceroses also would seem to be derivable from the
Palaeotheriidae. The very vagueness of the characters of these
creatures, considered from a classificatory point of view, has led to
much diversity in their placing. This though gratifying to the
evolutionist is tiresome to the writer who wishes to give a
methodical account of their various characters. It will be best
perhaps not to attempt an accurate placing or to reconcile con-
flicting opinion, bub to give some salient features of osteology
which lead to the belief in their relationship to existing groups of
Perissodactyles. A book upon the history of mammals would be
incomplete without some account of that well- ascertained series of
forms which seem to connect these primitive Perissodactyles with
the modern Horse. Equus, in fact, is not only the " show horse "
of the doctrine of evolution, but also the " stalking horse."

In the Eocene of both Europe and America are met with a
number of forms from which we may start. Hyracotherium,
regarded on the one hand as the type of a sub-family of the
Equidae themselves, and on the other as a member of the family
Lophiodontidae, was a small -sized animal, three feet or so in
length ; it possesses the complete Eutheriau dentition with a slight
diastema. The orbits are not separated from the temporal fossa ;
the fore-limbs were, four-toed, the hind three-toed, with moderately
long metapodia, especially on the hind-feet. The shoulder blade

Soc. 1900, p. 379) in the cave which produced the remains of Glossotherium. A
piece of skin covered with Fox-red hair, possibly spotted with paler areas, is believed
to be a relic of Onohippidium.

248' FROM OROHIPPUS TO MIOHIPPUS CHAP.

has a well -marked coracoid process. The radius and ulna are
separate ; so too are the tibia and fibula. EoJiippus, belonging
to the same sub-family, is slightly more primitive ; for the hind-
feet have a rudiment of digit I. Orohippus is a little nearer to
the Horses in that the molar teeth have acquired a little further
advance towards the equine type. Instead of the tubercles of the
teeth remaining for the most part separate, they have fused into a
set of ridges, of which, however, the pattern is less complex than
in the modern Horses. In other respects Orohippus is much like
Hyracotherium. Pachynolophus seems to be but a synonym.
. The next stage is shown by Mesoliippus, a Lower Miocene
form, usually referred to the neighbourhood of Palaeotherium. It
has nearly lost one of the toes of the fore-foot, a rudiment only
remaining ; the metapodials, at any rate of the fore-feet, seem to
be slightly increased in length. The orbit is not encircled by
bone, but there is a strong process from the frontal, which nearly
meets the zygomatic arch.

Anchitherium, from the Upper Miocene, is not far removed in
structure from the last-mentioned form ; it is a trifle nearer the
existing Horse in several points. The ulna is further reduced and
fused with the radius below ; the rudiment of digit V is still
more rudimentary ; the two lateral digits are smaller in proportion
to the central one than they are in Mesohippus ; the fibula is
fused below with the tibia. From this form to Equus is a small
series of steps, characterised by the still further reduction of all the
digits except III, by the still further reduction of the already
rudimentary ulna and fibula, and by the increasing depth of the
molar teeth, which are of course, in Equus, hypselodont.

Another interesting conclusion may seem to follow when we con-
sider the geographical range of the ancestral Horses. Hyracotherium
and Pachynolophus occurred both in the Old and New World.
From them may have arisen the Horses of both hemispheres.
After that point there is a^ division. Mesohippus is American,
and we get at Equus in that continent through Desmatippus and
Protohippus. On the other hand there are no remains known of
Mesohippus in Europe ; and unless subsequent researches prove
the existence of Mesohippus, we have to rely upon forms which
are placed with Anchitherium and Hipparion.

It seems that in America the next genus in the direct line
of equine descent to Mesohippus is Miohippus. It is smaller in

x FROM MIOHIPPUS TO PROTOHIPPUS 249

size than Anchitherium, to be considered immediately. The
odontoid process of the axis is just beginning to assume the
characteristic spout-like shape of that of the existing Horse and
many modern Ungulates. The median digit of both fore- and
hind -limbs has become greatly enlarged as compared with the
corresponding digit of earlier forms.

It is held, however, that Amhitherium is not on the direct
line of descent either in America or in Europe, in both of which
it occurs. Its teeth are in some respects less Horse-like than in
some of the more ancient genera, to which the converse would be
expected on the descent theory. Its hoofs are much elongated
and flattened, a mark of specialisation and not appropriate to
a creature holding an intermediate position in the equine series.
Both the American (A. equinum) and the European species (A.
aureliense) are of very large size, larger than its successors, and
such " alternations in bulk are unlikely."

The genus Desmatippus of Professor Scott1 fills in the gap
between Miohippus and Protohippus. The molars and premolars
are brachyodont, but there is a thin deposit of cement in the
tooth valleys, leading towards the more complete filling of these
valleys with cement, which is found in Protohippus. This genus
of Horses, of which there is at present but one species, D. crenidens,
was three-toed, and " the lateral digits, so far as can be judged by
fragmentary remains, were still fairly developed, and though
much more reduced than in Miohippus, appear to be somewhat
less so than in Protohippus."

To recapitulate, the following is the probable series of equines
in America — Mesohippus, Miohippus, Desmatippus, Protohippus.

The development of the limbs of the Horse shows a most
interesting series of stages, which correspond in part to the
ancestral forms which palaeontology seems to prove to be the
line of .the descent of our existing Equidae. This matter has
recently been elucidated by Professor Ewart, who details the
following facts and comparisons : —

In the youngest embryo (about 20 mm. in length) the
humerus is somewhat curved, and considerably longer than the
radius and carpus taken together. The first-named bone is
shorter in the adult, and the proportions of that bone in the
young as well as its curvature are suggestive of that ancient

1 Trans. American Phil. Soc. xviii. 1896, p. 55.

LU LIBRARY

250 TAPIRS CHAP.

Ungulate Phenacodus (see p. 202). In the next stage (an
embryo of 25 mm.) the humerus has slightly decreased in pro-
portionate length, and has come to be more like that of
Hipparion. In both of these embryos it should be noted that
the ulna is complete and separate from the radius. In the
second of the two it has more distinctly acquired the form which
it will possess in the adult. The second metacarpal — one of the
splint bones of the adult — is tipped with a small nodule of
cartilage, which is clearly the representative of one or more of the
phalanges belonging to that digit.

Fam. 2. Tapiridae. — The Tapirs may be distinguished from
the Horse and from the Ehinoceros tribe by a few characters,
which are as follows : —

The dentition is generally the full one of forty -four teeth.
The premolars in the more ancient forms are unlike the molars,
but like them in more recent forms. The molars of the upper
jaw have two crests parallel and united by an outer crest. The
fore -feet have four, the hind-feet three toes.

The family is fully as ancient as that of the Equidae, but the
specialisation of the toes never advances so far. The modern
representatives of the order are, so far as the feet are concerned,
in the condition of very early representatives of the equine stock.
Nor do the teeth of the Tapir's ever reach the complicated pattern
of that presented by at least the modern Horses, or indeed of the
Palaeotheres. Apart from this it is not an easy matter to dis-
tinguish accurately between these several families, including the
Lophiodontidae, which, as already mentioned, is placed nearer to
the Tapiridae than to the Palaeotheriidae. Indeed the differentia-
tion of these two families, the Tapiridae and the Lophiodontidae,
seems to be a matter of the greatest difficulty. The difficulty is
well emphasised by the fact that naturalists disagree most
profoundly as to the relations of various genera of extinct Tapir-
like animals. For Mr. Lydekker the genus Lophiodon includes
also the American genera Isectolopkus and Systemodon, which are
placed by Zittel in the sub -family Tapirinae as opposed to
Lophiodontinae, which contains Lophiodon and Helaletes. The
existing Tapirs can be differentiated from the existing Horses with
great ease, as the following account of the existing genera will
show.

The genus Tapirus is now met with only in South and

AMERICAN TAPIRS

251

Central America, and in the Malay Peninsula and the islands of
Java and Sumatra. This animal is in many respects the most
ancient of existing forms referable to the Perissodactyle order.
It has four toes on the front-feet, though only three on the hind-
feet. The number of teeth is 42 — nearly the typical Euther-
ian number. The Tapirs are always moderately-sized animals,
entirely covered with hair, and usually of a brownish -black
colour. The Malayan Tapir is, however, banded broadly with
white — a single band ; the young of the Tapir is spotted, and
striped with white. The nose and upper lip conjoined are pro-

FIG. 128. — American Tapir. Tapirus terrestris. x^.

duced into a short trunk, precisely comparable with that of the
Elephant. As in the Ehinoceros — and in this both contrast with
the other existing Perissodactyle genus Equus — the temporal fossa
is not separated from the orbit by bone. Of existing Tapirs
there are at any rate T. terrestris? T. roulini (the " Tapir Pinch-
aque" of Cuvier), T. dowi and T. lairdi in America (the last
two being sometimes separated into a distinct genus, Elasmo-
gnathus, on account of the prolongation of the ossified mesethmoid),
and T. indicus in the East. The tapir, probably T. terrestris, is
described by Buffon as "a dull and gloomy animal." It is
certainly mainly nocturnal in habit. The name terrestris was
given by Linnaeus, who placed it in the same genus as Hippo-

1 T. leucogenys and T. ecuadorensis are probably not distinct, the latter being
in reality T. terrestris, tbe former T. roulini.

252

THE ORIENTAL TAPIR

potamus amphibius ; hence the epithet applied to the Tapir. But
as a matter of fact it loves marshy neighbourhoods, and is in a
way amphibious. This does not of course apply to the Andesian
T. roulini, which inhabits the cordillera of Ecuador and Colombia.
The distribution of existing Tapirs is, as is so often the case,
restricted when compared with that of their extinct congeners and
allies. In Europe the remains of the genus Tapirus are abundant
from Pliocene strata, and its remains are there known from
as far back as the Miocene. The genus is thus one of the very
oldest forms of Mammalia at present inhabiting the earth.

Fio. 129. — Malayan Tapir. Tapirus indicns, young,

(From Nature.)

The Malayan Tapir is to be distinguished from the American
(T. terrestris — the other species have not been dissected) by the
greater development of the valvulae conniventes in the intestine,
the absence of a moderator band in the heart, and the less
elongated caecum, which is sacculated by only three bands, there
being four in T. terrestris.1 The animal frequents the most
retired spots among the hill woods, by which habit it seems

1 See Beddard, Proe. Zool. Soc. 1 889, p. 252, and other papers there cited, for
the anatomy of the Tapir.

x CHARACTERS OF RHINOCEROS 253

largely to escape the Tiger, its most formidable foe in those regions
of the world. Its quickness of senses enables it also to slip away
with rapidity. It can proceed at a great pace when disturbed,
and can readily push its way through obstacles. The young
animal, like that of the American species, is dark brown with
yellowish spots. It is stated by Mr. H. N". Eidley that the
young animal lies during the hot part of the day under bushes,
in which situation " its coat is so exactly like a patch of ground
necked with sunlight that it is quite invisible." It is interesting
to note that here, as with some other animals, it is the young
that are especially protected by such mechanisms. Moreover,
some of the spots are round and some are more elongated, so
that the resemblance to spots of sunlight which come in a direct
and in a slanting direction is greatly increased. Even the
colours of the adult are not so conspicuous when it is in its
native haunts as might be supposed. The breaking up of the
ground colour into tracts of two different colours prevent it from
striking the eye so plainly as if it were of one colour through-
out. " When lying down during the day it exactly resembles a
grey boulder, and as it often lives near the rocky streams of the
hill jungles, it is really nearly as invisible then as it was when it
was speckled."1

Fam. 3. Rhinocerotidae. — This family is to be distinguished
from the preceding by a number of characters, which though not
universal are general. In the first place, there are commonly
horns, or a horn, consisting of what appears to be an agglomera-
tion of hair-like structures fixed upon a roughened patch of bone
on the surface of the nasals. The incisors are diminished or
defective, and the upper canines are often wanting. The molars
and premolars are alike. The fore-feet are four- or three-toed,
but are functionally tridactyle ; the hind -feet are three -toed.
The skeleton in this family is massive, and the limbs relatively
short. The skull, as in the Tapirs, has a confluent orbit and
temporal fossa. The upper lip is generally more or less pre-
hensile ; the body is as a rule — to which the Pleistocene Hairy
Ehinoceros is of course an exception — rather sparsely covered
with hair. In this feature the Ehinocerotidae contrast both with
the Tapiridae and the Equidae. The family in reality contains
but one existing genus, though three have been instituted, viz.

1 Natural Science, vi. 1895, p. 161.

254 VISCERA OF RHINOCEROS CHAP.

Rhinoceros, Ceratorhinus, and Atelodvis. As there are so few
existing species the subdivision of animals which agree in so
many and such highly -characteristic features seems to be an
unnecessary procedure. The existing Rhinoceroses are but a
fragment of the total number of known forms from past epochs.
The family is very markedly on the wane.

The genus Rhinoceros is characterised by its heavy build and
thick, almost smooth, skin — smooth, that is to say, so far as con-
cerns the slight development of hair — which is often thrown into
folds. There is one or, there are two horns on the fore-part of
the head, which are, as has already been pointed out, structures
sui generis, and not exactly comparable with the horns of other
living Ungulates. There are three nearly equal toes on both
fore- and hind-limbs. The canine teeth of existing species have
disappeared ; the incisors are, or are not, present ; the molars and
prernolars are three and four in each half of each jaw.

The visceral anatomy of the Rhinoceros has been much inves-
tigated so far as concerns the Asiatic forms. A curious feature,
which serves to discriminate some of the Asiatic species from
others, is to be seen in the small intestine. In Rh. indicus *
this gut is furnished with numerous long cylindrical narrow out-
growths " like tags of worsted " ; in the allied Rh. sondaicus these
tags are present, but are natter and broader ; while in the two-
horned Rh. sumatrensis there are no tags at all, but only smooth
valve-like folds. Another mark by which these species can be
distinguished depends upon the variation in the presence or
absence of certain glands imbedded in the integument of the foot
— the so-called " hoof glands." These occur in Rh. indicus and
Rh. sondaicus, but are absent in Rh. sumatrensis.

Sir W. Flower2 studied some years since the skull features
which serve to differentiate the existing forms.

In Rh. sumatrensis the two long downward processes of the
squarnosal bone, termed respectively post-glenoid and post-
tympanic, do not unite below the auditory meatus. In this the
species in question agrees with the African forms but not with
the one-horned Asiatic species, where the two processes completely
fuse. Again, another character, though perhaps less important,

1 Garrorl, Proc. Zool. Soc. 1873, p. 92 ; ibid. 1877, p. 707. Beddard and Treves,
Trans. Zool. Soc. xii. 1887, p. 183.

2 Proc. Zool. Soc. 1876, p. 443.

x INDIAN RHINOCEROS 255

is the sloping backwards instead of forward of the occipital crest
in all two-horned species, whether African or Asiatic.

The Asiatic Rhinoceroses have, what the African animals
have not, functional incisor teeth throughout life. It has been
proposed on these and other grounds to separate generically the
African and Asiatic forms.

The Asiatic Rhinoceroses include three well -differentiated
species, in all of which the skin is much thrown into folds.
Rli. indicus is the largest form. It is one horned, and has
enormous folds of skin at the neck and hanging over the limbs.

FIG. 130.— Indian Rhinoceros. Rhinoceros indicus. x^.

So like artificial armour is this thick plating, that Albrecht Diirer
may be excused for having given the beast the appearance of
being actually mail-plated in a sketch which he made of a speci-
men sent over to the King of Portugal in 1513. This particular
beast, one of if not the first sent over to Europe, proved so in-
tractable in disposition that the king sent it as a present to the
Pope. But "in an access of fury it sunk the vessel on its
passage " ! The horn of this and of other species was held until
almost our times to have medicinal and other more curious values.
So recently as 1763 it was gravely asserted that a. cup made of
its horn would fall to pieces if poison were poured into it.
" When wine is poured therein," wrote Dr. Brookes in the year
referred to, " it will rise, ferment, and seem to boil ; but when

256

SONDAIC RHINOCEROS

mixed with poison it cleaves in two, which experiment has been
seen by thousands of people." John Evelyn also wrote of a well
in Italy which was kejpt sweet by a Rhinoceros' horn. This
species seems to be long-lived, even in captivity; a specimen
now to be seen in the Zoological Society's Gardens has been there
since the year 1864.

Rhinoceros sondaicus, the Rhinoceros of the Sunderbunds, has
a much wider range than the last species or Indian Rhinoceros

FIG. 131. — Sumatran Rhinoceros. Rhinoceros sumatrensis. x ^5. (From Nature.)

This is unknown out of India itself, and is there limited to a small
region ; the Sondaic form is found in Bengal and in the Malayan
Islands. It is a smaller species, and the armour has a tesselated
appearance. The female generally, if not always, is hornless.

The Sumatran species, Rhinoceros sumatrensis, is to be dis-
tinguished from the last two by its two horns. It is also covered

x HAIRY-EARED RHINOCEROS 257

by a much thicker coat of hairs, which are sometimes blacker
and sometimes redder. On account of its two horns it has
been proposed to separate it from the other Oriental species
into a distinct genus, Ceratorhinus. The animal has much
the same range as the last species, but extends to Borneo.
A variety of this species with hairy ears, from Assam, has been
separated as a distinct form, under the name of Eli. lasiotis,
by Mr. Sclater. The animal upon which that species was
founded was until quite recently living in the Zoological Society's
Gardens.

There are only two certainly-known species of Ehinoceros in
Africa. These are the White Khinoceros (Eh. simus) and the

FIG. 132. — Hairy-eared Rhinoceros. Rhinoceros lasiotis. x-fa.

Black Ehinoceros (Eh. licornis). The origin of the names is not
easy to understand, since the " white " animal is, if anything, darker
in colour than the Black Khinoceros. It is stated, however, that
in past years the specimens of Eli. simus found in the south-west
of Cape Colony were " paler and whiter in colour than those in
the north-east." At present there are no grounds for distinguish-
ing the species by their colour characters. But they are plainly
distinguishable on other grounds. Ehinoceros simus has a square
upper lip, and in relation to this crops the herbage upon the
ground. Eh. bicornis has a prehensile upper lip projecting beyond
the lower, and in a corresponding fashion feeds principally upon
the branches of shrubs. It has been pointed out by Mr.
VOL. x s

258 AFRICAN RHINOCEROSES < IIAI-.

Coryndon l that the calf of Eh. simus " always runs in front of
the cow, while the calf of Eh. bicornis invariably follows its
mother." Both animals of course have two horns, and upon the
varying proportions of the horns a large number of " species " have
been made in the past. It is stated that the longest horn of the
"White Rhinoceros" known measures 56^ inches; while that of

FIG. 133. — Head of Rhinoceros bicornis.

E. bicornis is shorter, 40 inches being apparently the maximum.
But the animal is smaller.

The possible third African species of Rhinoceros '2 has been
provisionally named after Mr. Holm wood, and is based upon two
horns 41 and 42 inches long, which may be abnormal horns
of Eli. bicornis ; but they are thinner and have a smaller pedicel.

Extinct Rhinocerotidae. — The existing Rhinoceroses are thus
confined to Africa, to certain parts of the continent of Asia, and
to some of the large islands lying to the south of that continent.
But formerly the genus, and allied genera, had a wider range.
As far back as the Miocene we meet with remains of Rhinoceroses
closely allied to existing forms. The more ancient forms have, as
is natural, more ancient characters. Thus in Eh. schleiermacJieri
of the Miocene, canines appear to have been present. The
Miocene Aceratherium, primitive in the absence of horns as its

1 Proc. Zool. Soc. 1894, p. 329. See also Mr. Selous' paper in Proc. Zool. Soc.
1881, p. 275.

'- P. L. Sclater. Proc. Zool. Soc. 1893, p. 514.

HORNLESS EXTINCT FORMS 259

name denotes,1 had also canines and, in one species, six incisors
in the lower jaw. This Aceratherium had, moreover, four toes in
the fore-feet. In the Miocene and later the Ehinoceros existed in
Europe and America. There was even a purely northern form,
the Rli. tichorhinus, which possessed a woolly covering and had
the same range as the Mammoth. This Ehinoceros was two-
horned.

The post-Pliocene and European Elasmotherium was a colossal
rhinocerotine creature. This great beast had two horns and a
body 15 feet long. Its limbs are not known, and as the teeth
are different from those of Rhinoceroses in general, it may not
have belonged to this group at all, though Osborn is inclined
to derive it from Aceratherium, admitting at the same time that
the evidence is " decidedly slender." The teeth in fact are like
those of a Horse in being hypselodont and prismatic in form. As
to the two horns, they were apparently not exactly like those of
typical Ehinoceroses ; there was an enormous horn posteriorly,
supported on a huge boss of bone, and in front of this a roughened
spot suggests a smaller or at least a much more slender horn.

It is important to notice that fossil Ehinoceroses belonging to
the restricted genus Ehinoceros were in Europe invariably two-
horned ; it is only in India, where they still exist, that one-horned
forms are met with in a fossil state.

The Ehinoceroses of America were mostly hornless. Dicera-
therium is an exception ; but in many cases it had two parallel not
successive horns, and these were, to judge from the slight promi-
nences, but feeble in development, and perhaps hardly exactly
comparable with the formidable weapons of the Old- World forms.
Aceratherium tridactylum, with indications of paired horns, may
be ancestral to Diceratherium. The American forms have weak and
slender nasals in correspondence with the absence of horns ; the
sagittal crest is retained in contradistinction to the great flattened
surface of the skull in the horned Ehinoceroses. Aceratherium of
both divisions of the globe probably represents the ancestral group
of the horned and the hornless forms. This being the case it is
highly interesting to note a distinct convergence in the quite

1 Quite recently, however, a species, A. incisivum, preserved at Darmstadt,
has been found by Professor Osborn to possess a slight rugosity upon the frontal
bones, which probably indicates the presence of a rudimentary horn, and the same
author is apparently inclined to place in Aceratherium the horned Teleoceras
(see p. 261).

260

HIPPOPOTAMUS-LIKE FORMS

separate American genera towards the European horned genera.
A genus sometimes united with Aceratherium, but still differing
from it in some points, is Aphelops (Teleoceras).1 This animal is
more nearly approximated to " the modern standard " of Khino-

ceroses than is its possible ancestor Aceratherium. The skeleton
in general is more robust, even surpassing that of modern forms,
and approaching the Hippopotamus. There is a reduction in the
upper incisors, which are limited to two pairs, and the lower molars
1 Osborn, Bull. Amer. Mus. Nat. Hist. x. 1898, p. 51.

ANCIENT RHINOCEROSES

are reduced to five. The lower incisors are only two. The
sagittal crest is less marked ; the fifth digit is reduced to a tiny
nodule representing the metacarpus. It had a small nasal horn.
There are numerous other details of likeness to modern Rhinoceroses
in this creature, which has only community of descent, with them
from the older hornless forms, such as Aceratherium and Caenopus.
In the genus Peraceras the upper incisors are as completely gone
as in the living African Rhinoceroses.

The most ancient rhinocerotine types 1 are the Hyracodonts and
the Amynodonts. They both date from the Eocene, and became
extinct in the succeeding Oligocene. Hyracodon2 (Fig. 134) was
" an agile, light-chested, and rather long-necked " type, resembling a
Horse in build. There were no horns present, but the hoofs were
more like those of the Horses than of the existing Ehinoceroses.
These animals were apparently plain dwellers and defenceless, which
is held to account for their compact hoofs and outward similarity
to a Horse. The genus is Oligocene. The dental formula is
IfCi-Pmf Mf.

It is surmised by Professor Scott that the number of dorso-
lumbar vertebrae was twenty-three or twenty-four. The radius and
ulna are complete and separate bones, but the latter is somewhat
reduced. There are four metacarpal bones, of which, however, the
fifth is much reduced. The animal is only three-fingered. The tibia
and the fibula are distinct, and show no tendencies towards fusion ;
but the fibula is much reduced. There are only three metatarsals
and three toes. Had this line, which is to be regarded as a side
branch of the Rhinoceros stem, not died out, it would probably
have resulted, thinks Professor Scott, in monodactyle — very Horse-
like types. It is later than the next genus to be described,
Hyrachyus, of which it is possibly a descendant. An intermediate
type, Triplopus, appears to bind together Hyracodon and Hy-
rachyus.

In Hyrachyus agrarius the skull is long and narrow, the
facial region being markedly longer than in existing Rhinoceroses.
The mastoid portion of the periotic bone is widely exposed upon
the outer face of the skull, which is, as has been said, not the
case with the existing genus Rhinoceros. The dentition is the
complete Eutherian dentition of forty -four teeth. The upper

1 See Osborn, Mem. American Mus. Nat. Hist. vol. i. pt. iii. 1898.
2 Scott, in Gegenbaur's Festschrift, ii. 1896. p. 351.

HYRACODON

263

molar teeth are strikingly like those of the genus Rhinoceros.
The fore-feet are pentadactyle, but functionally tetradactyle ;

the hind -feet tridactyle. The ulna is less reduced than in
Hyracodon, and the dorso-lumbar vertebrae are twenty-five.

The Amynodonts were short, heavy types, probably marsh -
haunting in habit, and possibly with a proboscis like that of the
Tapir. The orbit is higher than it is in the purely terrestrial

264 TITANOTHERES CHAP, x

Hyracodonts, and it is suggested that when swimming it was
raised above the surface as with the Hippopotamus. " This
feature," observes Professor Osborn, " with the long curved tusks,
undoubtedly used in uprooting, suggests the resemblance between
the habits of these animals and those of the hippopotami." There
were no horns in the Amynodonts. The face is shorter than in
the Hyracodonts, and the mastoid is covered as in recent Ehino-
ceroses. The canines are very strongly developed into tusks, but
the incisors show signs of disappearance. We know of the genera
Amynodon, Metamynodon, and Cadurcotherium. All except the
last, which is European, are American in range.

Fam. 4. Titanotheriidae. — These Oligocene Ungulates, often
attaining to large dimensions, are nearly peculiar, so far as is at
present known, to the North American Continent, and are at least
most abundant in it.1 Many generic names, such as Titanotherium,
Erontotherium, Brontops, Titanops, and Menodus, have been given
to them ; but a recent study of the entire material accessible for
description or already described has led Professor Osborn to the
opinion that there was but a single genus, to which the name
Titanotherium must be applied. Of this genus there are some
thirty well-characterised species, of which the gradual evolution
can be traced from the lowest strata of the White River beds
where their remains occur. An entire skeleton of T. robustum
enables us to understand the osteology of these forms and to
compare them with other Perissodactyles. This animal was more
than 13 feet long, standing some 7 feet 7 inches in height. It
seems to have presented during life the aspect of a Ehinoceros with
perhaps a touch of Elephant. The skull is not unlike that of a
Khinoceros in general dimensions and shape ; but there are a pair
of apparent horn cores anteriorly, which are smaller in the more
ancient forms and acquire a large size, a forward direction with a
divergence of the two in the later forms. A glance at the
accompanying figures of skulls (Fig. 137) of early and later
Titanotheres will exhibit the changes in this particular which the
skulls underwent in the lapse of time occupied by the deposition
of these Oligocene beds. The nasals are short in the later, longer
in the more early species, such as T. heloceras and T. coloradense.
The zygomatic arch projects much, and is " shelf-like " in the
later forms, the skull thus getting an immense breadth, which,

1 Remains of the genus have been met with in the Balkans.

FIG. 137. — Three figures showing the cranial evolution of Titanotherium. Upper figure,
T. trigonoceras ; middle figure, T. datum ; lower figure, T. platyceras. (After Osborn.)

266 HORNS OF TITANOTHERIUM CHAP.

together with the long and divergent horn cores, must have given
to the living animal a most bizarre appearance. It is an interest-
ing fact that this animal, though a Perissodactyle, agrees with the
Artiodactyla in the nineteen dorso-lumbar vertebrae, of which seven-
teen bear ribs.

The genus further agrees with the Artiodactyles in the
structure of the carpus. The toes of the fore -limb are four,
those of the hind-limb three ; but while the hind-limb is un-
doubtedly Perissodactyle in the arrangement of its component
parts, the fore -limb shows a hint of an Artiodactyle mode of
structure. This limb is paraxonic, the axis of the limb passing
between the two middle digits. It may be that this genus
represents more nearly than any other Perissodactyle or Artio-
dactyle the primitive stem from which both have diverged, though,
of course, it is not old enough to be very near to the actual
ancestor. The molar dentition is the typical one ; the incisors
seem to vary as to their presence or absence, and, if present, in
their numbers. In comparing the older with the more recent
forms it is noteworthy that there has been an increase of size
exactly as there has been during the evolution of the Camels and
some other groups of Ungulates. As already mentioned, the size
of the horn cores also increases until it culminates in the extra-
ordinary species, T. platyceras and T. ramosum, in which these are
half as long as the skull, flattened in form, and connected at
their bases by a " web " of bone. Arrived at this amount of
specialisation the genus Titanotherium apparently exhausted its
capacities for modification and ceased to be. The many
generic names may be explained by sexual differences on the one
hand and an incomplete knowledge of connecting links on the
other.1

Palaeosyops is somewhat like a Tapir in build, the skull
especially resembling that of the Tapir. As in Titanotherium
the molar teeth, instead of having an outer wall formed by fused
cusps, have a W-shaped outer wall on one side and two or one
cusps on the opposite side. It is, moreover, an Eocene form, and
in correspondence with its greater age is more primitive in some
points of structure, for example, in the absence of horns and in
the full dental formula. The fore-limbs are four-toed, the hind

1 See especially Osborn and Wortman, Bull. Amer. Mvs. Nat. Hist. vii. 1895,
p. 333, and Osborn, ibid. viii. 1896, p. 157.

x CAMEL-LIKE PERISSODACTYLES 267

three-toed. It w.as intermediate between a Tapir and a Rhinoceros
in size. It has been shown, too, from casts of the interior of the
skull, that the cerebral hemispheres are much less convoluted than
were those of Titanotherium.

Eelated to Palaeosyops is another primitive Titanothere, the
genus Telmatotherium. This is also Eocene, from the Uinta
Basin, the uppermost of Eocene strata. The skull of these
creatures was rather elongated, and not unlike that of a Titano-
there in general aspect. The dentition was complete and the
canines not very large. The horns, which acquire so prodigious a
development in the later Titanotheres, are just recognisable in at
any rate many species of this genus Telmatotherium, the name
being thus by no means an apt one. Better was that proposed by
Dr. Wortman, of Manteoceras or " prophet horned." The horns
are small elevations upon the frontals just at the junction of
these with the nasals, and, indeed, lying partly upon the latter
bones. In T. cornutum the horns are chiefly borne upon the very
long nasals, whose size contrasts with the same bones in the
more highly -developed Titanotherium. It appears to be quite
possible that Titanotherium was evolved from the genus
Telmatotherium.1

SUB-ORDER 9. LITOPTERNA.

Whether the Macraucheniidae should be considered as a
separate group of Ungulata is a matter of dispute. Cope
placed them in a special order of Ungulates which he called
Litopterna. Zittel, on the other hand, regards them as definitely
Perissodactyles. One curious point of resemblance to existing
Horses is shown — that is the presence of a pit in the incisor teeth.
This matter seems to be so important as to need a placing of
these forms in the neighbourhood of the Perissodactyles, even of
the Equidae ; it is so peculiar a character, and apparently so little
related to any obvious similarity in way of life, that it seems to
mark a special affinity. Not so the fact that in Macrauchenia
at any rate the orbit was entirely surrounded by bone as in the
Horse. We find that condition so frequently acquired in many
groups, — a development from an earlier condition where the cavity
for the lodgment of the eye is in continuity with the temporal

1 See Osborn, Bull. Amer. Mus. Nat. Hist. vii. 1895, p. 82.

268 ONE-TOED LITOPTERNA CHAP, x

fossa, that it cannot be regarded as anything more than a mark
of specialisation. It is, in fact, the case that the Macraucheniidae
are in many points specialised, while retaining many primitive
features of structure.

The chief primitive features are : the non-alternating positions
of the wrist- and ankle-bones ; these, of course, interlock in the
Perissodactyles of to-day and in many extinct families. Then
the absence of a diastema in the tooth series, coupled with the
presence in Mavrauchenia of a complete dentition. The small
brain may be referred to the same category. Macrauchenia
must have been a strange -looking animal. It walked upon
three toes on each limb; the skull was Horse -like in general
form, but the nostrils are removed to a point about as far back
as in the Whales or nearly so, the -nasal bones being correspond-
ingly reduced. This it is thought argues a proboscis. The
humerus is particularly compared by Burmeister1 to that of a
Horse. The radius and ulna though both well developed are
fused. The neck is long, and, as in the Camel, the vertebral
arteries run inside the neural arches. Since the fore-legs seem
to have been rather longer than the hind-legs, though only very
slightly, and the neck was long, the animal may have presented
some likeness to the Giraffe. It is interesting to note that in the
proportions of humerus to ulna this animal is more Lama -like
than Horse-like. On the other hand, the proportions of femur
to tibia are more Horse-like. The remains of the creature are
limited to South America, and to quite superficial deposits. It
is evidently a specialised type, and has pursued a course parallel
to that of the Horse. Much nearer to the Horse however, but
apparently by convergence only, is the genus Thoatherium,
usually placed in a separate family, the Protorotheriidae. In this
creature, which has many archaic characters, the toes are reduced
to one in each foot. In an allied form, Protorotherium, we have
the two lateral toes diminishing just as in Anchitherium.

1 N. Ada Acad. Cacs. Leap. Car. xxvii. 1885, p. 238.

CHAPTER XI

UNGULATA (continued} — ARTIODACTYLA (EVEN-TOED
UNGULATES) — SIRENIA

SUB-ORDER 10. AETIODACTYLA.
THE Artiodactyle or "Even -toed" Ungulates are to be dis-

IF 7T[

FIG. 138.— Bones of the Manus — A, of Pig (Sus scrofa). x £. B, of Red Deer (Cervus
elaphus). x \. C, of Camel (Camelus bactrianus). x £. e, Cuneiform ; I, lunar;
m, magnum ; mz, m5, second and fifth metacarpals ; R, radius ; *, scaphoid ; td,
trapezoid ; w, unciform ; U, ulna ; //- V, second to fifth fingers. (From Flower's
Osteology.)

tinguished from the Perissodactyla, and from other Ungulate
groups, by a number of trenchant characters. The most salient

2/0 SOLID-HOOFED PIGS

of these, and that which has given its name to the group,
concerns the arrangement of the digits. Instead of there being
but one prevailing digit — the third, in the hand and foot,
through which the axis of the foot passes, there are two, numbers
three and four, between which the same axis passes, and which
are perfectly symmetrical with each other. This type of foot has
been termed " paraxonic," as opposed to the " mesaxonic " Perisso-
dactyle foot (see Fig. 121 B, p. 235). It has been attempted
to prove that the single prevailing digit of the Horse's foot is a
fused pair of digits, and the state of affairs which characterises
the Camel, where the two metacarpals or metatarsals are to an
almost complete extent united, has been urged in proof; so,
too, certain abnormalities, such as those called " solid-hoofed
pigs." l These latter are simply Pigs in which the two central
metacarpals and the terminal hoofs are completely fused with one
another. In some of such cases there is not the slightest trace of
the union of the separate metacarpals and phalanges. Even the
sesamoid bones, attached behind to the toes, are two in number
instead of four. And, furthermore, the tendon supplying the
bones is single, though showing traces of its double origin.
Such Pigs often show the abnormality from generation to genera-
tion, and they proved convenient for those whose scruples would
not allow them to eat the flesh of a beast " dividing the hoof "
and not chewing the cud. More singular still, as showing a
pathological approach from another side to the Perissodactyle
condition in an Artiodactyle, is a calf, where the foot ended in
three equi-sized digits, of which the middle one lay in the longi-
tudinal axis of the limb. From the opposite side cases are
known of a Horse with a split hoof and phalanges, thus present-
ing the most striking likeness to a Camel.

There is, furthermore, in certain groups of Artiodactyles
(e.g. the Tragulidae) a tendency for the two middle metacarpals to
unite, quite apart from such " sports " as those illustrated by the
cases just set forth. And, as already mentioned, the union of the
two middle metacarpals culminates in the Camel, Ox, etc. There
is, however, absolutely no trace of such a fusion in the series of
Perissodactyle animals known to us ; and it would be by fusion
rather than dismemberment that, as it would appear on this
theory, the modern Ungulate foot has been arrived at. Of course

1 See Bateson, Materials for the Study of Variation, London, 1894, p. 387.

CARPUS AND TARSUS OF ARTIODACTYLES

271

the facts of Ungulate descent are absolutely destructive of any
such comparisons.

As is the case with the Perissodactyles, the Artiodactyles
show a historical series, the primitive five-toed condition being
almost preserved in Oreodon, up to the most modern modification
exemplified by the Ox, Sheep, etc., in which animals there are not
even vestiges of the fourth and fifth toes. It has been stated,
however, that the foetal Sheep has traces of
those rudiments. The so-called cannon bone
(the fused third and fourth metapodia) is
accompanied in its fusion by an increase in
length. At the same time the functional
middle metacarpals push aside the rudiments
and, forming a broad surface for that purpose,
articulate with the magnum and unciform
bones to the exclusion of the rudiments.
This has been termed an " adaptive reduc-
tion." In the " inadaptive reduction " there
is the same reduction of the metacarpals,
but the rudiments still articulate as in the
primitive Artiodactyle foot, i.e. Me II with
trapezium, trapezoid, and magnum ; Me III
with magnum and unciform ; Me IV and V
with unciform. This would appear to give FlG 139._Dors'lisurfaceof
greater solidity and consequently greater right tarsus of Red Deer
strength to the foot.

The carpal bones of the Artiodactyla
alternate in their articulation ; the primi-
tive state of affairs l is not retained even
in the earliest types. The femur has no
third trochanter, so prevalent in the Perissodactyles. In the
hind-foot the calcaneum has an articular facet for the fibula,
which is not characteristic of the Perissodactyla. In the more
modern forms, e.g. the Cervidae, the navicular and cuboid become
fused into one bone ; and there are even further fusions which
will be referred to later as characteristic features of different
groups. It is interesting to notice that the reduction begins
earlier and is clearer in the hind-foot than in the fore. One

(Cervus elaphns).
a, Astragalus ; c, cal-
caneum ; c3, cuneiform ;
cb, cuboid ; mill, mJV,
metatarsals ; ?<, navicu-
lar. (From Flower's
Osteology.)

1 See, however, p. 196, for a discussion as to which is the more primitive
arrangement.

2/2 STOMACH OF ARTIODACTYLES CHAP.

can see how this may be purely adaptive, the push of the hind-
legs in running needing a firmer support. In Hyomoschus this
is the case. The hind-limbs are provided with a cannon bone,
while the metacarpals of the fore-feet are still free.

The number of dorso-lumbar vertebrae is less in the Artio-
dactyle than in the Perissodactyle Ungulates. Whereas the former
have but nineteen, the latter have, as a rule, twenty-three such
vertebrae.1 The number of ribs varies from twelve (Camelus,
Hydropotes) through thirteen (Cervus, Gazella) to fourteen in
Dicotyles, Giraffa, etc.

The curious form of teeth known as " selenodont " is character-
istic of the Artiodactyla, though only found well developed in the
modern forms, and of those only in the Pecora. The more primitive
forms had " bunodont " teeth with typically four tubercles (if we
except the tritubercular and but little-known Pantolestes) ; and the
intermediate " buno-selenodont " type characterises such groups as
the Anthracotheriidae.

While the stomach of the Perissodactyles is always a simple

sac, it is complicated, or shows signs of complication, in the

Artiodactyles. That of the Hippopotamus is divided into two

chambers ; there are three in Tragulus, and four in the typical

. Ruminants such as Oervust Oms, etc.

Had we to deal only with the still living genera of Artio-
dactyles, it would be easy to sort them into two groups on the
characters of the teeth ; for the Pigs and Hippopotamus are pro-
vided with tubercular molars ; they are bunodont. The Deer,
Camels, Oxen, Giraffes, etc., have selenodont molars. Besides, the
latter are " Ruminants," and have a more complicated stomach.
The existing Chevrotains forbid a more trenchant division, since
they are, as will be pointed out in due course, somewhat inter-
mediate in structure ; the feet are more Pig-like, and the stomach
is not so typically Ruminant. In any case such a division is pre-
vented by certain extinct families which are perhaps ancestral to
both. They have teeth which are not quite bunodont and not
quite selenodont. These teeth have been termed buno-selenodont
or buno-lophodont.

The distribution of the living Artiodactyles presents us with
some interesting facts. The vast preponderance of species occurs
in the Old World — 34 in America as against over 250 species

1 Titanotherium (see p. 266) is exceptional.

xi ARTIODACTYLES OF MADAGASCAR 273

in Europe, Asia, and Africa. The Neotropical region has no Oxen,
or Sheep, or Antelopes. The latter are confined to Africa, Asia,
and certain parts of the Palaearctic region ; they are vastly more
prevalent in Africa, where they take the place of the totally
absent Deer. The Pig tribe is almost entirely Oriental and
Ethiopian in distribution, only one form, the European Wild Boar,
ranging into the Palaearctic region; and the two species of
Peccary are found in both North and South America. Broadly
speaking, the Ethiopian region is the headquarters of the Artio-
dactyla. But the great island of Madagascar has but one form of
Artiodactyle, a Pig of the genus Potamochoerus.1

GROUP I.—SUINA.

Fam. 1. Hippopotamidae. — The family Hippopotamidae con-
tains of existing genera only Hippopotamus, for the Liberian dwarf
Hippopotamus is not now regarded, as it was formerly, as the type
of another genus, Choeropsis. The reasons for its former separa-
tion were the loss of the outer pair of incisors and the different
proportions of various parts of the skull. This little Liberian
animal has, however, been shown by Sir W. Flower2 to possess
the missing incisors occasionally; and as to the proportions of
the skull, it is exceedingly common for small animals to vary from
larger relatives in this way. Hence, considering the characteristic
features of the Hippopotamus and the fewness of species, it seems
unnecessary to divide it up further. We shall therefore only
recognise one genus.

The Hippopotamus at present is African in range, and confined
to that continent. But quite recently it inhabited Madagascar ;
and further back still in time the existing African species,
H. amphibius, ranged into Europe ; there were also Indian forms,
which were contemporary with the Stone - age man. The
Common Hippopotamus is a great thick-skinned beast with but
few hairs. It has four toes on each foot, a complex stomach, but
no caecum. The strong incisors continue growing through life, as
.do the great canines. The number of incisors is two on each
side of each jaw. Some of the extinct species had six in each

1 Bones of Hippopotamus, however, indicate the very recent occurrence of that
animal in Madagascar.

2 "On the Pygmy Hippopotamus of Liberia,*' Proc. Zool. Soc. 1887, p. 612.
VOL. X T

274

HIPPOPOTAMUS

jaw, and they were distinguished as a genus Hexaprotodon, con-
trasting with Tetraprotodon, until intermediate conditions were
observed. Choeropsis, as already observed, was a still further
reduction of the tetraprotodont type. The molars (the formula is
Pm f- M |) when worn show a double trefoil pattern. The
orbital cavity is encircled by bone. As with many other aquatic
mammals the kidneys are lobulated.

FIG. 140. — Hippopotamus. Hippopotamus amphibius. x •£,.

A very singular fact about the Hippopotamus is the pro-
duction of a " bloody sweat," a carmine-coloured secretion, con-
taining small crystals and corpuscles, from the skin. This
coloured fluid has of course nothing to do with blood.1

The animal grows to a length of at any rate 14 feet. The
limbs and the tail are short. Like other aquatic animals the
nostrils are on the surface of the head, and can be closed when
the animal is under water. When it reaches the surface of the
water after a prolonged immersion, it spouts like a Whale. Sir
Samuel Baker says that ten minutes is the longest time that the
Hippopotamus can remain below the water. It is frequently a
dangerous animal to encounter, as it will capsize boats, and even
bite large pieces out of their bottoms ; with its huge teeth it

1 Tomes, Proc. Zool. Soc. 1850, p. 160.

ANATOMY OF PIGS

275

can and does attack and destroy human beings. The Hippo-
potamus not only swims, but can walk along the bottom of a
river with great rapidity. It occasionally puts out to sea from
the mouths of rivers frequented by it ; and it is supposed that in
this way Madagascar was populated with Hippopotamuses, whose
remains are now found in swamps in that island.

Fam. 2. Suidae. — The Pig family, Suidae, diner from the last
in their smaller size, in the terminal nostrils and mobile snout,
which is not grooved, except faintly as in Babirusa. They are
generally hairy, but the Babyroussa is an exception, while Phaco-

FlG. 141.— Wild Boar. Sus scrofa. x T'5.

choerus is but slightly haired. Though there are four digits, as
in the Hippopotamus, only two reach the ground in walking.
The stomach, furthermore, is simple, and (except in Dicotyles] there
is a caecum. The kidneys are smooth, and the liver is more
lobate than in Hippopotamus. The orbital cavity is confluent
with the temporal fossa. The typical genus, Sus, is distributed
over Europe, Asia, and the islands of the Malay Archipelago,
reaching as far as Borneo and Celebes. The dentition l is
complete. A single species, the so-called S. sennaariensis, is
from Ethiopian Africa, but it is not certain how far this animal
may be an escaped species introduced by man. A very large
number of " species " of Sus have been described, but Dr. Forsyth

1 There is, however, some doubt about the first premolars.

2/6

THE PYGMY HOG

Major is disposed to reduce them to four if not to fewer species.
He allows the widely - ranging S. scrofa, S. vittatus, and the
eastern Malayan S. verrucosus and S. barbatus.

The Pygmy Hog of the Bhotans seema to be not entitled to
specific rank, certainly not to generic (in the opinion of some),
though it has been termed Porcula salvania} The Wild Boar of

FIG. 142. — Pygmy Hog (from Nature). Sus salvania. x J.

Europe is Sus scrofa. It was formerly quite abundant in this
country ; not merely are its remains exhumed from fens, caves
and peat bogs, but there is ample evidence of its continuance
down to a comparatively late historic period. Enactments are
on record as to the hunting of these animals ; there are places,
such as Boarstall, whose names are clearly derived from the
name of the animal, presumably once a native of the locality ;
and various documents all show the presence of the Wild Boar

1 Dr. Garson has investigated its anatomy, Proc. Zool. Soc. 1883, p. 413, and
states that its differences from Sus are " unimportant and few."

XI

WART HOGS

277

in this country down to so late a period as the end of the
sixteenth century.

The African Wart Hog, genus Phacochoerus, is usually regarded

FIG. 143. — Wart Hog. Phacochoerus aethiopicus. x J.

as the type of a distinct genus of Pigs. This animal, " super-
latively ugly " with its huge tusks and great protuberances upon
the face, is chiefly to be
distinguished from the
genus Sus by these char-
acters, and by the com-
plexity of the last molar,
which, with the tusks,
are sometimes in aged
animals the only teeth
left. The complete for-
mula is Pm|^ Mf . There
are two species of this
genus, P. aethiopicus and
P. africanus. When en-
raged the Wart Hog is said to carry its tail directly up, and to
present a ludicrous as well as ferocious appearance.

The Celebesian Babyroussa, genus Babirusa, is an almost
hairless hog with enormously upturned tusks in both jaws of the

FIG. 144.— Heart of Wart Hog.

2/8

BABYROUSSA

male. In the Wild Boar there is a hint of this, which is carried
still further in Phacochoerits ; but in £nlirusa the upper tusks
turn upwards before they leave the substance of the jaw, for
which reason they appear to arise on its dorsal surface ; the
lower tusks are nearly as long. It has been found that the
young of this Pig are not striped as are those of other Pigs.
By means of the curved upper tusks this animal has been said by
old writers to suspend itself to branches of trees, just as does by
his downwardly-projecting tusks the male Chevrotain ! There is
but one species, B. alfurus.

From Sus proper the African and Malagasy Potamochoerus,
including the Red River Hog, is barely separable generically.
Their principal claim to generic distinction lies in the existence
of a horny outgrowth arising from a bony apophysis above the
canine in the male. These have been compared to the osseous
" horn cores " in the extinct Dinocerata. But the Javau Sus
verrucosus shows at least the beginning of a similar modification.
The popular name of the animal is derived from the fine rufous
colour of its pelage, not seen, however, in all the species. Dr.
Forsyth Major l recognises five species, of which only one is from
Madagascar.

Fam. 3. Dicotylidae. — The Peccaries are generally placed in

FIG. 145. — Peccary. Dicotyles lajcifu. x £.

a different family from that of the other Pigs. This family,
1 " On the Species of Potamochoerus," Proc. Zool. Soc. 1897, p. 359.

xi ERYMANTHINE BOAR 379

Dicotylidae, contains but one genus, Dicotyles, with at most two
species. The name of the animal is connected with the dorsal
gland ; the animal thus appeared to possess two navels. The
Peccaries, exclusively confined to the New World, differ from
the Old- World Pigs in one or more important characters. They
have only three toes on the hind-feet, and the stomach is com-
plicated. Though the Peccaries have but small tusks they hunt
in packs and are very dangerous animals to meet with. They
owe, too, their safety from many foes to their sociable habits.
Being nocturnal animals they are liable to the attacks of the
Jaguar, which will speedily overpower and devour a Peccary that
has strayed from its herd.

Fossil Swine. — The existing genera of the Pig tribe are also
known in a fossil condition. Sus itself goes back as far as the
Upper Miocene. Sus erymanthius, the Erymanthine Boar, is
known from beds of that age in Greece, England, and Germany.
This genus is not known to have had a wider distribution in the
past than it has in the present. Dicotyles occurs in the Pleisto-
cene of both North and South America, the regions which it
inhabits at the present day. The genus Listriodon, also Miocene,
is remarkable for having lophodont instead of bunodont teeth,
that is so far as concerns the molars, which resemble those of
the Tapir. It was European and Indian in range. A number
of genera, more remote from the existing Pigs than those which
have just been dealt with, are placed together in a special
sub-family, Achaenodontinae. The type genus, Achaenodon, had
a somewhat short skull for a Pig ; and it is in general aspect
and in the characters of the canine teeth highly suggestive of
that of a Carnivore. The bunodont molars, however, are Suine,
as is the form of the lower jaw with a rounded angle. This is
an Eocene animal found in Wyoming.

Elotherium1 occurs chiefly in the Miocene of both North
America and Europe ; but E. uintense is Eocene. The orbits
are completely encircled by bone in the more modern forms ; this
is not the case in the last-described genus, with which E. uintense
agrees. The skull is also longer and more Pig-like. The zygo-
matic arch is powerful, with sometimes a large descending process,
such as is found in Diprotodon, more faintly in Kangaroos, and in
Sloths and certain extinct Edentates. The lower jaw has a pair
1 Marsh, Aimr. Journ. Set. xlvii. 1894, p. 407.

280 EOCENE AND MIOCENE PIGS CHAP.

of dependent processes near the symphysis, which suggest pro-
cesses occupying a corresponding position in Dinoceras. The
skull and body are heavy, but the two-toed limbs are slender.
There is a smaller pair of toes behind these. The dentition
is complete, and the canines are not inordinately developed.
The brain is very diminutive. Perhaps E. uintense should be
separated as a distinct genus, Protelotherium.1

Hyotherium (which is regarded as identical with Palaeockoerus)
•has a sharp sagittal crest ; the orbit is nearly but not quite
closed. The canines are not strongly developed. The upper canines
have double fangs as in Triconodon among extinct mammals,
and as in the Hedgehog and other forms among living Mammalia.
The premolars have the cutting and serrated edge of those of some
other Pigs, a feature which gives them a curious resemblance to
the " grinding " teeth of Seals. The molars are tuberculate, and
like those of living Pigs. It is European and Indian in range,
and Miocene.

The genus Choeropotamus has a complete dental formula save
for the loss of a premolar in the lower jaw. Though it has lost
this tooth, it is from an older stratum than some of those
forms which have retained that premolar ; it has been found
in the Upper Eocene of the Isle of Wight and of the neighbour-
hood of Paris.

The American and Miocene Chaenohyus has lost the corre-
sponding teeth of the upper jaw.

ITomacodon2 is a genus consisting of several species, which
has a bunodont and complete dentition. The molars are sex-
tubercular in the upper jaw. H. vagans was of about the size
of a Eabbit, and it appears to have had a curved neck. The
limbs had five digits, as is so generally the case with Eocene
Ungulates. It is known from the Middle Eocene of Wyoming.

GROUP II. — EUMINANTIA.

The Selenodontia or Euminantia form the second division of

existing Artiodactyles. The characters of the teeth, which give

them their name, have already been referred to. They also

differ in that there are never more than a single pair of incisors

1 Osborn, Bull. Amer. Mus. Nat. Hist. vii. 1895, p. 102.

2 Marsh, Amer. Journ. Sci. xlviii. 1894, p. 262.

xi RUMINATION 28l

in the upper jaw, and very usually there are none. As a
general rule the third and fourth metacarpals and metatarsals
become united to form a cannon bone. To this there is but one
exception, the African Hyomosclms. Moreover, the second and
fifth digits are nearly always rudimentary, and may practically
disappear altogether. Here again the Tragulidae are an excep-
tion. The Kuminantia are so-called on account of the fact that
they "ruminate," that is, after the food has been rapidly

FIG. 146. — Stomach of Ruminant opened to show the internal structure, a, Oesophagus ;
b, rumen ; c, reticulum ; d, psalterium ; e, abomasum ; /, duodenum. (After
Flower and Lydekker.)

swallowed, it is forced back up the gullet and more thoroughly
masticated. Associated with this is a complex stomach, which
is divided into several compartments. This stomach has at least
three compartments, as in the Tragulidae ; but it has usually four.
Its characters are illustrated in Fig. 146. The majority of the
Selenodontia possess horns, which are partly formed of solid
protuberances of the frontal bones. In the Giraffe they are
somewhat different.

This group may be divided into — A. TRAGULINA, Chevro-
tains ; B. TYLOPODA, Camels, Lamas ; and C. PECORA, Deer, Ante-
lopes, Oxen, Giraffes, Goats, Sheep.

A. TRAGULINA.

As the Tragulina are undoubtedly the most ancient of the
Selenodontia it will be logical to commence with an account
of them.

282

DEERLETS

Fam. 4. Tragulidae. — This family comprises a number of
small Deer-like animals, which are really in many points more
related to the Pigs than to the true Deer. They are known as
Chevrotains ; and the term " Deerlet," introduced by Professor
Garrod, is certainly appropriate, since they have the aspect of
very small and hornless Deer. If it were not for their Artio-
dactyle feet one might at a glance confuse these creatures with
some Marsupial type. The family is Oriental and West African
in range. The two genera (whose individual peculiarities will

FIG. 147. — Indian Chevrotaiu. Tragulus tneminna. x ^.

be considered later) differ from other Artiodactyles in a number
of rather important characters.

They are absolutely hornless in both sexes. The canines are
present in both jaws, and are especially well developed in the
upper jaw. The dental formula is I § C ^- Pm § M §. In the
skull the tympanic bulla is usually, as in the non-ruminating
Artiodactyles, filled with loose bony tissue. The feet (usually) have
the four toes of the Suina, and are therefore in a more primi-
tive condition than in Deer and Antelopes. But as the middle
metacarpals are fused in Tragulus (though separate in Hyomoschus)
they are a stage further than are the Pigs, in the direction of the
typical Ruminants.

The stomach is comparatively simple, thus offering inter-

THE WATER CHEVROTAIN 283

mediate characters between the Pigs and the Euminants ; there
are but three separate compartments. A highly - interesting
character is afforded by the placenta. This is in the present
family of the diffuse kind, not presenting the separated and
tufted cotyledons of the Ruminant placenta. This we may fairly
assume is a further proof of the less-specialised characters of
this group * as compared with the Euminantia, a view, however,
which is not universally accepted. While the molars have the
selenodont character of other Pecora, the premolars are more
adapted for cutting, with sharp edges.

The genus Tragulus consists of several species (e.g. T. stanleya-
nus, T. napu, etc.), which have been aptly compared in external
appearance to certain Rodents such as the Agoutis. The legs
are delicate and slender, hardly " thicker than an ordinary cedar
pencil." These creatures have got among the Malays a consider-
able reputation for astuteness, embodied in the saying, " Cunning
as a kanchil." The male has tusks, which greatly contributed to
the confusion of this creature with the totally different Musk
Deer, Moschus moschiferus. It is even said to suspend itself by
their aid to the branches of trees, and so avoid danger.

Hyomoschus (or Dorcatherium as it should properly be called)
is West African. Its rich brown colour, with spots and stripes,
is much like that of the Chevrotains, but it has shorter limbs.
The only species is D. aquaticum, which is sometimes called, on
account of its frequenting the banks of streams, the Water
Chevrotain. Remains of this genus occur in Miocene and Pliocene
strata of Europe.

The separate metacarpals, comparatively simple stomach, absence
of horns, diffuse placenta, and spotted pelage are features which
argue the primitive position of these animals among existing
Artiodactyles.

Besides the two existing genera which have just been treated
of, there are a number of extinct genera undoubtedly belonging
to the same group.

Gelocus (Eocene and Oligocene in range) is a European
genus known from France. It differs from the living mem-
bers of the group by the fact that the second and fifth toes
on both hind- and fore-feet are represented, as in certain Deer,

1 For the structure of Tragulus, see Milne-Edwards, Ann. Sci. Xat. (5) ii. 1864,
p. 49.

284 EXTINCT TRAGULIDS CHAP.

by rudiments at the upper and at the lower end only ; they are
deficient in the middle. The middle large metacarpals, though
closely applied, are not fused. The metatarsals, on the other
hand, are, or are not fused, according to the species. A later form is
the genus Leptomeryx from the Miocene of North America. This
genus departs from the typical Traguline structure in more than
one point. The tympanic bulla is hollow instead of being filled
with cancellated bone ; the cuneiform is not fused with the cuboid
and navicular, though the latter are with each other ; the lateral
digits of the hind-feet are rudimentary. The magnum and
trapezoid, however, are fused. In the fore-feet the middle meta-
carpals are separate, and the lateral less perfect metacarpals have
toes. The metatarsals are fused.

Not definitely referable to the Tragulidae, but coming near
to them, are the Protoceratidae. Of this family there is but
one well-known genus, Protoceras,1 from the Miocene of North
America.

The skull is singularly reminiscent of Dinoceras, with which
this quite Artiodactyle genus has, of course, nothing to do. It
merely exemplifies the phenomenon of " parallelism." In general
form it is peculiarly long and low. There are three pairs of bony
protuberances : one, the largest, pair are situated on the maxillae
rising up just behind the implantation of the canine teeth ; the
parietals have a second pair ; and a third much more diminutive
pair of bosses are upon the frontals, near their junction with
the nasals. This description refers to the male ; the female has
only traces of the parietal bosses. These were all possibly tipped
or sheathed with horn or roughened skin. The dentition of this
genus is precisely that of the Tragulidae, i.e. I § C |- Pm -| M |- .
The orbit is completely encircled by bone ; the auditory bulla is
not swollen ; the premaxillae are small.

The nasal cavity is very large and open, the end of the nasal
bones anteriorly being situated at about the middle of the
skull ; this would seem to indicate at least a flexible and long
nose like that of the Saiga Antelope, if not a trunk.

The brain was of good size, and quite well convoluted.

The limbs are constituted on the Traguline plan ; in the fore-
limbs the middle metacarpals are quite free from each other,
and the more diminutive lateral digits are complete. The meta-

1 Marsh, Amcr. Journ. Sci. 1897, p. 16f>.

CAMELS AND LAMAS 285

tarsals are free, but with a tendency to fusion ; the lateral toes
are only represented at the upper extremity. The carpal bones
are separated.

This animal, which was about the size of a Sheep, though
of more delicate proportions, was allied not only to the Tragu-
lidae but to the Girafhdae ; it is impossible to refer it definitely
to either family.

B. TYLOPODA.

Fam. 5. Camelidae. — This small group of Selenodonts in-
cludes only the Camels and Lamas. The limbs are long and
have no traces of the second and fifth toes. The fused meta-
carpals and metatarsals diverge somewhat at their distal ends. In
the upper jaw is a single pair of incisors. The stomach differs
from that of the typical Euminants. The rumen has smooth and
not papillose walls, and from it are developed the " water cells,"
diverticula with narrow mouths provided with a closing sphincter
muscle. The psalterium is reduced to a mere vestige, and so the
stomach has, as in the Tragulina, but three chambers. This, so
far ancient, character in the structure of the Camel tribe is
associated with another, also seen in the more primitive Ungulates,
viz. the diffuse character of the placenta. A very singular
peculiarity of this group is the fact that the blood corpuscles
instead of showing the ordinary mammalian round contours
are elliptical.

The genus Camelus, confined to the Old World, is
made up of two quite distinct species, the Bactrian Camel,
C. bactrianus, with two humps, and the Dromedary, C.
dromedarius, with only one. The former species is Asiatic. It
is a singular fact that neither of the species is known to occur in
a genuinely wild condition. The so-called " wild " Camels appear
to be invariably feral. The two species will interbreed ; and there
is at the Zoological Society's Gardens such a hybrid, which has the
general appearance and shaggy brown hair1 of the Bactrian animal,
but the one hump of the Dromedary. It may be that the Bactrian
Camels of Lob-nor are really wild ; but the desert contains so many
remains of cities destroyed by sand-storms that these reputed wild

1 This is the winter dress. In the summer both camels lose their long rough
hair.

286

BACTRIAN CAMEL

Camels may be the descendants of animals belonging to the in-
habitants of those cities. A strayed herd of Camels has established
itself in a feral state in Spain. Otherwise the genus does not
occur in Europe. The Camels are also represented in the New
World. The genus Lama (Auchenia of many authors) belongs to
this family. These Camels differ from their allies in the Old
World by their smaller size, by the absence of the characteristic
hump, and by the dropping of one premolar, the dental formula

Flu. 148. — Bactrian Camel. Camelus bactrianus. x^.

being otherwise similar. A variety of names, Lama, Alpaca,
Huanaco, Vicuna, have been applied to these animals ; but it
appears that the names are in excess of the number of the species.
Mr. Thomas, who has lately inquired into the matter, will only
allow two, the Huanaco, Lama huanacos, of which there are two
domestic races, the Llama and the Alpaca, and the Vicuna, Lama
vicugna. They are both South American in range. Not only is
there a herd of escaped Camels in Spain, but the Spaniards
attempted to introduce and acclimatise the useful Lama. The first
Lama ever seen in Europe was brought in the year 1558 to

BEZOAR STONES

287

the town of Middelburg in Holland ; it was purchased and pre-
sented to the Emperor of Germany. Gesner gives a curious figure
of it, representing the animal as a comparatively colossal beast
submitting itself to the guidance of a dwarfish man. The habit of
" spitting " of the Laina is well known. Augustin de Zarate and
Buffon speak of the Lama as having no protection save this habit,
which is more than a mere ejection of saliva : the contents of the

FIG. 149. — Lama. Lama huanacos xTV.

stomach are forcibly shot at the object of its annoyance. It can
also kick and bite. In the intestines (as in those of some other
mammals) are found Bezoar stones, or Bezards as they are variously
spelt. These were once valued in medicine, and even so lately as
1847 were, according to Gay, the historian of Chili, in vogue;
these concretions, comparable to the ambergris of the Whales, were
supposed to be an antidote to poison.

Extinct Camels. — The earliest cameloid type is the genus
Protylopus? of which we are acquainted with an imperfect skull

1 Scs Wortman, Bull. Amer. Mia. Nat. Hist. x. 1898, p. 93.

288

PROFESSOR SCOTT ON POEBROTHERIUM

and the greater part of a radius and ulna belonging to one
individual, and most portions of the hind- limbs in other specimens.
The one species, P. petersoni, was about the size of a "jack
rabbit," and is late Eocene (Uinta formation) and American in
range. The teeth of this mammal are the typical forty-four, and
the canines are not pronounced, being incisiform in shape. In
the skull the nasals overhang, as in the genus Poebrotherium. The
orbit is not closed by bone. There is in this ancient Camel a trace
of the supra-orbital notch so characteristic of the Camel tribe.
" The vertebrae resemble those of the modern Lamas closely in
their general proportions." The lumbars have the usually Cameloid
formula of 7. This genus has but two functional toes on the hind-

FIG. 150. — Skull of Poebrotherium wilsoni. I1, I2, I3, Incisors 1-3. x -J-.
(After Wortman.)

feet, the second and fifth being reduced to vestiges. It is interest-
ing to note that the radius and ulna appear to remain distinct,
except in very old animals, in which they come to be co-ossified
in the middle only, thus foreshadowing their complete union in the
next genus, Poebrotherium. The present genus, moreover, as well
as Poebrotherium, was distinctly unguligrade ; it has not acquired
the characteristic phalangigrade mode of progression of the modern
types of Camels.

The American and Oligocene Poebrotherium has been recently
and exhaustively studied by Professor Scott.1 It was considerably
smaller than a Lama. Its neck was long as compared with other
Artiodactyles, but still shorter than that of the Lama. It was a
lightly-built, graceful creature, with apparently some external
likeness to a Lama. It is an important fact to notice that at this
1 "Osteology of Poclrothcrium," Journ. Morpli. v. 1891, p. 1.

xi ANCESTRY OF CAMELS 289

period, and for a long time after, there were no types referable to
the Camelidae in the Old World. Though a Camel in many
features of its organisation, Poebrotherium was " generalised " in
many ways. Thus the rnetacarpals and metatarsals were not
fused to form a cannon bone, and the two lateral digits were
represented by splint rudiments of metacarpals and meta-
tarsals. The dentition was complete. The skull though
distinctly Tylopodan, also shows more generalised characters.
Thus the orbit is not quite, though nearly, completed by bone.
In the Camel it is quite closed. The nasal bones are much
longer, reaching nearly to the end of the snout. The odontoid
process of the axis vertebra is not spout -like as in existing
forms, but cylindrical, though slightly
flattened upon the upper surface. The
scapula is described as being more like
that of the Lama than of the Camel,
though variations occur which approxi-
mate to the Camel. The brain, judging
of course from casts, has those sulci
" which are common to the whole series
of Ungulates, and closely resemble those
of a foetal Sheep."

* FIG. 151. — Anterior surface of

Later in historical sequence than axis of Red Deer. x|. o,
Poebrotherium, and structurally inter- ±^a^;; ^ E

mediate between it and ProtolablS, is men for second spinal nerve.
.1 TIT- /y -L J.T, • TJ- (From Flower's Osteology.)

the Miocene genus Gomphothenum. It

shows an advance in structure upon Poebrotherium, in that the
orbit is completely encircled by bone, though the posterior wall
is thin ; the lower canines instead of being incisiform are curved
back as in later Camels, and separated by a wide diastema from
the preceding and the succeeding teeth.

Later in age than Poebrotherium is Protolabis, a Tylopod in
which the full number of teeth is still retained ; its skull
presents no particular changes from the Poebrotherine type ; the
nasals, however, are somewhat shortened.

Later still in point of time is Procamelus. In this form we
have apparently an ancestral stock, whence both Camels and Lamas
were derived. The upper incisors are as in existing forms, but
the first and second persist for a somewhat longer time. The
skull shows two well-marked types of structure ; in P. occidentalism

VOL. x u

2QO EXTINCT GENERA OF CAMELS CHAP.

there are more points of likeness to the Lama, in P. angustidens
to the Camel. In both, the orbits are completely encircled by
bone. The nasals are much shortened. The odontoid process of
the axis is still more concave than in Poebrotherium, but not spout-
like as in existing forms. This fact shows that the spout-like
character of the Camels' odontoid process is not a point of affinity
to other Artiodactyles — in fact the occurrence of the same form of
odontoid process in Perissodactyles is enough proof of this. We
must come to the conclusion that the form is adaptive in all cases.
If we were not obliged on palaeontological evidence to come to this
conclusion, the structure in question is just one which would be
fastened upon as evidence of genetic affinity ; for it is a resem-
blance in a small though distinctive point of structure having no
obvious relation to utility. The metacarpals and metatarsals
have coalesced to form the cannon bones, though a rudiment of
one metacarpal seems to remain. The genera referred to appear
to be on the direct line of descent of the modern representatives
of the family. But there are other forms which are offshoots of
the main stem. Such are Homocamelus, Escliatia, and Holo-
meniscus. The last two are Pliocene and American ; the teeth
are much reduced.

C. PECORA.

The Pecora are a group which .possess so many characters in
common that it is not an easy task further to subdivide them.

In all there are but two functional digits on the feet, and the
metacarpals and metatarsals of these are fused. There are no
upper incisors, and canines in the upper jaw are not universal,
and generally small. Horns are confined to this group of the
Selenodontia.1 The premolar teeth are of a simpler form than
the molars. The stomach has four chambers, of which two
may be regarded as belonging to its cardiac half and two to
the pyloric. The former are, in the first place, a large paunch
or rumen, followed by a smaller reticulum, so called on account
of the network arrangement of the folds of its lining membrane.
Connected with the latter, and constituting the first part of the
pyloric half of the stomach, is the psalterium or " manyplies," so
called on account of the longitudinal folds, like the leaves of a
1 Unless Protoeeras (see p. 284) was furnished with horns.

CLASSIFICATION OF DEER

291

book, into which its lining membrane is raised. Finally there is
the abomasum, out of which proceeds the small intestine.
Garrod has observed that the chamber of the stomach which
varies most among the Pecora is the psalterium. This chamber
is specially large in Bos, and particularly small in the Antelopes
Nannotragus and Cephalophus. But its variation relates more
especially to the folds of its mucous membrane. These folds are
of varying lengths and have a definite, arrangement There may
be as many as five sets of laminae of regular depths. The most
simple psalterium is that of Cephalophus, where there are only
two sets of laminae of different sizes, a deeper set and a very much
shallower set ; this form is termed by Garrod " duplicate." Most
common is the " quadruplicate " arrangement, with four sets of
laminae of differing depths. In all Pecora the liver is but little
divided by fissures.

Fam. 6. Cervidae. — The Deer tribe is a very extensive one,
and, with the exception of Africa and Australia, world-wide in
distribution.1

The Deer are absolutely distinguished from all other Kuminant
animals by the existence of antlers, which are invariably present
in the male sex, save in the aberrant genera Moschus and Hydro-
potes ; in the Eeindeer alone are antlers present in both sexes.
The general characters of these appendages have been dealt with
on a former page (p. 200), where they are compared to, or rather
contrasted with, the horns of the Bovidae. These antlers, so
characteristic of the Cervidae, are very variously developed
among the members of the family. Thus in Elapkodus the
antlers are very small and entirely unbranched. In the Muiit-
jacs, Cervulus, the antlers are hardly larger, but they have a
small anterior branch arising from near the pedicel, the " brow
tine." In Cariacus antisiensis only one branch, the brow tine,
is present, but it is nearly as long as the main stem of the antler,
the " beam." In Capreolus capraea the beam bears two tines ;
in Cervus sika three ; in C. duvaitceli two of the three tines
present bear secondary branches. There are other complications
(some of which are illustrated in Figs. 152-157) of the. simple
antler which culminate in the complex antlers with their ex-
panded " palms " of the Elk and the Fallow Deer.

1 Sir Victor Brooke, "On the Classification of the Cervidae," Proc. Zool. Soe.
1878, p. 883.

292 ANATOMY OF DEER

Another highly-interesting fact concerning these same antlers
is their gradual increase in complexity of tines and palm from
the Miocene Cervus matheroni to the great Irish Elk of post-
Tertiary times.

Beyond the antlers there seems to be no character of universal
applicability which distinguishes the Cervidae from the nearly-
related Antelopes. There are, however, a number of structural
features which are nearly universally characteristic. Excepting
Moschus (which Professor Garrod would not allow to be a " Deer "),
no Cervine has a gall-bladder1 to its liver. All Bovidae (in-
cluding Antelopes) have, with the exception of Cephalophus.

A small but constant character of the Deer is the existence of
two orifices to the lachrymal duct. The genus Tragelaphus alone
among Antelopes shows this character.

So far as is known the placenta of the Deer has but few
cotyledons, that of the Bovidae many. But not many types are
known.

The navicular, cuboid and ectocuneiform are often united.
This is never the case in the Bovidae.

The first and second phalanges of the lateral (imperfectly
developed) digits are always present except in the Muntjacs ;
they are never found in Bovidae. The Deer always present a
light brown to a darker brown coloration. Elaphodus michianus
is almost black. There is commonly white on the under parts
and beneath the short tail. Some Deer, such as the Fallow Deer,
are spotted ; and the young of others that are uniformly coloured
when adult are spotted. In some cases a winter coat, darker
than the summer coat, is developed.

Altogether some sixty species of Deer are known, of which
the preponderance are Old- World forms. The Deer of the Old
"World are distributed among the genera2 Cervus (all Europe and
Asia) ; Cervulus, the Muntjacs (India, Burmah, China, etc.) ;
Hydropotes (Eastern China) ; Capreolus (Europe and Central
Asia) ; Elaphodus (Eastern China) ; there is one American
Cervus, the "Wapiti. The American genera are Cariacus and
Pudua. The Elk (Aloes') and the Reindeer (Rangifer) are circum-
polar. The principal structural modification which occurs within

1 It has been occasionally recorded in an Axis Deer, and in another species,
Cariacus superciliaris.

2 It is not every one that admits so many genera. I follow Sir Victor Brooke.

PERE DAVID'S DEER 293

the family Cervidae concerns the rudimentary fifth and second
toes. In Capreolus, Hydropotes, Moschus, Alces, Rangifer, and
Pudua there are considerable remains of the lower parts of
metacarpals II. and V. ; in the other genera smaller traces of
the upper ends of the same bones.

The two most abnormal genera are Moschus and Hydropotes,
more particularly the former, which neither Sir V. Brooke nor
Professor Garrod allow to be members of the family at all. Moschus
is usually placed in a special sub-family by itself, Moschinae, the
remaining Deer being referred to another sub-family, Cervinae.

Sub-Fam. 1. Cervinae. — The genus Cervus comprises rather
over twenty existing species, which, except the Wapiti (C.
canadensis), are exclusively Old World in distribution. The prin-
cipal features of variation in the genus, in accordance with which
it has been divided up into sub-genera, are (1) palmated (Fallow
Deer, Dama) or non- palmated antlers; (2) adults spotted with
white at all ages and seasons (Axis), or in summer only (Pseudaxis),
or not at all ; (3) spotted or unspotted young ; (4) existence or
absence of rudimentary canines in the upper jaw.

Among the members of this genus, Cervus (Elaphurus)
davidianus is interesting as having been first observed by the
missionary Pere David in a park belonging to the Emperor of
China near Pekin. Its horns are remarkable for dividing early
into two branches of equal length, of which the anterior again
branches into two. Specimens of this Deer were ultimately
obtained for the Zoological Society's Gardens.

The species of Cervus are fairly distributed between the Palae-
arctic and the Indian regions. The Palaearctic species, such as
Luhdorff's Deer (Fig. 152), are mainly Asiatic. Cervus elaphus
and Cervus dama alone are European and British. The former
of course is the Eed Deer, the latter the Fallow Deer. The
Eed Deer is reddish -brown in summer and greyish -brown in
winter, with the white patch on the rump so common in the
Deer tribe. The Ked Deer is genuinely wild in Scotland, in
certain parts of Devonshire and Westmoreland, and in the New
Forest. At the beginning of the last century, according to
Gilbert White, there were 500 head of deer in Wolmer Forest,
which were inspected by Queen Anne. The antlers may have as
many as forty-eight points ; and a stag with more than the three
anterior tines is termed a " Ptoyal Hart." The Fallow Deer has

294

LUHDORFF'S DEER

palmated antlers, and is usually spotted. It seems to be an
introduced species, common report pointing to the Eomans as
the introducers. It would be more correct to say " re-introduced,"
for fossil remains of this Deer have been met with.

FIG. 152. — Liihdorff's Deer. Cervus luehdorffi

Ulaphodus1 contains probably two species, E. cephaloplius of
Milne-Edwards and E. micliianus of Swinhoe, both from China.
The antlers are small and unbranched ; the canines in the male
are massive ; it differs from Cervulus, to which it is closely
allied, principally in the absence of frontal glands. The second

1 Garrod, "On the Chinese Deer named Lophotragus michianus by Mr.
Swinhoe," Proc. Zool. Soc. 1876, p. 757.

THE MUNTJACS

295

species has a dark iron-grey pelage, and the late Mr. Consul
Swinhoe described it as very Goat- like in aspect.

Capreolus. — The Roe Deer has fairly complex antlers. It is
a small Deer and has spotted young. The common Roe Deer,
C. capraea, is a native of this country. It is the smallest of our
Deer, and its antlers only have three tines in stags of the third
year. It is a singular fact about this Deer that though the pair-
ing season is in July and August, the young are not born until
the following May or June, a period which does not represent
that of gestation. The germ remains dormant for some time
before developing.

The Muntjacs, Cervulus, form a distinct generic type confined
to the Indian and the South-Eastern Palaearctic region. They

FIG. 153.— Mule Deer. Cariacus macrotis. x fe. (From Nature.)

are small Deer with spotted young, and short one-branched antlers
placed upon pedicels as long as themselves. The canines are
strongly developed in the males. There are about half-a-dozen
species.

Cariacus is exclusively American in range, and contains about
twenty species. There are or are not upper canines. The young

296

SIMPLE ANTLERS

are spotted. The antlers are occasionally very simple ; in C. rnfus
and a few allies (placed in a special sub-genus Coassus) they are
simple spikes without branches. In this genus, and in the nearly
allied and also New- World Pvdua, the vomer is prolonged back-
wards and divides the posterior nares into two. The bulk of the
species are South American.

FIG. 154. — Chilian Deer. Cariacus chilensis. xTV. (From Nature.)

Pudua, just mentioned, comes from the Chilian Andes. It is
a small Deer without canines and with minute antlers. Other
generic names have been proposed for various species of American
deer.

Hydropotes inermis is a small perfectly hornless Deer, living
on the islands of the Yang-tse-kiang. The male has tusks ; the
young are spotted. Though, like other deer, Hydropotes has no
gall-bladder, both Mr. Garrod a and Mr. Forbes 2 found the rudi-

1 Proc. Zool. Soe. 1877, p. 789.

- Proc. Zool. Soc. 1882, p. 636.

XI

ABSENCE OF ANTLERS

297

ments of one in the shape of a white ligamentous cord. Mr.
Forbes has especially dwelt upon the likeness of the brain to that
of Capreolus. The female has four teats, and produces three to
six young at a time. ^.V^

<£$ M?CS -^pb ;

i F ,,.- y ,n

•i

FIG. 155. — Water Deer. Hydropotes inennis. x r\. (From Mature.)

Alces machlis, the Elk or Moose, is a circumpolar species with
palmated antlers and is of large size. The young are unspotted.
This animal is the largest of the Deer tribe. The aspect of
this creature is by no means that of a Deer, the long, thick,
and rather prehensile upper lip not by any means suggesting
the family to which it belongs ; the legs, too, are ungainly
through their unusual length. The Moose has a curious method
of protecting himself from Wolves. Instead of moving about
during heavy snowstorms, and being thus on the heavy ground
an easy prey for these agile enemies, the animal forms what is
known as a " Moose yard." An area of ground is kept well

298

COMPLEX ANTLERS

CHAP.

trampled down, and the animal contents itself with browsing
upon the adjacent stems. The well -trampled ground gives an
easy footing, and by his powerful horns the great stag is able to
keep his enemies at bay.

FIG. 156. — Moose. A Ices machlis. x •£$.

Rangifer tarandus, the Eeindeer, is unique among Deer by
reason of the fact that both sexes wear antlers. These antlers
are palmated. The brow tine and the next or bez tine are also
palmated and are directed forwards and a little downwards. The
young are unspotted. The pelage alters in winter. Like the
Moose, the Eeindeer is circumpolar. As is well known, during the
Pleistocene period the Eeindeer extended its range as far as the
South of France. Even in the historic period it is said to have
been hunted in Caithness.

Reindeer, like so many other particularly Arctic animals,
have regular migrations. In Spitzbergeu, for instance, the animal
migrates in the summer to the inland region of the island, and in

REINDEER AND MUSK DEER

299

the autumn back again to the sea coast to browse upon the sea-
weed. These migrating herds have been stated to be led by a
large female.

FIG. 157. — Reindeer. Rangifer tarandus. x T'7).

Sub-Fam. 2. Moschinae. — Moschus moschifcrus l is a native
of the Asiatic Highlands. It is 3 feet or so high, perfectly
hornless, and with very large canines in the male. It is note-
worthy that in Hydropotes, where the canines are also very large,
horns are absent. These are examples, perhaps, of correlation.
The musk sac (whence the name) is present on the abdomen of
the male only. There is no crumen or suborbital gland, which is
so generally (though by no means universally) present in Cervidae.
But the male has, in addition to the musk glands, glands near
the tail and on the outside of the thigh. Unlike other Deer,
the lachrymal bone of Moschus bears but one orifice. The
feet, so far as concerns the preservation of the outer rudimentary

1 Sir W. Flower "On the Structure and Affinities of the Musk Deer (Moschus
moschifcrus)" Proc. Zool. Soc. 1875, p. 159; Garrod, loc. cit. 1877, p. 287; and
F. Jeffrey Bell, Proc. Zool. Soc. 1876, p. 182.

300

metacarpals, are of the more ancient type represented in Alces,
Hydropotes, etc. A gall-bladder is present. The young, as
in so many Cervidae are spotted ; but the adult is of a greyish-
brown colour.

FIG. 158. — Musk Deer. Moschus moschifems. xj. (From Nature.)

There is no doubt that Moschus is more nearly related to
the Cervidae than to any other Ruminants. It is regarded
by Sir W. Flower as " an undeveloped deer — an animal which
in most points (absence of horns, smooth brain, retention of
gall-bladder, etc.) has ceased to progress with the rest of the
group, while in some few (musk gland, mobile feet) it has taken
a special line of advance of its own."

The musk itself, which gives its name to the creature, is
found in a gland on the belly, about the size of a hen's egg.
The whole gland is cut out and sold in this condition. Such
quantities of musk deer have been and are killed for this purpose
that the rarity of the animal is increasing. In the seventeenth

MUSK PODS

301

century it was so common that the traveller Tavernier purchased
7673 musk "pods" in one journey, or, according to Buffon,
1663. The tusks, which recall those of Hydropotes, to which
Moschus is not nearly allied, and of Tragulus, with which
it has of course still less connexion, are said to be used for the
digging up of roots. Its feet, in relation to its mountain-ranging
habits, are very mobile.

Extinct Species of Deer. — It has been already mentioned
that the most primitive kinds of Deer had no horns at all,
resembling in this the modern Moschus and Hydropotes, and that
with lapse of time went hand in hand an increasing complexity
of antler ; the facts of palaeontology harmonising in the most
striking manner with the facts of individual development from
year to year. The oldest forms seem to be more nearly akin to the
living Muntjacs, and their remains occur in the lowest Miocene
beds of both Europe and America. At present the group is
confined to the warmer parts of Asia and some of the islands
belonging to that continent.

One of the oldest types is Amphitragulus. This genus,
which consists of several species, inhabited Europe, and differed
from living Muntjacs in being totally hornless in both sexes ;
the skull had no lachrymal fossa or deficient lateral ossification.

Nearly allied is Dremotherium of similar age and range.

The Middle Miocene has furnished the remains of the genus
Dicroceras. This is the earliest Deer in which horns have been
found. The horns are, as the name of the genus implies, bifid,
and have, like those of the living Muntjac, a very long pedicel.
This is also a European genus like the last. From this period
we come across true Deer, which commence in the Upper Miocene
and have branched horns. Moreover they belong, at least for
the most part, to the existing genera. One of the most remark-
able forms is Cervus sedgwicki (sometimes placed in a separate
genus, Polycladus) from the Forest Bed of Norfolk and from the
Upper Pliocene of the Val d'Arno. This creature was remark-
able for its multitudinously-branched antlers. These end in no
less than twelve points. No Deer exists or has existed in which
the horns are so completely branched.- They are like those of a
Eed Deer exaggerated.

Fam. 7. Giraffidae. — Undoubtedly the type of a distinct
family, Giraffidae, is the genus Gira/a. It is characterised by

302

GIRAFFE

the long neck, which, nevertheless, consists of only the normal
seven vertebrae, and by the " horns " which differ from those of all
other Kuminants ; they are small bony prominences of the frontal

FIG. 159. — Giraffe. Giraffa camelopardalis. x^0.

bones, which become fused with the skull, and which are covered
with unmodified skin. They are not shed. Between them is a
median prominence. This cranial armature is present in the
female as well as in the male, and is well developed even in the

" SPECIES " OF GIRAFFE 303

new-born young. The orbits are completely encircled by bone,
and there is no lachrymal fossa, so common in Deer and Antelopes.
There are no canines above ; but these are present in the lower
jaw. The rudimentary digits of other Ruminants have dis-
appeared in this genus. There are fourteen pairs of ribs as in
many other Artiodactyla. The liver of the Giraffe l is, as in
many, but not all, Ruminants, devoid of a gall-bladder ; neither
has it a caudate or a Spigelian lobe. The caecum is actually
largish (2|- feet in length), but is relatively very small, as the
small and the large intestines measure 196 and 75 feet in
length respectively. The Giraffe has a well-marked " ileo-
caecal " gland, found in many Ruminants ; its appearance in
Giraffa is especially compared by Garrod with its appearance in
Alces.

Considered by itself, Giraffa forms a very isolated type of
Ruminant. But after we have dealt with certain facts con-
cerning extinct forms clearly allied to Giraffa, the isolation of
the family will be found to be less marked.

The Giraffe (" one who walks swiftly," the word means in
Arabic) is, as every one knows, limited in its range to the African
continent. It is not, however, so familiar a fact that there are
two quite distinct species of Giraffe, one a northern form from
Somaliland, and the other South African. The distinctness of
these two, G. camelopardalis and G. australis, has been lately
worked out in some detail by Mr. de Winton.a The principal
point of difference between them consists in the large size of the
median horn in the Cape species, which is represented by the
merest excrescence in the other species. The Giraffe of West
Africa is held to differ from the northern and southern species,
coming nearer to the former. It appears in the first place to be
a larger animal, and slight differences in the skull have been
pointed out. This series of peculiarities may be expressed, for
those who do not object to trinomial nomenclature, by calling
this novel western form Giraffa camelopardalis peralta. The
existence of the three horns covered with unaltered skin is the
main characteristic of this Ungulate. But the Giraffe also differs
from other Artiodactyles by its enormously long neck, which
enables it to browse upon trees inaccessible to the common herd

1 For the viscera, see Garrod, Proc. Zool. Soc. 1877, p. 5, etc. ; and ibid. p.
289, etc. * Proc. Zool. Soc. 1897, p. 273.

304 INVISIBILITY OF GIRAFFE CHAP.

of Kuminants. The neck is often supposed to have some relation
to this method of feeding. But a more ingenious explanation of
its inordinate length is that it serves as a watch-tower. The
long grass of the districts inhabited by the animal swarms with
Lions and Leopards, which must be foes. The long neck allows
of a wide look out being kept, and it is noteworthy that the
Ostrich, living under similar conditions, is also renowned for its
length of neck. It is the spots upon the Giraffe which have
given it its name of Canieleopard ; these spots present in the
southern form a series of chocolate-coloured areas, sharply marked
off by white spaces. Of these spots it is asserted that they
serve as a means of concealing their possessor. Sir Samuel
Baker l wrote of it in the following words : " The red- barked
mimosa, which is its favourite food, seldom grows higher
than 14 or 15 feet. Many woods are almost entirely composed
of these trees, upon the flat heads of which the giraffe can
feed when looking downwards. I have frequently been mis-
taken when remarking some particular dead tree -stem at a
distance that appeared like a decayed relic of the forest, until
upon nearer approach I have been struck by the peculiar inclina-
tion of the trunk ; suddenly it has started into movement and
disappeared."

The Giraffe, remarked Pliny, " is as quiet as a sheep." The
Eoman public, to whom the first Giraffe ever brought into Europe
was exhibited, expected from its name " to find in it a combina-
tion of the size of the camel and the ferocity of a panther." As
a matter of fact, Giraffes in captivity are not always sheep-like in
temper. They will kick with viciousness and vigour, and will
even initiate an attack upon their keeper. At the same time
they are singularly nervous creatures, and have been known to
die from a shock. In moving, the Giraffe uses the fore- and hind-
limb of each side simultaneously ; this gives to its gait a peculiar
rocking motion, the singularity of which is heightened by the
curving movements of the long neck, which even describes now
and then a figure of eight in the air. Gira/a camelopardalis
and the species (?) already referred to are the only existing
Giraffes (of the genus Giraffa), and they are not found out of
Africa. Sir Harry Johnston has lately given a brief account
of a larger and more brilliantly coloured species from Uganda

1 Wild Beasts and their Ways, 1890, p. 151.

xi THE OKAPI 305

which will probably prove to belong to a distinct genus. It has
five horns, the additional pair being placed above the ears.

Sir Harry Johnston has quite recently made known another
genus of Giraffidae living in the Semliki forest, Belgian Congo
district. The skin and two skulls, as well as the bones of the
feet, are known from specimens sent by Sir Harry Johnston to
the Natural History Museum, and briefly described to the Zoo-
logical Society by Professor Eay Lankester.1 This creature, of
which the native name is " Okapi," is proposed to be called
Ocapia johnstoni. The first actual specimens which reached this
country were two bandoliers made from the skin of the flanks,
which were striped black and white, and were not unnaturally
held to be portions of the skin of a new species of Zebra. The
animal is of about the size of a Sable Antelope, and the back
and sides are of a rich brown colour ; it is only the fore- and
hind-limbs which are striped, the striping being longitudinal, i.e.
parallel with the long axis of the body. The head is Giraffe-
like, but there are no external horns ; wisps of curled hairs
seem to represent the vestiges of the horns of other Giraffes.
The tail is rather short, and the neck is rather thick and short.
The skull is clearly Giraffine. The basicranial axis is straight,
and the fontanelle in the lachrymal region is very large. Upon
the frontal bones near their parietal border is a large boss on
either side, which presumably represents the horn core or " os
cornu." On the mandible the great length of the diastema
between the incisors and premolars is a Giraffine characteristic.
The Okapi lives in pairs in the deepest recesses of the forest.

We are acquainted with a few extinct forms, belonging to
Giraffa, which are extra- African in range. G. sivalensis is from
the Pliocene of the Siwalik Hills in India, G. attica from Greece.
These remains, however, do not include the top of the skull, so that
it is doubtful whether their horns were as in G. camelopardalis.

A closely -allied genus is the extinct Samotherium. This
flourished in Miocene times, and its remains have been found in
the Greek island of Samos. The neck and limbs are shorter than
in the Giraffe, and the horns, longer than in Giraffa, are placed
just above the orbit upon the frontal bones alone, instead of
upon the boundary line of frontals and parietals as in Giraffa. In
several ways, therefore, the existing Giraffe is a more modified or

1 See also Sclater, Proc. Zool. Soc., 1901, ii. p. 3.
VOL. X x

306 EXTINCT GIRAFFES

specialised animal than its forerunner of the Miocene. In the
latter, the male alone carried horns, and in neither sex does
the unpaired median bony excrescence appear. The remains of
this genus (probably even the same species, S. boissieri) also occur
in Persia.

Helladotherium (there is but one species, H. duvernoyi} has
its four limbs of nearly the same length ; the skull of the only
known example is hornless ; the neck is shorter than in Giraffa.
It is known from the Miocene deposits of Pikermi in Greece.

Palaeotragus is a genus which is not referred to the Giraffidae
by all systematists. Its very name, given to it by the eminent
French palaeontologist M. Gaudry, indicates his opinion as to its
Antelopine affinities. The chief and indeed (according to Forsyth
Major a) the only reason for placing this Euminant with the Ante-
lopes is the large size of the horns. They undoubtedly suggest
the horn cores of Antelopes. But they are placed wider apart than
in those animals. It is thought that the hornless Camelopardalis
parva is the female of this species, which is from Pikermi.

Bather more different from Giraffa is the extinct genus
Sivatherium, from the Siwalik deposits of India. Here again
there has been some discussion as to its affinities. Some place
it in the neighbourhood of Antilocapra, but most palaeontologists
now regard it as a Giraffe. The main peculiarity of this large
beast was the existence of two pairs of horn cores ; the larger are
upon the parietal bones, and are of a palmated form, with a few
short tines, which are highly suggestive of those of the Elk
(Alces). The shorter anterior pair are upon the frontal bones.
The neck is short, the limbs of equal length, and there are no
additional toes upon the limbs. Sivatherium was almost as large
as an Elephant, and in restorations it is depicted as having a
fleshy dilated nose like the Saiga Antelope ; this view is based
upon the position and size of the nasal bones. Hornless skulls
have been identified as the female of Sivatherium.

Vishnutherium, Hydraspotherium, and JBramatherium are
allied genera.

Fam. 8. Antilocapridae. — This family contains but one genus

and species, the N. American " Pronghorn," Antilocapra americana.

This animal deserves a family to itself on account of the singular

structure of the horns, which are intermediate in character

1 Forsyth Major, Proc. Zool. Soc. 1891, p. 315.

THE PRONGHORN 307

between those of the Deer and those of the Antelopes. They are
unquestionably " hollow-horned " Ruminants, in that there is an
osseous horn core, upon which lies the actual horn. This, how-
ever, is softer than in Bovidae, and is seniicorneous. It is, indeed,
more like the velvet of the stag's horn. Moreover the horn is
branched, and there are sometimes even three prongs. Further-
more, it is now certainly known that the Pronghorn sheds its
horns not merely occasionally, but with definite annual periodicity.
It so far resembles the Deer. But it must be borne in mind that
in the Deer the horn shedding is a twofold process. There is first
of all the stripping off of the velvet, and secondly the shedding
of a portion of the horn core down to the burr. What happens
in the Prongbuck is the shedding of the true horn only ( = the
shedding of the velvet), not of the horn core. It appears, how-
ever, that occasionally (once in their lifetime ?) certain undoubted
Antelopes may cast their horns.1 Another external character of
this animal is the total absence of " false hoofs," the last vestiges
of the second and fifth digits. The Pronghorn is a gregarious
creature running in bands of six up to hundreds.

Fam. 9. Bovidae. — This family, more extensive than that of
the Cervidae, contains not only the Oxen, Sheep, and Goats, but
also the Antelopes, save only Antilocapra, which must be placed
in a family by itself. The only two points which distinguish
all Bovidae from all Cervidae 2 are the nature of the horns
already described, and the polycotyledonary condition of the
placenta. Moreover the horns are usually present in both sexes,
though there are exceptions, such as the Sheep and Goats, and
various genera of Antelopes (Tragelaphus, Tetraceros, etc.). There
are never the first two phalanges belonging to the rudimentary
digits II., V., as there are in all Deer excepting Cervulus. There
is as a rule but one orifice to the lachrymal duct. There are
never persistent upper canine teeth in either sex.

It is exceedingly difficult to separate the Antelopes from the
Sheep, Oxen, and Goats. Their inclusion along with these creatures
in one family, Bovidae, shows that no differences of an important
character exist. The term Antelope is rather of popular than

1 "On the Shedding of the Horns in the Prongbuck," see Bartlett, Proc. Zool.
Soc. 1865, p. 718 ; Canfield, ibid. 1866, p. 105 ; Murie, ibid. 1870, p. 334 ; and
Forbes, ibid. 1880, p. 540.

2 The distinction between the two families has been called " fanciful." It may
be admitted that it is not great.

308 ANTELOPES AND OXEN CHAP.

of zoological significance. As a rule there are horns in both sexes ;
but this rule is not without exceptions, of which one is the genus
Strepsiceros, the Koodoo. Many other Bovidae are horned in
the males only, e.g. Saiga, Tragelaphus. The Antelopes further
differ from the true Oxen in their more graceful build, and
in the fact that the horns, if they curve at all, generally curve
backwards towards the neck. In the Oxen, on the other hand,
the build is stouter, and the horns usually curve outwards. The
same remarks apply to the Sheep. Such an Antelope, however,
as the Eland (Orias) is very Ox-like in habit. Another feature
which may be remarked upon, though not of absolute differential
value, is that while the Antelopes are as a rule smooth and sleek
in their skins, the Oxen tend to be rough and shaggy. The Zebu,
however, in this, in its hump, and in general aspect, is far from
being unlike an Eland. But then the Zebu is a domestic race, and
we do not know what the wild stock was like. It is perhaps with
the Goats that the Antelopes have the nearest affinities, and it is
difficult to place such a form as Nemorrhaedus, and indeed some
others. In the Antelopes as a rule the middle lower incisors are
larger than the lateral ones ; in the Sheep and Goats they are alike
in size. The parietal bones, too, in the Antelopes are moderately
large and are much shortened in the remaining Cavicornia, especi-
ally in the Oxen. As the Antelopes are the oldest, so far as we
know, of all bovine animals, one would expect to find them com-
bining the characters of the rest. But they do this so effectually
that a disentanglement is really impossible. They date from the
Miocene. Antelopes are now limited to Europe, Asia, and Africa ;
they have always had the same range, though more abundant in
former times in Europe. They preponderate now in tropical
Africa, and abound in genera and species. Messrs. Sclater and
Thomas * allow altogether thirty-five genera, of which twenty-four
are exclusively Ethiopian in range.

In the following summary of the group Messrs. Sclater and
Thomas's work is followed. They commence with a section or
sub-family of which the type is the Hartebeest.

Bubalis, or Alcelaphus as it is sometimes called, is an African
genus, ranging however into Arabia. These Antelopes are
characterised by the long skull and the doubly -curved horns.
There are eight species of the genus, of which B. caama is the

1 The Book of Antelopes, London, Porter, 1894-1900.

GNUS

309

best known ; this is the animal known as the Hartebeest. The
Bontebok and Blessbok belong to a closely-allied genus, Damaliscus,
distinguished mainly by the fact that the bony base of the horn
cores is not extended upwards, and therefore the parietal bones
are visible when the skull is viewed from in front, which is not
the case in Bubalis.

The Gnus, Connochaetes, are familiar owing to their curious
aspect. The hairy face, and rump and tail like those of a pony

FIG. 160. — Brindled Gnu. Connochaetes taurinus. x^0.

are highly characteristic. The horns are bovine in appearance,
standing outwards and then curving upwards.1 There are three
species of Gnu, all from South Africa. They are C. gnu, C. taurinus,
and (7. albogulatus.

Of the Cephalophine section there are two genera :—
Ceplialophus is an African genus. These animals are known
as Duikerboks ; they are small, and have short non- curved
horns in the male sex only. Their general aspect is not un-
like that of certain Deer with simple horns, such as Cervulvs.
Messrs. Sclater and Thomas allow thirty -eight species. The

1 They are straight in the young.

3 I o WATERBUCKS AND KEEDBUCKS CHAP.

smallest species do not exceed the dimensions of a Hare. None
are really large.

Tetraceros is an Indian genus characterised, as its name
denotes, by the fact that it possesses four horns. It is the
posterior pair which correspond to the single pair of Cepha-
lophus. The anterior pair, which are much smaller and are
sometimes absent, are a new pair. The female of this Antelope is
hornless. Sheep are occasionally four-horned, and there is indeed
a breed of such in Kashmir. A four -horned Chamois was
described by the late Mr. Alston.

The Klipspringer, Oreotragus saltator, is the first type of a
third section ; as its name denotes, it is an Antelope with Goat-
like habits, being found particularly among rocks. The horns
are short and straight. This, the only species of the genus, is
African in range, of which its Dutch name gives evidence.
A specimen in the Zoological Society's Gardens (as has been
pointed out to me by Mr. Mercer) had the habit of depositing the
secretion of the tear gland upon a mass of concrete in its
enclosure, the secretion thus exuded forming a pointed heap of
hardish matter. It may be that the object of this is to guide its
fellows to its whereabouts.

Ourebia is a less-known genus, larger in size, but with horns
of the same character, though longer.

The Grysbok and the Steinbok, genus Rapliiceros, have similar
horns. This as well as the last two genera have horns in the
male only.

One of the smallest of Antelopes belongs to an allied genus ;
this is Neotragus pygmaeus. It is known as the Royal Antelope,
a name apparently derived from Bosnian's statement that the
negroes called it " the king of the harts." Its horns are very
small. The height of the animal is only 10 inches. Horns are
present in the male alone. The last three genera are African.

The Cervicaprine series, which is also African, includes the "Water-
bucks and Eeedbucks, so called on account of their water-loving
propensities. As in the last series, from which they are separated by
Sclater and Thomas, but with which they are united by Flower,
there are horns in the male only. These horns, though not twisted,
are long. The typical genus is Coins, of which there are eleven
species. The Waterbuck, C. ellipsiprymnus, and the Sing-sing,
C. unctuosus, are perhaps the best-known species ; the former is

xi THE SAIGA ANTELOPE 3 I I

blackish grey, the latter browner in colour. In C. maria and one or
two other species the horns are more curved backwards and again
forwards than in some of the others, where their form is sublyrate.

The Eeedbucks, Oervicapra, are closely allied to Coitus ; they
are, however, of smaller size. Here, as in that genus, the females
are hornless, and the horns of the males are of medium size.
Five species are referred to the genus. They are all of a
brownish fawn colour. A genus Pelea, with but one species, P.
capreolus, has been separated on account of the fact that the
horns are nearly straight and that there is no naked patch of
skin beneath the ears. This animal has received its name on
account of its resemblance to the Roebuck.

The Antilopine section includes a number of genera.

The genus Antilope is Indian in range. It includes but one
species, A. cervicapra. This Antelope is of medium size, with a
brown pelage getting blacker with years; it is thus known as the
Black-buck. The female, which is hornless, is lighter brown. The
horns are long, spirally twisted, and closely ringed.

Aepyceros, with two species, is African. The Palla (Ae. melam-
pus) is a large Antelope, with longish lyrate horns in the male,
which are half-ringed.

The Saiga Antelope, genus Saiga, is one of the most remark-
able types of Antelope in its outward appearance. Its nose
is very large and inflated, the two nostrils being quite widely
separated, a depression indeed lying between them dorsally.
The horns are lyrate in the male, absent in the female. The
" ovine expression " of this bovine animal is more pronounced in
the female. Corresponding with the clumsy nose are very short
nostrils, the commencement of the narial aperture being therefore
very far back. It is almost suggestive of Macrauchenia in this
respect. The fleece is also Sheep-like. The genus occurred in this
country during the Pleistocene. It is now an inhabitant of Eastern
Europe and Western Asia. The only species is S. tartarica.

The Chiru, Pantholops, is allied to the Saiga. The horns of the
male are long and nearly straight ; they are ringed in front. The
muzzle is swollen in the male ; the nostrils are large, and provided
with extensive sacs internally. The colour of this animal, which
is exclusively Thibetan in range, is a pale fawn. The hair, in
accord with its habitat, is very woolly. No living specimens have
ever been brought to Europe. This creature has accumulated much

3I2

LODER'S GAZELLE

legend. Its blood is believed by the Mongols to possess- virtues,
and by means of the rings on the horns fortunes are told. Natur-
ally the animal is on these grounds hard to stalk and shoot.

The Gazelles, genus Gazella, are fairly numerous in species,
which are both Palaearctic and Ethiopian. There are altogether
twenty-five of them. The genus as a whole is characterised by
the small or moderate size, the sandy coloration, with white belly,

FIG. 161. — Loder's Gazelle. Gazella loderi. x jV

the presence of dark and light stripes on the face and on the
flanks. These streaks, however, are not always present, and their
presence or absence serves to differentiate some of the species.
The horns are usually present in both sexes. The horns are of
fair length, ringed, and of lyrate form.

The Springbok is separated from the rest of the Gazelles, to
which genus it is clearly most nearly related, as a genus Anti-
dorcas. This genus differs from Gazella by having only two lower
premolars as in Saiga. Otherwise it resembles the Gazelles ;
there is but a single species, A. euchore, which is African.

XI

GAZELLE-LIKE FORMS

313

Ammodorcas is closely allied to the Gazelles, but differs from
them in having an elongated neck and also a long tail. A.
clarkei, the only species, is limited to Somaliland.

Lithocranius, not unlike the last, has a still longer neck,
which makes it almost Giraffe-like ; its -tail, however, is short.
The scientific name is derived from the " solid stony character of
the cranium." In running, this Gazelle carries the head forward
in a straight line with the body. It is African.

Dorcotragus with one species, D. megalotis, is a pigmy Gazelle
restricted to Somaliland. Its likeness, on account of size and in
some other superficial features, to the Klipspringer, led to its
original confusion with that genus (Oreotragus).

A sub-family Hippotraginae, or Hippotragine section, includes

FIG. 162.— Sable Antelope. Hippotragus niger. x -fa. The horns of the specimen
figured have not nearly reached their full dimensions.

a number of Antelopes which agree in the possession of four
mammae, and of molars more like those of the true Oxen, of

314

SABLE AND BEATRIX ANTELOPES

horns of some length, present in both sexes, and of a longish
tail. They are all African in range.

The type genus Hippotragus has its horns placed above the
orbits ; they are not twisted, but curved backwards. There are
three species in the genus. Of these the best known is H. niger,
the beautiful Sable Antelope. Its general colour is a rich, dark,
glossy brown with white stripes on the face, and with a white
belly. The other species are the Eoan Antelope, H. equinus, and
the Blaaubok, H. leucopkaetts, of which the last specimen was
probably killed in 1799.1

The genus Oryx (chiefly African, but also Arabian and Syrian)

FIG. 163.— Beatrix Antelope. Oryx beatrix. x TV (From Nature.)

also contains a number of species, which are fairly familiar through
the fact that several of them are always on view in the Zoo-
logical Society's Gardens. The genus differs from Hippotragus
in that the horns, present in both sexes, are placed behind the
orbits, arid slant backwards in a line with the face. They are
aunulated. The Leucoryx (0. leucoryx) is of a pale colour, but
1 W. L. Sclater, The Fauna, of South Africa, Mammals, i. 1900.

xi KUDUS AND ELANDS 3 I 5

this is not so marked as in 0. Beatrix, which is largely white
with, however, brown legs. The Gemsbok is a handsome creature
with greyish tawny colour, much darker on the legs, and with a
Gazelle-like, dark, side stripe. It has received its vernacular name
on account of its supposed likeness to the Chamois (" Gemse "),
just as the Rehbok was so-called from its supposed likeness to the
Eoe Deer, and the Eland to the Elk. The Beisa (0. beisa) is of
a similar tawny colour to the last, and also with darker stripes.

The Addax (Addax) of North Africa, Arabia and Syria, has
but one species (A. nasomaculatus). The horns are spirally twisted.

The Tragelaphine section includes the Kudus, Elands, Nilgais,

FIG. 164. — Speke's Antelope. Tragdaphits spekii ( 9 ). x TV

and Harnessed Antelopes. They are all long-horned (when the
horns are present in both sexes), the horns being twisted ; the
nose is naked with a slight median groove, and all are Ethiopian
or Oriental in range.

The genus Tragelaphus includes the Harnessed Antelopes, so
called on account of the direction of the stripes suggesting
harness. The females are hornless, and the colours of the two
sexes are different. The hoofs are long and the toes rather
unusually separable, which state of affairs is in accord with the

LIVINGSTONE'S ELAND

swampy country affected by many. rl\ grains and T. spekei are
larger forms ; the Boschbok, T. sylvaticus, is smaller.

The Kudus, genus Strepsiceros, have more markedly twisted
horns, which are absent in the female. The body is vertically
striped with white. The largest species is S. kudu; a smaller
form, S. imberbis, is from Somaliland.

The last genus of this section or sub-family is the African Eland,
genus Oreas a (which it appears should be spelt Orias). The Elands

FIG. 165.— Elan

are perhaps more Ox-like in appearance than the other members of
this group, and in both sexes have horns, in which the spiral
twisting is more close. Orias canna is the name of the common
Eland. 0. livingstonii has been applied to an East African
variety, which has thin and faint lateral stripes like the other
members of the group to which it belongs.

The genus Boselaphus includes only B. tragocamelus, the
Nilgai, which is purely Indian in range. The female is hornless,
and the horns of the male are smooth and not long.

1 Taurolragus oryx has unfortunately been discovered to be the correct name
for the Eland.

HYBRID OXEN

317

The members of the Bovine section or Oxen are to be dis-
tinguished from other hollow-horned Euminants by their stouter
build and by the fact that the horns stand out from the sides of
the skull and are simply curved, not twisted ; and smooth, not
annulate like those of other Euminants. The muffle is naked,
broad, and moist. The Oxen are widely distributed ; but are
entirely absent from the Australian region and from South
America and Madagascar.

The true Oxen are perhaps best considered to form but a
single genus, Bos. They have, however, been divided into a number
of genera. Even the supposed aberrant Anoa depressicornis of
Celebes hardly differs sufficiently to warrant its separation. In
favour of this view, too, is the extraordinary ease with which
different " genera " will cross with each other and produce fertile
offspring. The following is the pedigree of an animal lately
living in the Zoological Society's Gardens. The female offspring
of a male Zebu and a female Gayal was mated with a male Bison.
The female calf was again mated with a Bison and produced a
calf, also a female, which contained therefore the three species,
Bos indicus, Bibos frontalis, and Bison americanus. It is clearly
unwise in view of this fact to insist too much upon generic dis-
tinctions in any of those types.1

Of this genus the Oriental Gaur (Bos gaurus), the Gayal (B.
frontalis), and the Banteng (B. sondaicus) form a well-marked
section, characterised by their dark coloration and by the some-
what flattened horns.

The Gaur, Bos gaurus, has a more concave forehead than its
allies ; the horns are less curved than those of the Banteng, and
less so than the horns of the Gayal (Bos frontalis'). It inhabits
the Indian Peninsula ; and extends through Burmah to the ex-
tremity of the Malay Peninsula. The Malay name of this
animal is Sakiutan, which simply means wild cattle. It chiefly
frequents wooded hills and is an excellent mountain climber.

Bos frontalis, the Indian Gayal, has a white caudal disc like
the last species, but the forehead is flat and the horns curve but
little. It is chiefly known as a tame animal, and its occurrence
in the wild state has been doubted. It has furthermore been
suggested that it is merely a tame race of the Gaur altered

1 A. D. Bartlett, " On some Hybrid Bovine Animals bred in the Society's
Gardens," Proc. Zool. Soc. 1884, p. 399.

AUROCHS AND WISENT

slightly through domestication. It is, however, said not to cross
in a state of nature with the Gaur.1

FIG. 166.— Gayal. Bos frontalis.

The Banteng, B. sondaicus, is distributed through Chittagong,
Tenasserim, and the Malay Peninsula to Java and Borneo. There
are apparently two races of this animal. The species differs from
the others by the fact that the horns are smaller and more curved ;
there is a white caudal disc ; the forehead is narrower and the
skull longer than in the others.

The American Bison and the European Aurochs form another
section ; they are indeed extremely alike, specific differences being
hardly recognisable. The Bison of America, formerly present in
such numbers that the prairies were black with countless herds,
has now diminished to about a thousand head.

One of the largest of existing Bovidae is the Aurochs, Wisent, or
European Bison, Bos bonasus (or Bison europaeus). It is exceedingly
like its American relative. Formerly the animal was much more
widely spread than it is now, extending its range from Europe into
North America. It is now limited to certain districts on the Urals,
in the Caucasus, and a herd of them are kept up through the
fostering care of the Emperor of Kussia in the forest of Bielovege
in Lithuania. The term " Aurochs " should not really be applied to
this species but to the Wild Cattle, Bos taurus. It is, however, so
generally used for the Wisent (which is the German name) that it
1 See Proc. Zool. Soc. 1890, p. 592.

EXTINCT OXEN

319

is not necessary to change it. The Sclavonic name is Zubr or Suber.
It is a great beast, standing 6 feet or so in height at the
shoulder. It ranged further over Europe well within the historic
period. In the days of Charlemagne it was spread over Germany
and was a beast of the chase. In the year 1848 the Emperor
of Russia presented a pair of these Oxen to the Zoological Society
of London. At the time of their presentation an interesting
communication was made to the Society by M. Dolmatoff, on the
method of the capture of these two examples. The creature is
not easy to capture and is alarming to confront. " The eyes,"

FIG. 167. — Bison. Bison americanus. x ^5.

says an old writer, " are red and fiery ; the looks are furious and
commanding." It has of course the shaggy mane and hump of
the American animal. The herd in Lithuania was said to be
1900 in the year 1856. Mr. E. N. Buxton,1 who has lately
visited the forest, quotes M. Neverli to the effect that at present
the numbers are not more than 700.

Allied to this animal, and apparently still nearer to the American
Bison, is the extinct R priscus of Europe. The Pleistocene Bisons
of North America, R antiquus and R latifrons, are not remote from
the living forms. Finally, the Miocene R sivalensis from India,
and the Pliocene Rferox and R alleni of North America, take back
this group to as remote a period as any other genus of Oxen.

1 Proc. Zool. Soc. 1899, p. 64.

-•>* «,

320

THE THIBETAN YAK

The Yak, Bos grunniens, is a long-haired peculiar type, confined

FIG. 168. — Yak. Bos grunniens. x

to the Thibetan plateau. B. (Anoa) depressicornis of Celebes is
characterised by its straight horns ; allied to it is B. mindorensis

FIG. 169. — British Wild Ox. Bos taurus. From Vaynol Park, Bangor. x-jV

(Philippine Islands), supposed, however, to be a hybrid between

CHILLINGHAM CATTLE

321

it and some other species. Africa has at least two Buffaloes. We
may finally mention the Wild Ox of Europe, B. pritnigenius, the
supposed progenitor of our domestic cattle, believed to be still
surviving in the herds at Chillingham, Chartley, and elsewhere.
This animal is sometimes called the Aurochs. The Romans spoke
of it as the Urus, and it appears to have formerly attained to
more gigantic proportions than at present. It is the small size
of the present race that is the chief objection to tracing them
back to the large Oxen existing near London in 1174, and found
sub-fossil in the Cambridgeshire fens.

Of the true sheep, genus Oms, there are a considerable number

FIG. 170.— Punjab Wild Sheep. Ovis vignei.

of species. The Sheep are to be distinguished from the Goats by
their rather stouter build and by the absence of the beard in the
male. The horns are developed in both sexes, and are usually
twisted and often of large size.

The Sheep are almost entirely Palaearctic and Nearctic. They
only just get into the Oriental region. One of the finest species
is the great Pamir Sheep, 0. poli, whose length reaches 6 feet

Y

VOL. X

322

AMERICAN SHEEP

CHAP. XI

7 inches, and height 3 feet 10 inches. The horns of this fine
Sheep may measure more than five feet round the curves.
The Kocky Mountain Bighorn (0. montanci} is a Sheep ranging
along the Eockies as far south as New Mexico, and also to the
far north ; they are not confined to the chain of mountains
mentioned, but occur also on the mountains of British Columbia
down to those of California. The horns are not quite as large

FIG. 171.— Himalayan Burrhcl Sheep. Ovisburrhd.

. r-i
V (From Nature.)

as those of the last species, but measurements give a length
(along the curve) of 32 to 40 inches.

Just as the Goats are often limited to islands and small
stretches of country, so are the Sheep. Thus Cyprus has a
species, 0. ophion, peculiar to itself. This, which is known as
the Cyprus Mouflon, is limited to a range of mountains, the
Troodos, in that island. In 1878 it was believed that the
animal was nearly exterminated, a flock of twenty-five members
alone surviving. They have, however, since increased. Confined

'i- XiV (From Nature.)

FiG.,173.— Barbary Sheep. Om tragelaphus. x

324

SHEEP AND GOATS

to the Thibetan plateau are 0. hodgsoni and 0. naliura. Corsica
has the Mouflon, 0. musimon ; and the Barbary Sheep or Arui,
0. tragelaphus, is found only in Northern Africa. Ovis Iturrliel
and 0. Uanfordi are Indian forms.

Ovis nahura is chiefly responsible for the impossibility of
strictly separating the Sheep and Goats. It has no suborbital
glands or lachrymal fossae, which are as a rule present in the

FIG. 174. — Thar. Capra jemlaica. x

(From Nature.)

Sheep and absent from the Goats. On the other hand inter-
digital glands are present, which is the case with Sheep. Its
habits, too, are a blending of those of the Sheep and the Goat.
It lives largely on undulating ground like Sheep, and frequently
lies down during the day on its feeding ground. On the other
hand it is, like the Goats, a splendid climber.

The Goats, genus Capra, differ from the Sheep in their slighter
build and in the fact that the horns are not spirally curved, but
arched over the back There is also the characteristic beard,

xi THE WILD GOAT 325

and the male is odorous. The true Goats are almost exclusively
Palaearctic in range. They show the limited distribution of the
Sheep, a distribution which follows from their mountain-loving
habits.

Thus we have the Spanish Ibex (C. pyrenaicd), limited to
the Pyrenees and other mountain ranges of the peninsula;

FIG. 175. — Sinaitic Ibex. Capra sinaiiica. x j^.

C. ibex, the Steinbok of the Alps and the Tyrol ; the Markhoor,
C. falconeri, of certain mountain ranges of Afghanistan ; the
Caucasian, Sinaitic, and Cretan Ibexes, and the Thar.

Capra aegagrus, the Persian Wild Goat, ranges from the
Caucasus to Sind. It is this animal which produces the true
" bezoar stone." The substance in question is a secretion appar-
ently found in the stomach. It is still, according to Mr. Blan-
ford, regarded as an antidote to poison in Persia. Buffon called
this Goat the " Pasan," which is evidently a corruption of the
word bezoar. When the substance was in repute as a medicine
of the " alexipharmic " kind, the supply naturally came up to
the demand. Thus the bezoar stones of the Lama in South
America gained repute, and there were " Oriental bezoar, cow

326

BEZOAR STONES

bezoar, hog bezoar, and monkey bezoar " ! As concretions of one
kind or another are not uncommon objects in the alimentary
tract of mammals it was easy enough to obtain a fair amount of
some substance which was sure to sell well. It is said that a
stone weighing four ounces was once sold in this country (or at.
any rate in Europe) for £200.

" There can be no doubt," observes Mr. Blanford, " that
C. aegagrus is one of the species, and probably the principal, from
which tame goats are derived."

The Chamois (Rupricapra) and the Goral (Nemorrhaedus) are

FIG. 176. — Japanese Goat Antelope. Nemorrhaedus crisjms. XT\. (From Nature.)

best described as Goat-like Antelopes ; but, as already said, it is
difficult to split up the Bovidae satisfactorily. The Eocky
Mountain Goat, Haploceros montanus, is a large Goat-like creature,

THE MUSK OX

327

which has the peculiarity of having the shortest cannon bones of
any Ruminant. Its name denotes its range

O

= — -r-g^TOS ^^*^3(t^S«^SP^*^ S*. • 'f

FIG . 1 7 7 . — Goral . Nemvrrh aedus goral.

(From Nature.)

The Musk Ox, Ovibos moschatus, has been thought to be on
the borderland between the Sheep and Oxen, as indeed expressed
in its scientific name. It is a purely Arctic creature, now con-
fined to the Nearctic region ; but it formerly existed in the
Arctic regions of Europe.

The anatomy of the " soft parts " of this genus has lately
been investigated by Dr. Lonnberg.1 The animal has no foot
glands such as occur in Ovis. Its kidneys, however, are non-
lobate, and it has orbital glands. The cotyledons of the placenta
are unusually large, and the cow has the " primary four " teats.
It cannot, in fact, be definitely referred to either the Caprine or
the Bovine section of the Cavicornia, and while possibly most allied
to Budorcas, it may be regarded, at least for the present, as
entitled to form a separate sub-family of its own. The muzzle

1 Proc. Zool. Soc. 1900, p. 142.

328 EXTINCT PIG-LIKE ANIMALS CHAP.

has a slight naked strip above the nostrils, as in the Sheep, but
there is no fissure of the upper lip.

Extinct Families of Artiodactyla.

The origin of the Artiodactyla is placed by Cope in the family
Pantolestidae,1 allied to the genus Protogonodon of the Condy-
larthra. As, however, this family is represented by but a few
back teeth and a fragment of the hind-foot, it seems premature to
regard it as the necessary starting-point of the Bunodont and
Ruminant groups.

Fam. Anthracotheriidae. — This well-known and ancient
family consists of creatures of for the most part a Pig-like form,
with teeth approaching the selenodont shape, and a complete
dentition. The carpals, tarsals, metacarpals, and metatarsals are
all free. The toes are four (or five) to each foot, with the outer-
most beginning to be reduced. These of course are all generalised
and primitive characters, pointing nowhere in particular, except, of
course, to an Artiodactyle stock, on account of the teeth . and the
two predominating toes.

The type genus of the family, Anthracotherium, is not, as its
name might seem to denote, a relic of the Carboniferous period ;
its remains were found in lignite, which may also show that it
was at least semi-aquatic in habit. Its form, however, must
have been Pig-like, so at least one would presume from the
elongated skull and shortish legs. There were species as great
as a Ehinoceros, and smaller forms. The genus began in the
Oligocene and continued down to the Pliocene. It is known
from Europe, Asia, and America.

The skull is long with a prominent sagittal crest. The
facial part is also very long, and the orbits are not closed by a
bony ring. The premolars are simple teeth ; the molars dis-
tinctly bunodont with a tendency in one or two to the seleno-
dont condition. The canines are powerful, as are also the
incisors. The scapula has been specially compared with that
of the Camel. It has no acromion, which is usually though
not always absent in Ungulates. An ally of the present animal,
for instance, the Hippopotamus, has the acromion developed.
The radius and ulna, the tibia and fibula, are all fully developed.

1 The name Triyonolestes has to be substituted for Pantolestes.

xi EXTINCT PECORINES 329

Ancodus (or Hyopotamus, as it has been called) is also Oligo-
cene in range, and its remains have been found in the same
countries as have those of Anthracotherium. Both genera are
indeed closely allied. Ancodus seems to be a more slightly-
built creature. The skull looks weaker, but presents much the
same features of organisation. In A. velaunus, a species found in
French rocks, a metacarpal of digit I. was present in the man us,
while A. hrachyrhynchus had a completely five-fingered manus.

The Miocene genus Merycopotamus (from the lower layers of
the Siwalik formation in India) is more distinctly selenodont
than the forms already discussed. On this ground it has been
placed in a separate sub-family. As, however, in other respects
it does not depart from the Anthracotherian type of structure, this
proceeding seems to be hardly necessary. There are two species
known, of which one, M. nanus, is, as its name denotes, a dwarf
form.

Fam. Caenotheriidae. — While the last family consisted
of animals rather more akin to the Pigs, the present is more
Pecorine in its characters. The molars are selenodont ; but as in
the Tragulidae the premolars are more of the nature of cutting
teeth. The dentition, like that of so many of these early
Ungulates, is complete, and the canines are not prominent. The
feet are four-toed, the lateral toes not reaching the ground.

The principal genus is the Eocene and Miocene Caenotherium.
Of this genus there were a considerable number of species all
European in range, and of small size — not more than a foot or so
in length. Their small size is suggestive of the Chevrotains.
In the skull the orbital cavity is nearly or quite surrounded
by bone, and the tympanic bulla is large and inflated. A
common feature of Artiodactyles, a failure of the nasals and
maxillae to meet at the side of the face, is to be seen in this
ancient forerunner of the Pecora.

Plesiomeryx, also European, and from the same geological
horizon, is a very closely allied form.

Fam. Xiphodontidae. — This family consists of slender,
small Artiodactyles which are, like the Caenotheriidae, related
to the Pecora. They are confined in their range to Europe.

The type genus Ziphodon has selenodont molars and elon-
gated, slender, cutting premolars. The dentition was complete
and the canines not highly developed. Like Caenotherium,

33O OREODON AND MESOREODON CHAP.

Xiphodon was a hornless creature, but with only two toes, the
two lateral digits being represented by the merest rudiments of
rnetacarpals. The other metacarpals were unusually long.

Amphimeryx (also called Xiphodontotlierimti) is much more
imperfectly known, but belongs to this family or to that of the
Caenotheriidae. Dichodon is another member of the same family.

Fam. Oreodontidae. — This family, consisting of numerous
genera, is limited to the North American continent. Its range
in time is from the Eocene to the Lower Pliocene. The family
as a whole is to be distinguished by a number of primitive
characters. The dentition is complete ; the feet are four- or
even five -toed ; the orbit is sometimes open behind. The canines
of the lower jaw are not more pronounced than the incisors.
The characteristics of the group will be further developed by
a consideration of some of the principal genera which are in-
cluded in this family.

Oreodon is a Miocene form about as large as a Peccary. The
skull has a short face with a completely-closed orbital cavity.
In front of the orbit is a deep pit, not a mere deficiency of
ossification, such as occurs in many Artiodactyles. This is
placed on the lachrymal bone, and is in fact a lachrymal fossa,
such as occurs in other forms. The odontoid process of the axis
vertebra is somewhat cheese-taster shaped, as in recent Artio-
dactyles. There are fourteen dorsal vertebrae and a very large
number of caudals. The radius and the ulna are completely
separated, as are the carpals. There are five digits to the fore-
limbs. The fibula is complete and independent. The hind-foot
is four-toed. Several species of the genus are known.

Merycochoerus is an allied Miocene genus. It is more
massive in form than the last, but otherwise does not present
differences of importance.

Mesoreodon is another genus of this family which presents
some curious features of organisation. In the skull and teeth
there is nothing very noteworthy, but the hyoid is remarkable.
This appendage of the skull is by no means always preserved,
and when it is, it might be denied that it belonged to any
particular skull. In the present case there appears to be no doubt
as to the identity of the bones, which resemble the corresponding
bones of the Perissodactyla much more than they do those of
other Artiodactyles. Associated with the bones an ossified

xi " HOWLING " OF MESOREODON . 331

thyroid cartilage of the larynx was found. As the skull was
that of a male, this character may be a sexual one. It is quite
comparable to the ossification of the same cartilage in the
American monkey CallitJirix. " The function of the bone,"
observes Professor Scott,1 " was probably similar to that per-
formed by the enormously -inflated basihyal of the howling
monkeys, and must have given to these animals most unusual
powers of voice." Another important anatomical fact about
Mesoreodon is the apparent existence of a clavicle. It is of
course conceivable that the remains of some other animal have
got mixed up with that of the individuals upon which the
present genus is founded ; but failing that, here is a clavicle in
an Ungulate. The spine of the scapula possesses a metacromion.
This greater development of the spine of the scapula in Artio-
dactyles than in Perissodactyles is, it is suggested, to be cor-
related with the earlier loss of the clavicle in the latter group of
Ungulates.

Cyclopidius (synonymous with Bracliymeryx) is a kind of pug
form of Oreodon. The skull is short and broad, and the end of
the snout a little turned up. The upper incisors are small and
drop out early. On each side of the nasals is a large oval
vacuity which is perhaps to be compared to the lateral deficiency
to be found in other Artiodactyles. One species of this singular-
looking form is appropriately called C. simus.

Other allied genera are Merychyus and Leptauchenia. The
former extends as far down as the Lower Pliocene, and is thus
one of the newest forms of Oreodontidae.

AgriocJioerus2 (Fig. 1 78) is placed in a separate sub-family from
the types which have just been considered. It is Miocene in range.
It differs from Oreodon and its closer relatives by the fact that the
orbit is open behind and not closed. The most remarkable fact
about this creature is that the terminal phalanges of the digits
(five in the fore- and four in the hind-feet) being pointed, seem
to suggest their encasement with claws rather than hoofs. The
pollex, though small, seems to have been opposable. As with
other Oreodonts, the molars are selenodont. The premaxillae
are toothless — at least in adults, for two teeth are present in the

1 Trans. American Phil. Soc. xviii. 1896, p. 125.

2 For complete osteology see Wortman, Bull. Amer. Mus. Nat. Hist. vii. 1895,
p. 145.

332

PROTOREODON

CHAP.

young. There are
several species. Agri-
ochoerus, like Oreodon
and primitive Ungu-
lates in general, had a
long tail. The genus
thus shows a mixture
of ancient and special-
ised characters.

The most ancient
form of Oreodont is
Protoreodon. This is
Eocene, and became
extinct during that
period. It had a com-
plete dentition, open
orbit, and no lachry-
mal fossa. The fore-
feet were five -toed,
the hind four-toed.

Fam. Anoplo-
theriidae. --This
family is entirely
Eocene in point of
time, and is unknown
outside Europe. The
dentition of the
group is complete ;
the molars are seleno-
bunodont, like those
of the Anthraco-
theriidae. The bones
of the carpus, tarsus,
metacarpus and meta-
tarsus are all free ;
the toes are four to
two in number on
each foot. The orbit
is widely open behind.
The tail is long, as in
Xiphodon, etc.

xi EOCENE ARTIODACTYLES 333

These general characters only just serve to differentiate the
family; but they illustrate its archaic character, in which it
resembles the Xiphodontidae, and even more the Anthraco-
theriidae. A survey of some of the genera which have been
assigned to the family will bring out other features in the
organisation of these very ancient Artiodactyles.

Anoplotherium is so called on account of the fact that it is,
like all ancient Artiodactyles, without horns or claws. Tusks it
might have, but as a matter of fact has not. There are, as in
Artiodactyles generally, nineteen dorso-lumbar vertebrae; the
long tail has numerous chevrons. The shoulder blade has a well-
marked acromion and a distinct coracoid process; it is wide
proximally. The bones of the fore-arm and fore-leg are, as is
usual in primitive Artiodactyles, separate.

In the skull the chief features, in addition to that mentioned
in the definition of the family, are the large size of the par-
occipital processes ; there is no fossa lachryrnalis or deficiency in
the side of the face. The animal is three-toed, both in the fore-
and hind-limbs. The second toe is nearly as large as the Artio-
dactyle third and fourth. There are tiny rudiments of the two
remaining fingers. The hind-foot is also three-toed, and there is
a trace of the hallux. The fingers are so widely separated and
divergent from each other that it has been suggested that the
animal had webbed feet and inhabited marshes, in which it swam
by the aid of its long tail. * The creature was the size of a Tapir.

Closely resembling Anoplotherium are a number of other genera.

Diplobune ( = Hyracodontotherium) was much like the last,
but was a more delicately-formed animal. The fingers and toes
(three of each) end in such sharply-pointed phalanges that claws
seem to be almost suggested. There are several species of this
genus. Dacrytherium differs by the presence of a lachrymal fossa.

Dicliobune has four-toed extremities, of which the lateral ones
are more slender and shorter than the two middle ones. As in
other Anoplotheriidae, the anterior premolars are furnished with
a sharp cutting edge.

Order V. SIRENIA.

Aquatic Mammalia, with but few scattered hairs ; hind-limbs
absent; fore-limbs paddle-shaped ; tail flattened, and either Whale-

334 LIMBS OF SIRENIA

like or rhomboidal to circular in form. Nostrils on upper surface
of not specially -elongated snout. Clavicles are absent. The
scapula has the normal mammalian form, with a well-developed
and roughly median spine. The bones of the arm and hand articu-
late together, as in land animals ; the phalanges show at most
traces of increase in number above the normal. Pelvis represented
by a vestige, more highly developed in some fossil than in recent
forms. Stomach complex, consisting of several chambers. Lungs
simple and not lobulated. Diaphragm oblique and very muscular.
Brain peculiar in form and but slightly convoluted. Testes ab-
dominal. Teats two, and pectoral in position. Placenta noii-
deciduous and zonary.1

This limited group consists of purely aquatic forms, which are
both marine and fresh-water in their proclivities. They have
been placed in the immediate vicinity of the Whales ; but it is
now believed by most zoologists that the likenesses which they,
undoubtedly show to the Cetacea are of an adaptive kind and
related to their similar mode of life. The group is a readily -
definable one. Externally they are marked by their dark
coloration, somewhat Whale-like though of clumsier build, and by
the total absence of external ears and hind-limbs ; the latter are,
however, as will be pointed out shortly, marked by certain rudi-
mentary bones. There is a flattened tail, which in the Dugong and
Rhytina is precisely like that of a Whale. It is interesting to
note that the former genus, whose tail is, judging it at least by
the standard of the Whales, more completely modified for the
aquatic life, should also show other features which indicate their
longer life as marine creatures. For the flippers are more Whale-
like in that the fore-arm is completely enclosed within the body,
or nearly so, and the nostrils have a more decidedly superior
position than in the Manatee. The fore-limbs of this group, as
may be inferred from what has just been said, are flipper-like ;
but, contrary to what we find in Whales, the phalanges do not as
a rule show any traces of multiplication, so characteristic a feature
of the Cetacean hand, and the individual bones are connected by
well-formed joints. Beneath the thick skin, which is sparsely
provided with stout hairs in the Dugong, is a layer of blubber.
Dr. Murie has called attention to the fact that this layer in the

1 In Halicore ; probably also in Manatus. See Turner, Trans. Roy. Soc. Edinb.
xxxv. 1889, p. 641.

xi DISTRIBUTION OF MANATEE

335

Manatee l differs from the blubber of the Whale ID that there
is no free oil anywhere.2

The skeleton of the Sirenia is strong and massive, thus con-
trasting with the loosely-textured bones of the Cetacea. The
cervical vertebrae are, as a rule, free, but the second and third are
fused in Manatus and the extinct Halitherium. It is noteworthy
that in Rhytina the cervical vertebrae have the exceedingly thin
centra that characterises the neck vertebrae in Whales. The ribs
are most of them firmly articulated by two heads. The breast-
bone is generally reduced, as in AArhales ; and but few ribs are
attached thereto. The vertebrae, moreover, are well locked to-
gether by zygapophyses, and not loosely attached as in Whales.

The shoulder blade is long and narrow, and not unlike that of
the Seals. It is totally unlike the peculiarly-modified scapula
of the Whale tribe. But, as in the latter, there are no clavicles.

The hind-limbs are only represented by the pelvis ; and this
is a rudimentary structure, varying, however, in the degree of its
degeneration. That of the extinct Halitherium recalls the pelvis
of the Korqual. There is a single triradiate bone, with an aceta-
bular cavity for the rudiment of the femur in the centre ; it
suggests that here the three normal elements of the pelvis have
become fused into a single bone. In the Dugong there are two
small bones on each side.

The Manatees (Manatus} 3 are found in the fresh-waters and
along the Atlantic coasts of South America and Africa. It
appears that there are four species, of which one only is African,
the others American. Report asserts the former occurrence of
this genus on the shores of St. Helena.

The Manatee is provided with only six cervical vertebrae, a
fact which distinguishes it from the other existing genera of its
group. A remarkable feature which it exhibits is the large
number of molar teeth. These apparently go on increasing in-
definitely during its life, the suggestion being that they are worn
away by the nature of the food — algae with much sand intermixed.
As many as twenty molar teeth have been counted in one half of
the jaw, and there is no reason to forbid the assumption that they

1 Kiikenthal has discovered a thick coating of rudimentary hairs in the foetus
of the Manatee, thus showing that it is the descendant of an animal furry like
a Seal.

2 "On the Manatee," in Trans. Zool. Soc. vol. viif. 1872, p. 127.

3 Hartlaub, " Beitrage zur Kenntnis der Manatus- Arten," Zool. Jahrb. 1886, p. 1.

336

UPPER LIP OF MANATEE

may get still more numerous.
This large number of grinding
teeth is obviously suggestive of
the Whales, with which the
Sirenia are believed by some to
be allied. It is at least a re-
markable coincidence that these
two aquatic groups of mammals
should both have assumed the
same indefinite tooth formula.
It is correct to say assumed, since
extinct forms of Manatees, such
as Halitherium and Prorastoma,
have not a continuous succession
of molars. The brain of the
Manatee is, contrary to the
usual arrangement among aqua-
tic mammals, smooth, and only
marked by one or two fissures.

The Manatee * is black in
colour, its thick skin being
wrinkled. The animal is assisted
in feeding by a curious mechan-
ism of the upper lip ; this is split
in two, and the two halves, which
are furnished with strong bristles,
can play upon each other like
the points of a pair of forceps.
The flippers are furnished with
nails, save in M. inunguis, but in
the nailed forms it is not every
finger which is thus armed.

Halicore? the Dugong, is an
entirely Oriental and Australian

1 Beddard, " Notes upon the Anatomy
of a Manatee (Manatus inunguis)," Proc.
Zool. Soc. 1897, p. 47.

2 See Kiikenthal in Semon's "Zoolog.
Forschungen," Denkschr. Jen. 1897 ;
Langkavel, "Der Dugong," Zool. Gartenf
1896, p. 337.

THE DUGONG 337

form ; there appears to be but a single species, though more than
one name has been given to supposed distinct species. As already
mentioned, it differs from the Manatee in the possession of a
Whale-like tail ; the nostrils, too, are more upon the upper surface
of the head, and there are no nails upon the flipper. The peculiar
cleft lip of the Manatee is not so well developed in the Dugong,
but there are traces of it ; and in the foetus the likeness to the
Manatee in this respect is very striking. It would thus appear
that Halicore is a stage in advance upon Manatus; that the
remarkable mechanism of the lip of the latter has been possessed,
but has been lost, by the Dugong. The skull of the Dugong is
distinguished by the stout premaxillary bones, which bear a tusk
in the male. In the female the tooth is there, but is lodged within
the bone. This incisor has a milk forerunner. The back teeth
of the Dugong (there are no canines) are few in number (four or
five, even six), thus showing a gradual reduction when compared
with Manatus ; and this culminates in the toothless Rhytina. It is
also interesting to notice that in the massive lower jaw there are
traces of an incisor. Were this to be developed into a tusk, the
jaw would present a curious resemblance to that of Dinotherium.

The Dugong, H. dugong, has the reputation of being the
original of the mermaid legends, since the young is held to the
pectorally-situated breast with one nipper. " But it should be
remembered," justly observes Dr. Blanford, " that stories of
beings half man or woman, half fish, are as common in temperate
as in tropical seas, and that some of them are more ancient than
any European knowledge of the Dugong."

Extinct Sirenians. — The earliest genus that can be with
certainty referred to this order is the Oligocene Prorastoma.
This genus, though offering no particular skull-characters that
assist in the determination of the much-debated affinities of the
Sirenia, shows a remarkable condition of the teeth that may
afford a clue. The species P. veronense, recently described by
Mr. Lydekker,1 is founded upon a fragment of the skull which
contains two teeth apparently representing the third and fourth
upper milk molars. The interest attaching to these teeth lies in
the fact that they clearly exhibit the buno-selenodont condition
characteristic of certain early Artiodactyles, e.g. Merycopotamus.

Halitherium is a later genus, which is known- by the nearly

1 Proc. Zool. Soc. 1892, p. 77.
VOL. X z

338 STELLER'S SEA-COW

complete skeleton. The skull is like that of other Sirenia, with
the down-turned premaxillary region. But the nasal bones, lost,
or at least rudimentary, in recent forms, are well developed ; the
likeness of ancient to living forms in this respect being exactly
paralleled by the Zeuglodonts, when compared with recent Whales.
The vertebral centra exhibit distinct epiphyses, which have dis-
appeared in living Sirenians. The cervical vertebrae are seven, of
which the second and third are occasionally fused. There are
nineteen pairs of ribs, and there are three lumbar vertebrae.
The sternum consists of three separate pieces. There is a
rudimentary femur.

The recently -extinct Steller's Sea-cow, belonging to the genus
Rhytina, was a huge beast, seen in the flesh up to nearly the end
of the last century. It frequented the shores of Bering's Straits.
Its remains occur in the peat on the shores of those seas. It reaches
a length of some 20 to 30 feet. The external characters were
much like those of other recent Sirenians. The nostrils were above
the fore part of the snout, the latter being truncated and obtuse.
The tail was of the Cetacean pattern, and thus like that of Halicore.
The head of this Sireniau was small, and the teeth had entirely
vanished save for the apparent existence as transitory structures
of two small incisors in the upper jaw. The absence of teeth
was compensated by the presence of a horny palate for the
trituration of the sea-weeds which constituted the food of Steller's
Sea-cow. The fore-limbs seem to have possessed no nails, but
were covered at the extremity with short, bristly hairs, no doubt
serving the purpose of keeping the animal moored in safety to
the slippery beds of Fucus upon which it browsed.

There are nineteen pairs of ribs. The vertebrae of the
cervical region are the customary seven, and the centra are thin
and plate-like as in the Cetacea, the animal being thus short-
necked like those marine creatures.

CHAPTER XII

CETACEA — WHALES AND DOLPHINS

Order VI. CETACEA.1

AQUATIC Mammalia of fish -like form; tail expanded into
horizontal flukes ; a fatty dorsal " fin " present in most species ;
anterior limbs converted into fin-like paddles ; posterior limbs
only represented by skeletal rudiments. Hairy covering re-
duced to a few isolated hairs in the neighbourhood of the
muzzle. Nostrils represented by the single or double blow-hole,
nearly always situated far back upon the skull. Bones of loose
texture and much impregnated with oil. The skull has a greatly-
developed facial portion ; supra-occipital bones meeting the frontal
by overgrowing, or growing in between the parietals; bones
surrounding the organ of hearing loosely attached to the skull,
the tyinpanics of peculiar cowrie-shell form. Coronoid process of
mandible absent, or very feebly developed. Teeth, when present,
few or numerous, always of simple conical form, with at most
traces of additional cusps {Inici} ; if absent their place taken by
whalebone. Cervical vertebrae of short antero-posterior diameter,
often more or less completely welded together into a single mass.
Articulations between dorsal and other vertebrae feeble. Scapula
peculiarly flattened ; acromiou strongly developed as a rule, but
arising from a slightly- marked spine ; coracoid process generally
strongly developed. Phalanges of digits always more numerous
than in other mammals. Clavicles absent. Stomach complex,
consisting of at least four and often more chambers. Lungs
simple and non-lobulated. Diaphragm obliquely set and very

1 See van Beneden and Gervais, Osttographie des Cetaces ; and for a more general
account Beddard, A Book of IVTiales, London, Murray, 1900.

340 SIZE OF WHALES

muscular. Brain much expanded transversely and well con-
voluted. Testes abdominal Teats two, inguinal in position.
Placenta diffuse and non-deciduate.

The Whales and Dolphins, which constitute this order, form an
assemblage which is easily characterised by reason of the fact
that their affinities to other groups of Mammalia are so doubtful
that they furnish matter rather for speculation than for authori-
tative statement. Some hold that they resemble in certain points
the Ungulata ; while others again see in them the culminating
term of a series which commences with such a form as the Otter,
and of which the Seals and Sea-lions are intermediate stages. A
third opinion is that the Whales have arisen from some low
mammalian stock, too primitive to be assigned to any existing
order of mammals. Palaeontology, as will be seen later, throws
no light whatever upon their origin. This matter has already
been referred to (see p. 120) in considering the position of the
Cetacea

The Whales include the most gigantic of all the orders of
vertebrated animals. No creature living or extinct is so large as
the Sibbald's Eorqual, which attains to a length of some 85 feet,
or perhaps even rather more. On the other hand we have what
are by comparison minute forms. Apart from the possibly pro-
blematical Delphinus minutus, stated to be only 2 feet in length,
we have as a minimum 3 or 4 feet. The size of the Cetacea
has been subjected to much exaggeration. The first duty of a
Whale, observed the late Sir William Flower, is to be large ; and
Natural Historians, in the recent as well as in the remote past,
have not hesitated to put very round numbers upon the dimen-
sions of the larger members of the order. We may perhaps
pass over Pliny's " fish called balaena or whirlpool, which is so
long and broad as to take up more in length and breadth than
two acres of ground," and a number of analogous exaggerations,
which gradually dwindled down to the dimensions just stated of
the great Eorqual. M. Pouchet has made the ingenious sug-
gestion that the statements of the ancients may have been nearer
the truth than observations of to-day would have us believe ;
he pointed out justly that in former times Whales were not so
relentlessly pursued as during the last century ; the inference
being that they may have lived to a greater age, and attained
a more colossal bulk. The more modern exaggerations in the

HAIR OF WHALES

341

dimensions of the bigger Whales are probably due to the fact that
measurements have been taken, not in a straight line from snout
to tail, but along the bulging sides of the Cetacean, rendered even
more convex than in nature by decomposition, and by pressure
due to the immense tonnage of the creature.

The Cetacea are the most perfectly aquatic of all mammals ;
they never leave the waters which they inhabit. It is true that
legends have represented them as pasturing upon the shore —
Aelian spoke of Dolphins basking in the sun's rays upon the
sand ; and the " Devil Fish " of California, Rhachianectes (see
p. 357) has given rise to improbable stories — but they are

FIG. 180.— Killer. Orca gladiator. x-4V (After True.)

apparently only legends. Indeed a stranded Whale cannot
live long, for it is unable to breathe, the comparatively feeble
breast being crushed by its own weight. In accordance with
the purely aquatic habit, we find a modification of the outward
form of the body (and as we shall see later of many of the in-
ternal organs), which renders the Cetacea externally unlike all
other mammals. The form is fish:like, the fore-limbs are paddles,
the tail is expanded into two horizontal flukes, which serve to
propel the creature through the water.

The skin is smooth and shiny, so smooth and so shiny that it
1ms often been compared to coach leather. But nevertheless
they are not entirely without that most essential character of the
class Mammalia, a coating of hair. The hairy covering is, how-
ever, reduced to the very smallest proportions ; it is represented

342 DISAPPEARANCE OF HAIR CHAP.

. _> _

by a few hairs only — so few that they can be counted with ease
— in the neighbourhood of the muzzle. These hairs are not
present in all Whales ; they are absent, for example, in the White
Whale or Beluga. When present they are not furnished either with
sebaceous glands or with muscular fibres, which are such universal
concomitants of the hair follicles in the Mammalia generally.
This appears to be conclusive evidence that the hairs, few as they
are, are still undergoing degeneration. The need for a furry
coat is removed by the presence of a thick coating of fat im-
mediately underlying the skin. This is known as the blubber,
and is the main incentive to the pursuit of Whales. It must
not, however, be assumed without further argument that the hair
is absent because its place is taken, as a mechanism for retaining the
heat, by the blubber ; for the Seal tribe possess both fur and blubber.
Another conceivable explanation is quite at variance with such a
view of economy. It may be noticed that among Ungulates there
is a tendency to lose hair, particularly among more or less aquatic
forms. Thus the Hippopotamus is almost naked (as is indeed the
Walrus) ; the Ehinoceros, too, often a frequenter of marshy soil, is
almost as denuded as is the Hippopotamus. It is not, however,
settled that the Whales have anything to do with the Ungulata ;
otherwise an additional argument might be used, that is, the
secular loss of hair in some members of this group. The Hairy
Rhinoceros, Rh. tichorkinus, was, as its name denotes, a hairy
beast ; the Mammoth was equally so. The descendants, or at
least the modern representatives of both these creatures, are but
scantily clad with hairs.

A final reason for the naked character of the skin in exist-
ing Cetacea is closely connected with a feature in the organisa-
tion of three or four living species which must first be
described.

Some years ago the late Dr. J. E. Gray of the British Museum
described from the sea, off Margate, what he considered to be a
new species of Porpoise, characterised by the presence on the
dorsal fin of a row of stony tubercles. As a matter of fact it
was subsequently shown that the Common Porpoise has the same
structures, so that there was no need for a Margate species,
Phocaena tuberculifera. Moreover, in the Indian Neomeris, a
close ally of the Porpoise, a more abundant calcified covering of
scales exists along the whole back of the animal. These plates,

BONY SKIN PLATES 343

it has been discovered, are larger ii> the foetus, a fact which
naturally points to their being an inheritance from the past, now
undergoing retrogressive changes. Such a way of looking upon
the facts is confirmed by the finding, many years ago, by the
naturalist and physiologist Johannes Miiller, of bony plates in
connexion with the remains of a Zeuglodont Cetacean. It looks,
therefore, very much as if the Eocene ancestors of the modern
Cetacea had a skin studded with bony plates, as have the arma-
dillos. This being the case, the disappearance of hair is not
surprising. The room would be taken up by the calcified plates,
and when the latter disappeared, as they have in the vast majority
of existing Whales, the naked skin alone would be left.

Whales possess no externally-visible hind-limbs ; rudiments
of these appendages are present, which will be dealt with under
the description of the principal features of the skeleton. But
it has been discovered that in the Porpoise, external vestiges
of hind-limbs do appear in the foetus, a fact which, be it ob-
served, does away with the old view that the flukes of the Whale
are the last term in the series of vanishing hind-limbs, of which
the Seals, with their hind-limbs and tail bound up together, offer
an intermediate step.

The tail is fish-like in form, but the flukes are horizontal
instead of vertical as in fishes and Ichthyosaurus. This arrange-
ment is no doubt associated with the need for rapid return to
the surface waters after a prolonged immersion in search of food.
A downward stroke, such as is given by the powerful and large
tail flukes, would naturally bring about this result rapidly.
The tail, moreover, is under all circumstances the swimming
organ. Its motion has been stated to be slightly rotatory, like
that of a screw, and it is the case that the two flukes are often
alternate in shape like the flanges of a screw ; one being convex
upwards, the other convex downwards.

The fore-limbs are in the form of paddles, but they do not
apparently serve as organs of locomotion so much as balancers.
When a Whale is killed, it falls over on to one side, the office of
the flippers being to maintain the proper position, It is be-
lieved, however, from the fact that the embryo often shows a
relatively larger pectoral fin than that of the adult — the differ-
ence being due to a reduction in the adult of the number of
phalanges — that the fin was once an organ of progression.

344 FOUR-FINGERED WHALES CHAP.

The pectoral fin of "Whales exists in two forms. In the
Toothed Whales it is shorter and rounder; in the Whalebone
Whales longer and narrower. Structural differences accompany
these outward dissimilarities. In the first -named group the
humerus and the beginning of the radius and ulna are within the
body, and do not form a part of the fin. In the Whalebone
Whales, on the other hand, the fin contains all the bones of the
fore -limb. Another remarkable contrast between the hand in
the two groups of Whales is that while the Toothed Whales have
five fingers, thus justifying the prevailing opinion that they are
the more primitive of the two groups, the Whalebone Whales have
only four fingers. Actually the Eight Whale, Balaena, seems to
have five fingers ; and, indeed, the fact that it has, is often used to
distinguish it from the Humpback, which has undoubtedly only four.
But a careful consideration of the state of affairs which prevails in
the foetus of Balaenoptera dispels this idea. Between what are
apparently the second and third fingers, a rudimentary finger,
consisting of four phalanges, appears. This is not produced, as is
an additional finger found in the White Whale or Beluga, by a
splitting of a finger. Accordingly the four-fingered condition of
the Whalebone Whales is produced by the dropping out of a finger
in the middle of the series, — a very remarkable fact. When
fingers disappear, as, for intsance, in the Horse, etc., it is at the
two ends of the series that the digits vanish. If this view of
Professor Kiikenthal's * be accepted, it follows that the pre-
sumed thumb of the Right Whale is what has been termed the
prepollex.

The hand of the Whales, like those of some other aquatic
creatures, e.g. the reptile Ichthyosaurus, has a larger number of
phalanges than have terrestrial animals. The result of this is,
of course, to increase the length of the fin and its utility as a
paddle. It is commonly not all the fingers that have developed
this great number of accessory phalanges. Eudimentary nails
have been found upon the Cetacean hand ; but in no case are
they functionally developed. In the Manatees we have the
disappearance of the nails still imperfectly accomplished. In
M. latirostris there are nails ; these have vanished, apart from
possible traces to be seen with a microscope, in M. inunguis.

A very characteristic feature of certain Whales are the furrows

1 Vergleichend-anatomische Untersuckungen an Walthiere, Jena. 1889-93.

BLOWHOLE AND SPOUTING

345

to be seen on the throat. This is especially the case with the
Rorquals, in which group the Humpback Whale. Megaptera, is
to be included. The whales of
these two genera (Bcdaenoptera
and Megaptera) have a large num-
ber of the throat furrows — as
many as sixty have been counted.
Some other Whales have a smaller
number ; thus Rhachianectes has
but two on each side, and the
Physeteridae have not many more.
These furrows are absent in very
young embryos. It is thought
by Professor Kiikenthal that they
allow of a wide opening of the
mouth.

The blow-hole of Whales is, of
course, the aperture of the nostrils,
which are not so far back in the
foetus as in the adult. By the
characters of the nostrils the
Toothed Whales can be distin-
guished from the Baleen Whales ;
in the latter the orifice is double,
in the former single. In embryos
of Dolphins, however, the two aper-
tures are quite independent. The
phenomena of spouting have often
been misinterpreted.1 When the
Whale breathes, the expired air
rushes out through the nostrils.
The water vapour in the breath
condenses into drops of water in
the cold Arctic regions where
the phenomenon has been mainly
observed. Hence the idea that water taken in at the mouth is
expelled through the blow-hole. As the Whale approaches the
surface to breathe, it may be that some of the water of the sea is

1 "And at his gills draws in, and at his trunk spouts out, a sea," wrote Milton,
and think many others.

JI

FIG. 181. — Dorsal surface of bones of
right anterior limb of Round-headed
Dolphin (Qlobicephalus mdas). x *fo.
The shaded portions of the digits
are cartilaginqus. c, Cuneiform ; H,
humerus ; I, lunar ; R, radius ; *,
scaphoid ; td, trapezoid or magnum ;
U, ulna ; u, unciform ; //- V, digits.
(From Flower's Osteology.)

346 TOOTH GERMS IN WHALEBONE WHALES CHAP.

driven upwards by the forcible expulsion of air from the lungs.
But for the most part the water which is spouted is simply
condensed breath.

Like some, but not all, other aquatic Mammalia the Whales
have apparently no external ear. Indeed the opening of the ear
is excessively small. In a huge Eorqual it will " admit a quill " ;
and although " a quill " is rather vague, we may fairly allow any
sized quill without proving that the orifice of the auditory
passage is anything but exceedingly minute. As a proof, added
to so many, that the Whales are the progeny of terrestrial creatures,
we have the occasional traces of external ears.1

FIG. 182. — Left lower jaw of foetus of lialaenoptera rostrata. Inner aspect,
natural size, showing teeth. (After Julin.)

Whalebone Whales never possess permanent teeth as well as
the baleen ; but in the foetus are more than traces of true teeth,
which, however, never arrive at maturity. The whalebone itself is
described later (p. 354). That the Whalebone Whales possess teeth
while in the foetal condition was discovered so long ago as 180V.
It has since been confirmed by many observers. Not only is there
one set of teeth developed in the foetal Balaenoptera but two,
of which one comes to a greater maturity ; the other, in fact,
remaining at a very early stage of development. The more
complete dentition belongs to the milk series, as is the case with
the Toothed Whales. A very interesting conclusion with regard
to the derivation of the simple conical teeth of Whales seems to
follow from the development of these structures in Balaenoptera.
There are in the young foetus fewer teeth than in the more
advanced embryo. Now in the younger embryo some of the
teeth are furnished with more than one cusp ; they are bi- or
even tri-conodont. As Sir E. Owen observed, the teeth — some of

1 These have heen recorded by Professor Howes in the Porpoise.

THE STOMACH 347

them — are literally double teeth. This is a suggestion of the
more complicated teeth of the Zeuglodonts, and shows so far
that the simple conical teeth of existing Whales (of. however the
Platanistidae) are not by any manner of means so primitive as
their actual structure would undoubtedly lead one to believe.
Further than this, the greater number of teeth in the older
embryo coincided with the disappearance of these double teeth,
which seem to split up into the simple conical teeth.

The Toothed Whales are not furnished with baleen, but with
teeth only. These teeth are more or less numerous, their
arrangement being of value in the classification of the group ;
a matter which is dealt with later.

In the Narwhal, whose dentition in the adult is reduced to
the well-known tusk or tusks (properly developed only in the
male), there is a complete foetal dentition. A very curious fact
has been elucidated by Professor Klikenthal about the dentition
of the Common Porpoise. It appears that in this Cetacean the
two teeth corresponding to each other of the two dentitions may
fuse into a single tooth, which has in consequence a double crown.
It may be that this is the case with the Platanistid Inia, and
that its diconodont teeth are not, therefore, a reminiscence of the
comparatively complicated teeth of the ancient Zeuglodonts.

The internal organs of Whales which show the greatest
peculiarities as compared with other mammals are the stomach,
the lungs, and the diaphragm. Whales always possess a com-
plicated stomach divided into many, but into a variable number
of, chambers : there are as few as four in some, as many as fourteen
in Ziphioids.

On account of its complication the stomach l has been
compared to that of Kuminants — it has even been alleged that
Whales "ruminate" — but the comparison will not hold good.
Nor, on the other hand, is there a very close resemblance to the
equally-complicated stomach of the Sirenians.

The Eorqual has a stomach with as few compartments as any.
The only Whale which appears to have fewer is Balaena mysticetm,
where there are but three. In the Rorqual the oesophagus opens
into a more or less globular sac; from the upper end of this,
i.e. close to the entry of the oesophagus, arises the second chamber,
long and narrowish ; then follows an extremely short third sac,

1 For details and literature see Jungklaus, Jen. Zeitschr. xxxii. 1898, p. 1.

348 LUNGS AND DIAPHRAGM CHAP.

then a larger fourth, after which comes the dilated commence-
ment of the small intestine. The latter might be regarded as
a chamber of the stomach were it not for the fact that the ducts
of the liver and the pancreas open into it. This represents one
type of the Cetacean stomach, which seems to be found in all Whales
except the Ziphioids. In the latter, the oesophagus opens into
the first compartment as usual ; but the second division of the
stomach arises not close to the entrance of the oesophagus, but
at the opposite end. It would seem, therefore, as if the first
division of the stomach, found in most Whales, were missing in
Ziphioids. This way of looking at the matter is confirmed by
the fact that in Hyperoodon a remnant of the missing first
stomach is found in the shape of a small diverticulum of the
oesophagus just before it enters the stomach.

The essential difference between the Whale's and the Eumin-
ant's stomach is this : in the latter the stomach is primarily
divided into two portions, of which the first is uon- digestive
and is clothed with oesophageal epithelium. The second, the
abomasum, is the digestive region. The first part is again
divided into three compartments. In the Whales, on the other hand,
it is the digestive part which is again subdivided, while if the
first part is divided it is not markedly so as in the Euminants.

The lungs are remarkable for their unlobulated character ; in
this they agree with the lungs of the Sirenia. The thoracic
cavity in which they lie is barrel-shaped, and not, as is usual in
terrestrial mammals, boat -shaped, i.e. nairower sternally than
above. The alteration of the shape of the thoracic cavity is
associated with the aquatic life ; so at any rate the fact that it is
also marked in Seals and even in the Otter seems to show. The
Whales are also characterised by the great obliquity of the
diaphragm, which is extremely muscular. In this character
again we find an agreement with the Sirenia, and also with
other aquatic mammals ; it is not therefore a character of
Whales so much as evidence of an adaptation to the aquatic life.
The advantage is, it appears, in the increased capacity of the
thoracic cavity, and the consequent greater possibilities of expan-
sion of the lungs, which it must be remembered serve as
hydrostatic as well as breathing organs.

Some of the internal arteries of Whales break up into
retia mirabilia. Their kidneys are lobulated ; whether this has

ASYMMETRY OF SKULL

349

anything to do with the
aquatic life is not so clear.
It also characterises the
Sirenia, more or less, and
the Otters ; but, on the other
hand, the terrestrial Bears
show the same structure as
do also some Ungulates. It
must be borne in mind, too,
that the kidneys of foetal
Man are lobulated.

The liver is a compact
organ not showing such lobu-
lation as is common, but not
universal, among mammals.

The bones of Whales have
a somewhat loose structure,
,and are much impregnated $^
with oil. In many features
the skeleton of Whales is
highly distinctive of the
order.

The brain case is small
proportionately and rounded.
The " face " is therefore long,
and in some cases, especially
among the fossil forms of
Platanistidae, the rostrum
is extraordinarily elongated.
The asymmetry of the
Whale's skull is one of its
most remarkable features ;
this, however, is entirely
limited to the Toothed
Whales, and among them
is more pronounced in some
forms than in others. Thus
the Platanistidae and many
Ziphioids are not nearly so
asymmetrical as the Dol-

350

NASAL BONES

phins and, especially, Physcter. This asymmetry affects parti-
cularly the premaxillae, the maxillae, and the nasals. The
base of the skull is symmetrical. The Whale's skull has very
long premaxillae which, however, do not, except in the extinct

FIG. 184. — Under surface of the cranium of a young Ca.a'iiig Whale (Globicep/talus
melas). x £. AS, Alisphenoid ; BO, basioccipital ; cf, coudylar foramen ; £xO,
exoccipital ; Fr, supra-orbital process of frontal ; gf, gleuoid fossa of sqiiamosal ;
Ma, body of malar ; MX, maxilla ; OS, orbitosphenoid ; Per, posterior (mastoitl)
process of periotic ; PI, palatine ; PM;c, premaxilla ; Pt, pterygoid ; Sq, squamosal ;
tg, deep proove on squamosal for meatus auditorius externus, leading to tympanic
cavity ; Ty, tympanic ; Vo, vomer ; ZM, zygomatie process of malar. (From
Flower's Osteology.)

Zeuglodonts, bear any teeth. The nasal bones, whether sym-
metrical or the reverse, are very small in existing Whales,
which arrangement, together with the long and broad maxillary
bones, removes the anterior nostrils, the blow -hole, far back-
wards. The roof of the skull is not at all formed by the
parietals externally. These bones form a portion of the side
of the cranium, but are replaced or covered by the enormously-

FUSION OF CERVICAL VERTEBRAE

351

developed supra-occipital in the adult. Here again the Zeuglo-
donts are more typically Mammalian, for in them the parietals
have a normal development and situation, rising even into a
median crest as in so many quadrupeds. The hones related
to the organ of hearing, the tympanics and petrous hones, are
very solid and dense in structure. Moreover they are but

Pa

30

ExO

cd

ih

FIG. 185. — A section of a skull of a young Caa'ing Whale (Globicepnalus melas). x £. a,
Angle ; an, anterior nares ; AS, alisphenoid ; bh, basihyal ; BO, basioccipital ; BS,
basisphenoid ; cd, condyle ; cp, coroiioid process ; ExO, exoccipital ; Fr, frontal ;
id, inferior dental canal ; IP, interparietal ; ME, ossified portion of the meseth-
moid ; MX, maxilla ; Na, nasal ; Pa, parietal ; Per, periotic ; PI, palatine ; PMx,
prernaxilla ; pn, posterior nares ; PS, presphenoid ; Pt, pterygoid ; *, symphysis
of inaudible ; sh, stylohyal ; SO, supra-occipital ; Sq, squamosal ; th, thyrohyal ;
Vo, vomer. (From Flower's Osteology.)

loosely attached to surrounding bones, and are thus easily and
frequently lost. Nearly the only mammals which resemble the
Whales in the fact that the pterygoids sometimes meet in the
middle line below are the Edentata (Anteater and Armadillo, see
p. 167). But in both groups this peculiarity is not universal.

The vertebral column is remarkable for the fact that more or
fewer of the cervical vertebrae may be fused together into a short
and compact mass. This is seen at its maximum in the genera
Bcdaena, and Neobalaena. The odontoid process of the second

352

BREAST BONE

CHAP.

vertebra, though hardly at all marked, is nevertheless really
present and developed from a bony centre of its own, as in other
mammals. The dorsal and lumbar vertebrae are, of course, to be
distinguished by the presence of ribs attached to the former ; but

FIG. 186. — Section through middle
line of united cervical vertebrae
of Greenland Eight Whale (Bal-
aena mysticetus). x J. a, Arti-
cular surface for occipital condyle ;
e, epiphysis on posterior end of
body of seventh cervical vertebra ;
sn, foramen in arch of atlas for
first spinal nerve ; 1, arch of atlas ;
2, 3, 4, 5, 6, conjoined arches of
the axis and four following verte-
brae ; 7, arch of seventh vertebra.
(From Flower's Osteology.)

as there is only a rudimentary pelvis, not attached to the
vertebral column, no sacral region can be detected. The caudal
vertebrae are to be recognised by the V-shaped chevron bones
below.

FIG. 187. — A, Sternum of Greenland Right Whale (Balaena mysticetus). x
Sternum of Common Rorqual or Fin Whale (Balaenoptera musculus).
(From Flower's Osteology.)

B,

The sternum in the Whale tribe is much more modified in the
Whalebone Whales than in the Odontocetes. In the latter it is
made up of several pieces, as in other mammals, which often,
however, become coalesced. In the Mystacoceti this bone is a
single piece, to which only one pair of ribs is attached, and its

DIVISIONS OF CETACEA

353

form is characteristic of the genus. It is heart-shaped, more or
less, in Balaena, and somewhat cross- or T-shaped in the genus
Balaenoptera. In the Odontocetes the ribs have, some of them,
the normal attachment by capitulum and tuberculum. In the
Mystacocetes the at-
tachment, where it
exists, is very loose,
and the tuberculum
alone is attached to
its vertebra. This
allows of the freer play
of the ribs during re-
spiration. The scapula

has a very character- FlG" l88-~^ view of bones of 'posterior extremity of
* Greenland Right Whale (Balaena mysticetus). x £.

i, Iscbium ; /, femur ; t, accessory ossicle repre-
senting the tibia. (After Eschricht and Reinhardt)
(from Flower's Osteology.)

mion, where it exists,

is placed near the anterior margin of the shoulder blade, and
overlaps the generally long coracoid process. Clavicles are totally
absent. The pelvis is very rudimentary, consisting merely of a
single bonelet, to which are attached the rudiments (in some cases)
of a femur, and, in Balaena (Fig. 188), of a tibia also.

Whales are to be divided into three great groups: — (1) the
Whalebone Whales or Mystacoceti ; (2) the Toothed Whales or
Odontoceti ; and (3) the entirely -extinct Archaeoceti or Zeu-
glodonts.

.
istic form in these

SUB-ORDER 1. MYSTACOCETI.

This division is thus characterised : — Teeth are never function-
ally developed ; they are present in the young, but replaced in
the adult by the baleen or whalebone ; the external respiratory
aperture is double ; the skull is perfectly symmetrical ; the rami
of the mandible are arched outwards and do not form a true
symphysis ; the sternum is always composed of a single piece of
bone ; the ribs articulate only with the transverse processes of
the vertebrae.

The Mystacoceti are nearly invariably huge creatures, the
sole exceptions being the Pygmy Right Whale, Neobalaena, and

VOL. x 2 A

354

WHALEBONE

a small Korqual. But even these are larger than the majority
of Toothed Whales.

The most characteristic feature by which the Whalebone
Whales are to be distinguished from other Whales is that which
gives to them their name, the presence of whalebone. Whale-
bone is a horny product of the epithelium lining the mouth, and
is comparable to an exaggeration of the transverse ridges which

are found in the mouths of all
mammals upon the palate. In
rion - Cetacean mammals these
ridges vary in depth, and are
arranged as a rule transversely,
but with an oblique inclination.
This is precisely how the plates
of baleen are disposed in the
mouth of a Whale. Each piece
of " bone " is triangular in shape,
the broader end being that of
attachment while it narrows
gradually ; the inner side of the
blades is frayed out into a number
of threads which form the strain-
ing apparatus. The plates vary-
in length up to as great an ex-
treme length as 13 feet, which
occurs in the Eight Whale at
times. The colour is black or

FIG. 189. — Section of upper jaw, with

baleen plates, of Balaenoptera. a, paler, even white. Ihe number
Bone of jaw; b gum ; ; c, straight Qf thege plateg in the mouth is
edge of baleen plate ; a, e, frayed

out surface of baleen plates. (After very great. As many as 3/0

blades have been counted. They

diminish in length towards both ends of the series. Though
whalebone has been in use for a long period, whence the whale-
bone came was formerly one of those things not generally
known.

A very prevalent notion was that the whalebone formed the
eyelids or perhaps the eyelashes of the creature. Scaliger, com-
menting upon Aristotle, held that the whale had " lamellae upon
the eyebrows, which, when the head is plunged below the surface,
were raised by the water ; but when the animal raised its head

xii A STRANDED RORQUAL 355

above the waves the lamellae fell and covered the eyes." Whale-
bone, too, has been often spoken of as " the fin of a whale," " the
finnes that stand forth of their mouths." The value of whale-
bone is still great, in spite of various substitutes which are now
used in its place. In the year 1897, for example, the value of
this article was £2000 per ton. As a single Whale may produce
several tons of this material, it is not surprising to find that the
results of a whaling voyage may be very profitable.

Fam. 1. Balaenopteridae. — This genus Balaenoptera includes
the Rorquals, which are Whalebone Whales of large size, differing
from the Eight Whales in three important external characters :
the head is comparatively small ; there is a dorsal fin ; the throat
is marked by numerous longitudinal furrows. The bones of the
cranium are not so arched as in the Eight Whales, and as a
consequence the plates of baleeii are shorter. The hand is only
four-fingered. The cervical vertebrae are for the most part all
free. One of the earliest records of a Whale stranded in the
Thames was probably of a species of this genus in the year 1658,
and is thus described by. John Evelyn : — " A large whale was
taken betwixt my land butting on the Thames and Greenewich,
which drew an infinite concourse to see it, by water, horse, coach,
and on foot, from London and all parts. ... It was killed with
a harping yron, struck in the head, out of which spouted blood
and water by two tunnells, and after an horrid grone it ran
quite on shore and died. Its length was 58 foot, heighth 16 ;
black skinn'd like coach leather, very small eyes, greate taile,
onely two small films, a picked snout, and a mouth so wide that
divers men might have stood upright in it ; no teeth, but suck'd
slime onely as thro' a grate of that bone which we call whale-
bone, the throate yet so narrow as would not have admitted the
least of fishes ... all of it prodigious, but in nothing more
wonderful that an animal of so greate a bulk should be nourished
onely by slime thro' those grates."

Professor Collett has recently given1 an elaborate account of the
characters and habits of this great Whale (Balaenoptera musculus).
Though a large beast (44 to 67 feet in length) it is exceeded by
other Eorquals ; it is of a dark grey blue colour above, white, for
the most part, below. The dorsal fin is large and high ; the
flippers relatively slender and small. The whole throat from the

1 In Proc. Zool. Soc. 1886, p. 243.

356 RORQUALS CHAP.

symphysis of the jaws to the middle of the belly is, as in other
species, marked by furrows, forty to fifty-eight in number. The
hairy covering is reduced (in an adult female) to thirteen hairs
on each side of the lower jaw ; in a foetus there were also seven
hairs on each side of the upper jaw, as well as rather more on
the lower jaw — altogether, forty-eight. This Whale appears to
feed chiefly upon small Crustacea, especially the Copepod, Calanus
finmarchicus. The number of baleen plates is about 330 on each
side of the jaw. This Whale sometimes swims singly, but usually
in schools of even as many as fifty.

Rudolphi's Rorqual (B. lorealis) seems to be a perfectly
inoffensive beast ; it is said to be able to stay under water for
as long a time as twelve hours.

A smaller species than the last is B. rostrata — at the outside
33 feet in length. Here the hairy covering is reduced1 to " two
small hairs on the integument covering the apex of the lower
maxilla." The colour is greyish black above, the underside
white. On the other hand, B. sibbaldii, the Blue Whale, is the
giant of its race, reaching a length of 85 feet. Its colour is a
dark bluish grey, with small whitish spots on the breast. The
dorsal fin is small and low with straight margins.

B. musculus, the Finner, is intermediate in size — not more
than 70 feet. It seems doubtful whether the " sulphur bottom,"
B. australis, of Antarctica and B. pataclionica differ specifically
from this.2

The genus Megaptera is very near Balaenoptera, but differs
from it mainly in the following external and internal characters.
The dorsal fin is not very prominent, and its place is taken by a
lowish hump, whence, indeed, the common name of this Whale,
" Humpback." The pectoral fin is unusually long, and the
creature uses it to beat itself, the surrounding water, and, more
playfully, its mates. The general outline of this Cetacean is
more clumsy than that of Balaenoptera. The most important
internal difference is in the form of the scapula, which has at
most a slight acromion and coracoid process. These are rather
more pronounced, according to Messrs, van Beneden and Gervais,3

1 Perrin, "Notes on the Anatomy of B. rostrata," Proc. Zool. Soc. 1870, p. 805.

2 von Haast, "Notes on a Skeleton of Balaenoptera australis," Proc. Zool. Soc.
1883, p. 592.

3 Osteographie des Cttacte, Paris, 1880, p. 130.

xii THE CALIFORNIAN WHALE 357

in the southern form of the genus, which is known as M. lalandii.
The head, it should also be remarked, is studded with large
tubercles about the size of an orange, which seem to be hyper-
trophied rudiments of the hairs, which should be present in this
region of the body. As is the case with other Whales, numerous
species have been made out of individuals of Megaptera. Captain
Scammon, who observed many " gams " or herds of these Whales,
remarked x that he had extreme difficulty in finding any two
individuals precisely alike ! The best-known species in any case
is the northern M. longimana, which occurs on our own coasts.
The genus is, like so many Cetaceans, world-wide in range ; and
it is possible that the difference in the scapula already referred to
may justify the separation of a southern M. lalandii (with which
in that case, perhaps, M. capensis and M. novae zelandiae will be
synonymous). Quite recently M. Gervais has insisted upon a
Megaptera indica from the Persian Gulf. Megaptera grows to
a length of 50 to 60 feet. Seventy-five feet have been stated,
but measurements of Whales have usually to be received with
caution.

Ehackianectes, with but one species, -B. glaucus,2 the " Cali-
fornian Grey Whale," is the last genus of the family Balaenop-
teridae. This Whale is but imperfectly known anatomically ; but
quite sufficient has been ascertained to show its great divergence
from Balaenoptera or Megaptera. The dorsal fin is completely
absent, and the throat pleats, so characteristic of the typical
Balaenopteridae, are reduced to two. It has, however, the general
outline of a Eorqual, with a relatively small head. In osteo-
logical characters it tends to unite the two families Balaenop-
teridae and Balaenidae (if they are really necessary subdivisions).
The skull is on the whole Rorqual - like ; but its fore -part is
narrow as in the Greenland Whale, and the premaxillaries are
pinched up in the middle line so as to be visible from the side ;
this again is a Balaenid character. The cervical vertebrae are
free as in Eorquals, and the sternum is quite as in that group.
The scapula has more the shape of that of Balaena.

Rhachianectes glaucus is confined to the Pacific, and has been
extensively hunted from the shore. It is not, however, a very
valuable Whale, since the baleen is short as in Rorquals, and the

1 Marine Mammals of the Xorth-West Coast of North America, 1874.
2 Cf. Scammon, loc. cit.

358 SPECIES OF RIGHT WHALES CHAP.

beast, moreover, appears to be fierce, a somewhat rare attribute
of Whales. It has been spoken of, indeed, as " a cunning,
courageous, and vicious " animal. Rhacliianectes is essentially a
coast Whale, and loves to lie in the surf in quite shallow water
waiting for the tide to float it off. This Whale varies much in
colour from black to mottled grey and black, and reaches a length
of about 40 feet.

Fam. 2. Balaenidae. — The Eight Whales of the genus
Balaena are to be distinguished from Neolmlaena and from
the Eorquals by the following characters : —

The size is large, 50 to 60 feet. There is no dorsal fin. The
head is more than or nearly one-fourth of the entire length of
the animal. The baleen is very long. The throat is not grooved.
The orbital process of the frontal is not wider than the down-
ward process of the maxilla. The cervical vertebrae are all
fused. The scapula is rather high. The hind- limb has the
rudiment of a tibia. The intestine has no caecum.

A vast number of different genera have been founded on
detached bones, bits of whalebone, and more or less complete
skeletons of Eight Whales coming from different parts of the
world. In Dr. Gray's catalogues we find the following allowed,
viz. Balaena, Eubalaena, Hunterius, Caperea, Macleayius. The
number of " species " distributed among the genera is some
thirteen or more, with whose names we shall not trouble the
reader. As a matter of fact there are not more than two species
which can with certainty be identified and distinguished, both of
which are so close that they cannot possibly be placed but in
the same genus, Balaena. In no group of Whales — in no group
of animals probably — has imagination run riot to so terrible an
extent in the formation of genera and species as in these Eight
Whales. This multiplication or rather division of genera has
arisen from an old idea that Whales coming from different seas •
must be of different kinds, a notion now thoroughly exploded.

The term " Eight Whale " simply means that the Whales of
this genus are the right kind of Whale for the whaler to pursue.
Their whalebone is longer and more valuable, while the oil is
not only more abundant but of a superior quality. The two
species demand a separate account.

The Greenland Whale, Balaena mysticetus, is one of the rare
instances of a Whale which has an exceedingly limited range in

GREENLAND WHALE 359

space. It is absolutely confined to the Arctic Ocean, and reported
occurrences on our coasts are due to a confusion with B. australis,
to be presently described. At the " Devil's Dyke," near Brighton,
there is, or was, the skull of a most flagrant Eorqual, which is
carefully labelled " Greenland Whale." This Whale grows to a
length of 50, 60, rarely 70 feet. It is black in colour, save
for a white patch on the under side of the jaw. The head is
quite one-third of the body in length. There are a few scattered
hairs at the extremity of the jaws. The length of time which
this Whale can endure immersion has been variously stated. The
utmost limit of endurance is stated by Scammon to be one hour
and twenty minutes. The pursuit of this Whale is attended by
dangers, not in the least because the animal is itself fierce and
ready to attack, but simply on account of the velocity with
which, and the great depth to which, it will dive, and also to the
huge muscular force which is exerted in its struggles to free itself
from the harpoons. It is indeed an extremely timid beast. It
has been remarked that "a bird alighting upon its back some-
times sets it off in great agitation and terror." Combined with
this timidity of disposition is an intense affection for its young,
" which would do honour," observed Scoresby, " to the superior
intelligence of human beings." Yet that trader and observer
goes on to remark that " the value of the prize . . . cannot be
sacrificed to feelings of compassion " ! The fact that this Whale
and its congener, B. australis, feed among swarms of minute
pelagic creatures, which they engulf in their huge mouths, led
the ancients to believe and assert that they fed upon water only.
When the Whale feeds it moves along with some velocity, taking
in huge mouthfuls of sea water with the contained organisms,
which are then strained off by the whalebone and left stranded
upon the tongue.

Unlike its congener, the southern Eight Whale, B. australis,1
is world-wide in distribution, avoiding only the Arctic regions.
Where the Greenland Whale is found B. australis does not exist.
The principal differences which it shows from B. mysticetus are
firstly in the relatively shorter head and shorter and coarser
whalebone. In the second place it has more ribs, fifteen pairs
as against thirteen ; but there is apparently some little confusion
in the matter of ribs. An additional rib at the end of the series

1 The name that has priority seems to be glacialis.

360 WHALING IN THE BAY OF BISCAY CHAP.

is apt to get lost, and in the skeleton of so huge and unmanage-
able a beast there is nothing more unwise than to insist upon, as
specific characters, what may be due merely to defective prepara-
tion. This Whale has often, and the Greenland Whale also, a
rough horny protuberance upon the snout known as the " bonnet."
The causation of this is not clear. It has been spoken of as " a
rudimentary frontal horn." But this suggestion of an Ungulate
affinity can hardly be accepted. It seems to be more like a kind
of corn.

This Whale was once more abundant on the coasts of Europe
than it is to-day ; it was much hunted by the Basques in past
time. The Whale which frequented the Bay of Biscay was usually
called the Biscayan Whale or B. Biscay ensis ; but there is prob-
ably no specific difference. Among the small towns which fringe
the Bay, it is very common to find the Whale incorporated into
the armorial bearings. " Over the portal of the first old house
in the steep street of Gruetaria," writes Sir Clements Markham,1
" there is a shield of arms consisting of Whales amid waves of
the sea. At Motrico the town arms consist of a Whale in the
sea harpooned, and with a boat with men holding the line."
Plenty of other such examples testify to the prevalence of the
whaling industry on these adjoining coasts of Spain and France.
It appears that though the fishery began much earlier — even in
the ninth century — the first actual document relating to it dates
from the year 1150. It is in the shape of privileges granted
by Sancho the Wise to the city of San Sebastian. The trade
was still very flourishing in the sixteenth century. Eondeletius
the naturalist described Bayonne as the centre of the trade, and
tells us that the flesh, especially of the tongue, was exposed for
sale as food in the markets.

M. Fischer,2 who, as well as Sir Clements Markham, has
given an important account of the whaling industry on the
Basque shores, quotes an account of the methods pursued in the
sixteenth century. It was at Biarritz — or as Ambroise Pare,
from whom Fischer quotes, spelt it, Biaris — that the main
fisheries were undertaken. The inhabitants set upon a hill a
tower from which they could see " the Balaines which pass, and
perceiving them coming partly by the loud noise they make, and

1 Proe. Zool. Soc. 1881, p. 969.
2 Actes Linn. Soc. Bordeaux, 1881.

xii NEOBALAENA 361

partly by the water which they throw out by a conduit which
they possess in the middle of the forehead." Several boats then
set out in pursuit, some of which were reserved for men whose
sole duty it was to pick out of the water their comrades who
had overbalanced themselves in their excitement. The harpoons
bore a mark by which their respective owners could recognise
them, and the carcase of the animal was shared in accordance
with the numbers and owners of the harpoons found sticking in
the dead body of the Whale. At this period the fishery was at
its height. But it continued to be an occupation along those
shores until the beginning of the eighteenth century, after which
it gradually declined. The fishery of Whales began to be carried
farther afield than the shore, and for a long time the Basques
furnished expert harpooners to whaling vessels proceeding to the
Arctic seas. A curious example of the continuance of the fishery
until at least 1712 is given by Sir C. Markham. In the parish
records of Lequeito for that year, it is noted that a couple were
married who possessed between them all the necessary outfit for
a whaling cruise.

The genus Neobcdaena is interesting from more than one
point of view. Its size compared with its gigantic relatives is
small, some 16 or 17 feet. The genus bears the same kind of
proportion to Balaena that Kogia does to Physeter among the
Physeteridae. It is one of those Whales which are very restricted
in habitat ; up to the present it is only known from the Antarctic
region in the neighbourhood of New Zealand and South Australia.
Structurally it is in a few points intermediate between the Eight
Whales and the Eorquals. The head is proportionately (as well
as, of course, actually) not so large as in Balaena. There is a
falcate dorsal fin ; but the head in outline is not Eorqual-like in
spite of its similar proportions. The whalebone is long. The
throat is not grooved. Neolalaena has forty-three vertebrae, of
which the cervicals are all fused. There are as many as seventeen
or eighteen dorsal vertebrae, the largest number in any Cetacean
as far as is known. With these are articulated not eighteen but
only seventeen ribs. The first dorsal vertebra appears to be with-
out a rib. The ribs are very broad and flat. The body thus
gets an appearance of a Sirenian. The lumbar vertebrae are
fewer than in any other Cetacean, being only two. The scapula
is more like that of the Eorquals than that of the Eight Whales ;

362 TOOTHED WHALES

that is to say, it is long and not very high. The skull is most
like that of Bcdaena, but the process of the frontal arching over
the eye is broader relatively than in Balaena, and thus approaches
Balaenoptera. Nothing is known of the viscera of this Whale.
The whalebone is white, and the animal was first described by
Dr. Gray from pieces of "bone." It is not always that so
fortunate a diagnosis of specific or generic difference has been
made from a structure which apparently offers so little aid for
discrimination.

There is but a single species of the genus which is named
Neobalaena maryinata.1

SUB-ORDER 2. ODONTOCETI.

The Odontoceti have teeth but no whalebone ; the blow-hole
is single ; the skull is not symmetrical ; some of the ribs are
two-headed.

Fam. 1. Physeteridae. — This family of the Odontocetes may
be thus defined : — All or most of the cervical vertebrae are fused
together. The costal cartilages are not ossified. In the skull the
pterygoids are thick and meet in the middle line ; the sym-
physis of the mandible is long. Teeth, more or fewer, are found
in both jaws, but those of the mandible are alone functional
(?exc. Kogia}. The pectoral limb is smallish. The throat is
grooved by two or four furrows.

This family of Whales is again susceptible of division into
the two sub-families — Physeterinae or Sperm Whales and the
Ziphiinae or Beaked Whales. Professor P. J. van Beneden was
strongly against any subdivision of what is here regarded as a
perfectly natural family, embracing the Physeters and the Beaked
Whales. There are, however, some reasons for the subdivision. The
Ziphiinae have a reduced series of teeth, never exceeding two on
each mandible, which contrasts with the fully-toothed mandibles
of both Physeter and Kogia. The stomach of the Ziphioids is
extraordinarily complicated even for a Cetacean. The small
head of the latter group, which recalls in a curious way that of
Mosasauroid reptiles and some Dinosaurs, is in contrast to the

1 For osteology see Hector, Trans. New Zeal. Inst. %-ii. 1876, p. 251 ; and
Beddard, Trans. Zool. Soc. xv. 1901, ix 87.

SPERM WHALES 363

enormous head of the Cachalot and the very fairly -developed
skull of the "Pygmy Sperm Whale." Both, however, furnish
spermaceti, and in various osteological details come near together.
On the whole we incline towards separating the Cachalots from
the Ziphioids, and shall therefore commence with the former as
being in some respects the more primitive members of the family
Physeteridae.

Sub-Fam. 1. Physeterinae. — This sub-family may be thus
defined : — Teeth in lower jaw numerous. Xo distinct lachrymal
bone. Stomach with only four compartments (? as to Kogia*).

Of this sub-family the best-known genus is Physeter, including
the Sperm Whale or Cachalot. Of other reputed species we shall
speak later. The genus is characterised in the first place by its
large size — as much as 82 feet of length have been assigned to
Physeter macrocephalus ; but Sir William Flower thought that
55 or possibly 60 feet might be a better approximation to the
greatest length of the Cachalot. The head is enormous, a third
of the length of the body, and terminates in a massive and
bluntish snout. This is, however, not so abruptly truncated as
is often represented in figures. According to Messrs. Pouchet
and Chaves,1 it slopes forward two metres beyond the end of the
lower jaw ; the mouth is thus ventral and almost shark-like in
position, as is the case also with the Pygmy Sperm Whale, to be
considered later. In connexion with this peculiar position of
the mouth, it has been asserted — Mr. F. T. Bullen figures it2-
that the Sperm Whale turns over upon its back to bite. The
blow-hole is single, and shaped like the sound-rhole of a violin ;
it lies upon one side, and is not median in position. The throat
is grooved as in the Ziphioids by two grooves. The dorsal fin is
represented by a whole series of lowish humps, decreasing in
elevation from before backwards. The pectoral fins are not large
relatively speaking. The great square head is not occupied
entirely by the skull ; the cavity lying above, which is of course
traversed by the tube ending in the blow-hole, is filled with the
spermaceti, which is fluid fat during the life of the animal.
Spermaceti also occurs in other Whales ; and that of Hyperoodon,
whence it has been extracted for commercial purposes, is said to
offer no differences of importance from the spermaceti of the

1 Journ. de I'Anat. xxvi. 1890, p. 270.
2 The Cruise of the Cachalot, London, 1900.

..

364 SPERMACETI AND AMBERGRIS CHAP.

Sperm Whale. Spermaceti as a drug appears to have been first
mentioned in the pharmacopoeias of the famous medical school of
Salerno towards the year 1100. But it was confounded with a
totally distinct substance, viz. ambergris. The confusion was also
made by the famous alchemist Albertus Magnus, and by the
observant Archbishop of Upsala, Olaus Magnus, in his work
De gentibus septentrionalibus. It was supposed in fact by these
writers to be the liberated sperm of the Whale, hence obviously
the name. Later on, the substance in question was regarded as the
brain of the Cachalot, in fact as late as the middle of the eighteenth
century. It was Hunter and Camper who really discovered the
true nature of the substance, oil of course, in the cavities of the
skull.1 The huge skull of Physeter " is perhaps the most modified
from the ordinary type " of skull in the whole mammalian class.
The top of the skull rises into a huge crest lying transversely,
and from it slope forward two lateral crests formed from the
maxillary bones ; in this great basin lies the spermaceti already
referred to. The skull, as in Toothed Whales generally, is ex-

O v *

ceedingly asymmetrical. The right premaxillary and the left
nasal bones are much larger than their fellows ; indeed the right
nasal is hardly present as a separate bone. The parietal if pre-
sent is fused with the supra-occipital. The jugal is large, and is
not divided into two pieces as it is in the Ziphioids. The ptery-
goids meet below for a considerable distance, as in many Dolphins,
and in the Edentata among other mammals. The symphysis of
the lower jaw is very long, but the bones do not appear to be
ankylosed. The length of the symphysis recalls that of the
Gangetic Dolphin, Platanista.

In the vertebral column the atlas alone is free, the remain-
ing cervicals being fused. There are only eleven dorsal vertebrae,
eight lumbars, and twenty-four caudals. The breastbone of this
Whale is a roughly-triangular bone made up of three pieces.
Four cartilaginous sternal ribs are attached to this bone. The
scapula is remarkable for the fact that it is concave on the outer
and convex on the inner surface ; otherwise it is quite typically
Cetacean in form. The shortness of the pectoral limb is shown
by the phalangeal formula, which is as follows : — I 1, II 5,
III 5, IV 4, V 3.

1 See Pouchet, "Contribution a 1'histoire du spermaceti,' Bergens Museums
Aarbogfor 1893, No. 1.

xii FEROCITY OF THE CACHALOT 365

One of the reasons for the pursuit of the Sperm Whale is
the desire to obtain that extremely valuable product ambergris.
This substance has long been known ; but its true nature was
for centuries in dispute. In Dr. Johnson's Dictionary (so
recently as the edition of 1818 !) ambergris is provided with
alternative definitions : it is either the excrement of birds washed
off rocks, or honeycombs that have fallen into the sea !

An old writer asserted of ambergris that it was " not the scum
or excrement of the whale, but issues out of the root of a tree,
which tree, howsoever it stands on the land, alwaies shoots forth
its roots towards the sea, seaking the warmth of it, thereby to
deliver the fattest gum that corns out of it, which tree other-
wise by its copious fatness might be burnt and destroyed."
These " explanations " were caused by the fact that ambergris is
sometimes found floating in the sea. Ambergris is, of course, a
product of the intestinal canal of the Sperm Whale ; it seems to
be of the nature of cholesterin, and its place of origin was con-
clusively proved by finding the beaks of cuttle-fish imbedded in
it. When first extracted from the alimentary canal it is of
greasy feel and consistency ; later it hardens, and acquires its
characteristic sweet earthy odour. Ambergris is used mainly as
a vehicle for scents, and is a costly substance. A piece weighing
130 Ibs. was valued at £500. Though now entirely used in
connexion with perfumery, it was held by the ancients to be of
great value as a specific in certain diseases.

The Sperm Whale is chiefly a tropical animal. Examples
that have been cast up on our shores are strayed individuals. It
often goes about in herds, which seem to be composed of females.
Its food is chiefly cuttle-fishes, and it is said to have a pre-
dilection for those colossal cuttle-fishes whose existence has
until recently been doubted. Mr. Bullen has sketched a conflict
between these two giants of the deep. On the other hand it is
said that its large throat, more than big enougli to swallow a man
(the Whale is credited with being that which swallowed Jonah),
does not usually admit fishes larger than Bouitos and Albacores.

The ferocity of the Cachalot has been denied and affirmed.
It certainly has great strength, for it am throw itself com-
pletely out of the water. Captain Scammon thinks that ships
which are mysteriously lost at sea, with no obviously assignable
cause, are sometimes the victims of the furious rushes of a bull

366 THE IIIGH-FINNED CACHALOT

Sperm Whale. Marco Polo took much the same view, but
suggested that the Whale did not deliberately attack the ship,
but was deceived by the foam following in its wake into thinking
" there is something to eat afloat, and makes a rush forward,
whereby it shall often stave in some part of the ship." ]

Sir W. Flower and many others are of opinion that there is
but one species of Cachalot. But many names have been given
to supposed other forms. The genus itself has even been
divided, and to a set of vertebrae from the south Dr. Gray gave
the perfectly superfluous name of Meganeuron kreffti. The " High-
finned Cachalot " rests mainly upon the suggestions of Sir Efobert
Sibbald. It is supposed to have a high dorsal fin, and teeth in
the upper as well as in the lower jaw. Common though it was
asserted by its describer to be, there is not a bone, not a fragment
even of a bone, alleged to belong to Physeter tursio in any
museum in the world ! It seems premature, therefore, to include
this mysterious creature in any list of Cetacea, though that was
done by no less a naturalist than the late Mr. Thomas Bell. It
is this creature round which most of the stories of ferocity con-
gregate. It is held to be the monster from which Perseus
delivered Andromeda, and which was about to devour Angelica
upon the shore of Brittany. The fact of the matter is, that the
Sperm Whale, like so very many other Whales, is world-wide in
range ; and those naturalists who did not believe in so wide a
distribution found themselves obliged, in order to satisfy their
own views, to create new species for those of distant localities.
Hence the dozen or so of synonyms which refer to what is to be
called Physeter macrocephalus.

The genus Kogia (sometimes written Cogia), the so-called
" Pygmy Sperm Whale," is a southern form of much smaller
dimensions than its gigantic ally just described. Kogia does
not exceed 15 feet or so in length. It differs from Physeter
also in the well-marked and falcate dorsal fin, in its generally
delphinoid form, in the short snout, and the more normal (for a
Whale) shape of the blow-hole, which is crescentic.

There are also a number of osteological characters in which
the two Physeterines differ from each other. In Kogia all the
cervical vertebrae are ankylosed together ; the skull is short,
though equally asymmetrical ; the ribs are as many as twelve or

1 Yule, Travels of Marco Polo, ii. London, 1874, p. 231.

xii BEAKED WHALES 367

fourteen ; the scapula has not the concave face that it has in
Physeter. The functional teeth of the lower jaw seem to be
reinforced by two on each side of the upper jaw. Moreover, the
articulation of the ribs with the vertebrae does not show the
very anomalous state of affairs that characterises Physeter, where
the two heads of a rib may be upon one vertebra.

While there is no doubt as to the generic distinctness of
Kogia, there is again the same difficulty that is met with
throughout the whole of the order in settling into how many
species the genus requires dividing.

We can dismiss, as unnecessary, additional generic names
(Euphysetes, Callignathus), but there do appear to be reasons fdr
allowing two species, if the accounts of their osteology are to be
depended upon. One of these is K. breviceps, with thirteen pairs
of ribs, no teeth in the upper jaw, fourteen or fifteen on each side
of the lower jaw, vertebral formula C 7, D 13, L 9, Ca 25, and
phalangeal formula I 2, II 8, III 8, IV 8, V 7.

The other will then be K. simus, with fourteen pairs of ribs,
two teeth in the upper jaw, nine in each ramus of the lower jaw,
vertebral formula C 7, D 14, L 5, Ca 24, and phalangeal formula
I 2, II 5, III 4, IV 4, V 2.

A Californian species has been called K. floweri, whose teeth
seem to be particularly long and recurved. And the New Zealand
K. pottsi has been held to be also a distinct form. There seems
to be nothing of special interest to record about the way of life
of these Cetaceans, which are but imperfectly known.

Sub-Fam. 2. Ziphiinae. — Teeth in the lower jaw not more
than two on each side. A distinct lachrymal bone. Stomach
with very numerous compartments.

These Whales are all of moderate size, not exceeding 30 feet
or so in length. They have a falcate dorsal fin rather near the
end of the body ; the muzzle is prolonged, hence the name often
given to them of " Beaked Whales." The throat is grooved ; the
blow -hole is single and median, crescentic in form, with the
concavity pointing forwards. A character possibly differentiat-
ing the Ziphioids from other Whales is the fact that the body
ends in a rounded projection between the flukes of the tail.
This has at any rate been noted in Mesoplodon, Ziphius, and
Hyperoodon. The Ziphioid Whales are by no means common ;
indeed of Berardius but four or five specimens have ever been

368 WHALES AND " SEA-SERPENTS " CHAP.

met with. Most of them are southern in range, and the
vast stretches of desolate coast which occur in these regions of
the world account possibly for the rarity of their remains.
These Whales have done duty more than once for the " Sea
Serpent." Quite recently an alleged sea serpent turned out to
be a couple of Mesoplodon lying head to tail ! The head in
these Whales is small compared to the body. The skull is
characterised by the strong maxillary crests, enormously deve-
loped in the male Hyperoodon. The vertex of the skull too is
raised, forming a pronounced prominence behind the aperture of
the nares (blow-hole) ; in many forms the rostrum is made of
very dense bone, and is thus relatively abundant in rock strata.
The pterygoids meet in the middle line as in the Cachalot.
In addition to the few functional teeth in the lower jaw there
are more numerous but small teeth in the upper jaw. These
are not always to be recognised, as they are not attached to the
bone, but merely imbedded in the gums, so that they come away
when the skull is prepared.

The genus Berardius l differs from Mesoplodon by its rather
more symmetrical skull, of which the vertex is formed by the
nasals. The mesethmoid is only partly ossified. The teeth
are two on each side of the mandible, with their apices directed
forwards. The vertebral formula is C 7, D 10, L 12, Ca 19.

B. arnouxi, from the seas of New Zealand, is the only species
of this genus which is well known. It is 30 or 32 feet in
length, and is of a velvety black colour, with a greyish belly.
Instead of lowing like a cow, this Whale has been described
as " bellowing like a bull " ! A singular and somewhat inex-
plicable fact has been stated of this species. The teeth were
said to be protrusible, and Sir James Hector stated that the
teeth were imbedded " in a tough cartilaginous sac which
adheres loosely in the socket of the jaw, and is moved by a
series of muscular bundles that elevate or depress it." Sir
William Flower justly observed that these statements " accord so
little with anything hitherto known in mammalian anatomy
that further observations on the subject are extremely desir-
able." Like other Ziphioids, Berardius feeds mainly, if n6t
entirely, upon cuttle-fish, a prey eminently suited to their almost/
toothless mouths. It is not known whether Berardius has the

1 See Flower, Trans. Zool. Soc. viii. 1872, p. 203.

xii VOICE OF MESOPLODON 369

Ziphioid grooves upon the throat. Nothing is known of the
structure of the internal viscera of this Whale. It appears not
to be really limited to the region of New Zealand, as is often
stated, for Malm has lately described a skull (JSerardius vegae}
from Bering's Straits.1

Mesoplodon2 is a world-wide genus embracing a number of
species ; on the lowest estimate seven species can be distinguished,
and Sir W. Flower would add two more. These are moderate-
sized Whales, 1 5 to 1 7 feet in length. In the skull the mesethmoid
is ossified ; the nasals are sunk between the upper ends of the
premaxillae. There are but a single pair of teeth in the mandible
attached to nearly the middle of its length (whence the generic
name). The vertebral formula is C 7, D 9 or 10, L 10 or 11,
Ca 19 or 20. The sternum consists of four or five pieces. The
amount to which the cervical vertebrae are fused varies ; but
some are always fused.

The only species which has ever been stranded on the shores
of this country is M. Widens, an example of which was described
many years ago as the " Toothless Whale of Havre " ; it was an
old animal which had probably lost its teeth. Nevertheless it
received the separate generic and specific name of Aodon dalei.
The animal lived for two days out of the water, and made a sound
like the " lowing of a cow." An instance of the rarity of the
Whales of this genus is afforded by M. europaeus, of which only a
single skull is known ; this was extracted from a dead body, found
floating, about the year 1840. It has never appeared since.
M. layardi is remarkable on account of the very large size of its
strap-shaped teeth ; these curve over the upper jaw in such a
way as to prevent the animal from fully opening its jaws. The
case is curiously paralleled by the Sabre-toothed Tiger. This
. species is antarctic in range. From the opposite extremity of the
globe comes M. stejnegeri, again known by but a single skull. It
is singular on account of the large size of the brain case, and is -a
native of Bering's Straits. M. hectori has its two teeth situated
quite at the extremity of the mandible, and in this feature
approximates to the genus Berardius. It was, indeed, confounded
with that genus by one naturalist.

1 Bihang Svensk. Akad. Handl. viii. 1883.

2 Flower, Trans. Zool. Soc. x. 1878, p. 415 ; and H. 0. Forbes, Proc. Zool. Soc,
1893, p. 216.

VOL. X 2 B

37O ZIPHIUS CHAP.

Ziphius is a genus which is also of world-wide range. Here
again the number of species is at present merely a matter of
opinion. The prevalent impression, however, is that but a
single species exists, which will therefore have the name of
Z. cavirostris. The genus (and for the matter of that the species
too) may be thus characterised in comparison with its allies.
The mesethmoid is ossified as in Mesoplodon, but the nasals joined
together form the vertex of the skull. There are two teeth near
the symphysis of the mandible, besides the usual small and
" functionless " teeth in the upper jaw. The vertebral formula is
C 7, D 9 or 10, L 11, Ca 21.

The throat of a Ziphius from New Zealand was described by
Messrs. Scott and Parker l as having three grooves on each side.
Whether this form is the same as von Haast's Z. novae zelandiae
is a matter of doubt ; but the individual to which his name has
been applied was 26 feet long, and had but a single groove on
each side. Even in the external characters of many Whales many
points require clearing up. Our knowledge of Ziphius dates from
the year 1804, when a skull " completely petrified in appearance "
was picked up upon the Mediterranean coast of France, and
described by the great Cuvier. It was forty years before another
specimen was found. In the New Zealand specimen of von
Haast already referred to, the body was scored by numerous
lacerations. These wounds may have been due to fights among
the Whales themselves ; the for wardly- situated teeth would be
capable of inflicting such wounds. But it has also been stated
that the armed suckers of gigantic cuttle-fish are responsible for
these scratches.

Hyperoodon is the most easily - distinguishable genus of
Ziphioid Whales. Its characters are the following : — The skull
has enormously-developed maxillary crests in the adult male ; the
mesethmoid is not fully ossified. There is but a single tooth to
each ramus of the lower jaw, besides, of course, the usual small
teeth in the upper jaw. The vertebral formula is C 7, D 9, L 9,
Ca 18. The cervicals are fused into one mass, more or fewer
being free in other Ziphioids. The sternum consists of three
pieces only, the last of which is bifid posteriorly.

The name Hyperoodon was given to this Whale by Colonel
Lacepede on account of the rough papillae upon the palate, which

1 Trans. Zool. Sot. xii. 1889, p. 241.

xii RANGE OF HYPEROODON 371

were mistaken by that observer for teeth. It is curious that the
name is really appropriate in spite of this mistake, though of
course it would be so to all the Ziphioids. In more than one
feature this genus comes nearest of all the Ziphiinae to Physeter.
Its enormous maxillary crests are paralleled in that Whale ; but
in Jfyperoodon their great thickness contrasts with the thinness
of those of the Cachalot. The correspondence in the attachment
of a rib to its vertebra by both heads is noteworthy. It is
remarkable that in this particular Hyperoodon is more like
Physeter than the supposed nearest ally of the latter — Kogia,

Of this genus two species are known. The best known is the
common northern H. rostratum (with many aliases) ; the second
species from the southern hemisphere, If. planifrons, is only
known from a single water- and pebble-worn skull. Its identifica-
tion, however, depends upon the known accuracy of the late Sir
William Flower.

The northern species (Jfyperoodon rostratuni) has often been
recorded upon our own coasts ; the first record of the stranding of
this Whale was in the year 1717. In that year an example was
found at Maldon, in Essex. Like the Beluga, Hyperoodon rostratum
gets lighter in colour with advancing years. The young are
black ; the old animals pale brown with some white about them.
The under surface, however, is always greyish white. The length
of this Whale reaches to at any rate 30 feet. But John Hunter
had a specimen which he believed to be 40 feet in length.
The specimen, however, consisted only of a skull, so that error
might have crept in. It has already been mentioned that the old
males have enormous maxillary crests. According to M. Bouvier,
who has lately made an exhaustive examination of the anatomy
of this Whale,1 the females occasionally exhibit the same crests,
which are thus presumably of the nature of spurs sometimes seen
in old females among the Gallinaceous birds. The number of
grooves upon the throat is in dispute in this Whale as in Ziphius.
One pair is the usual allowance ; but Kiikenthal found four in
some embryos studied by him. Attention has already been called
to the voice of Ziphioid Whales. Hyperoodon neither " lows " nor
" bellows," but " sobs " ! Hyperoodon rostratum is a gregarious
Whale, going about in herds, or " gams " as they should techni-
cally be termed, of four to ten or even fifteen. This Whale

1 Ann. Sci. Nat. (7), xiii. 1892, p. 259.

3/2 DOLPHINS

can leap right out of the water, and while in the air can turn its
head from side to side, a capability which has not been mentioned
in any other Whale. It can also stay under water for an unusually
long period. Captain Gray,1 who has made an accurate study of
this species, states that so long a period as two hours is the limit of
endurance ; this event occurred in the case of a harpooned Whale.

Fam. 2. Delphinidae. — This family, which includes the
greater number of Cetacea, may thus be characterised : — Whales
of small to moderate size. Teeth as a rule numerous, and
present in the upper as well as in the lower jaw. Maxillae
without large crests ; the pterygoids, often meeting in the middle
line, enclose an air space open behind. The anterior (five to eight)
ribs are two headed, the posterior with tubercular head only. The
sternal ribs are ossified.

The Dolphins and Porpoises, as already stated, embrace the
greater number of existing species of Whales. Sir W. Flower and
others who have followed him, allow nineteen genera. But as to
the exact number of known species there is much uncertainty.
That very careful observer, Mr. True, considers2 that there are
fifty which demand recognition. As many as one hundred have
received names. The matter is one which is perhaps barely ripe
for decision. All the Dolphin tribe are, for Whales, smallish
animals. The Killer Whale, Orca, is the only genus (or species ?)
which usually attains to more than moderate bulk. The rather
mysterious Delphinus coronatus, 36 feet in length, of M. de Fre-
minville, would seem to be a Ziphioid ; it was described as having
a very pointed beak, and as having the dorsal fin -situated near
the tail; such characters suggest a Mesoplodon.

The genus Delphinapterus, the Beluga or White Whale, con-
sists of but a single species, though as usual more than one
name has been given to supposed different species. It is char-
acterised as a genus by the following assemblage of structural
features : — It has only eight to ten teeth occupying the anterior
part of the jaws only. All the cervical vertebrae are free and
unjoined. The vertebral formula is C 7, D 11 (or 12), L 9,
Ca 23. The pterygoids are wide apart, though they converge
as if about to meet at their posterior ends. There is no dorsal
fin. The colour is white.

1 Proc. Zool. Soc. 1882, pp. 722, 726.
2 Bull. U.S. Nat. Mus. No. 36, 1889, p. 7.

xii THE BELUGA AND NARWHAL 373

The Beluga is a northern species purely. The reputed form,
D. kingii, was said to come from Australian seas; but there
seems to have been an error in this statement. It is interesting
to note that the white colour, so characteristic of the genus
and species, is not found in the young, which are blackish. They
gradually pale as they advance towards maturity. Delphin-
apterus leucas reaches a length of 10 feet, and like other
Porpoises will ascend rivers in search of food. It is said to
be specially addicted to salmon. Among the contents of the
stomach have been found quantities of sand. But this habit of
swallowing sand or pebbles has been noted in other Whales.
Whether it is or is not accidental (taken in with ground-living
food), it seems hardly likely that it is used for purposes of
ballast ! The Beluga has a voice ; but the name " Sea Canary "
is hardly suitable to it. A specimen of this species, recently
described from the shores of Scotland (it is often thrown up
upon our coasts), which had got entangled in the stakes of a new
net, was regarded by the natives, on account of its white colour,
as a ghost. Externally, besides its colour, the Beluga is remark-
able for possessing a distinct neck, which is correlated of course
with the freedom of the cervical vertebrae, and is also seen in
Platanistidae.

The Narwhal (Monodon) is closely allied in structure to the
last genus. It has the following . anatomical characters : — The
teeth are reduced to a single " horn " in the upper jaw, which
is rudimentary in the female. The neck vertebrae are free. The
vertebral formula is C 7, D 11, L 6, Ca 26. The pterygoids
are as in Delphinapterus, and, as in that genus, there are no hairs
upon the face or dorsal fin.

This genus is of course most obviously characterised by the
twisted tusk of the male, which is occasionally double. This
tusk has given to the only species of the genus, M. monoceros,
both its generic and specific name. The animal has a spotted
colour ; but, as in the case of the Beluga, old animals tend to
become white. The use of its horn to Monodon has been
debated. In the first place it is clearly a secondary sexual
character. The males have been observed to cross their horns
like rapiers in a fencing match. It may be that they are used in
more serious combats. An ingenious suggestion is that the long
and strong tusk enables its possessor to break the thick ice and

3/4 THE PORPOISE

make a breathing hole. A third suggestion is due to Scoresby,
who was led to make it from having taken out of the stomach of
a Narwhal a large skate. He held that with its tusk the Whale
empaled the fish and then swallowed it. The Narwhal is not
large, 15 feet or so in length. But Lacepede, who was apt to
compile with lack of discrimination, speaks of 60 feet long
Narwhals. Monodon is purely Arctic, and but three or four
specimens have ever been cast up on our shores.

Of true Porpoises, genus Pkocaena, there are apparently
several species. The genus itself has the following characters : —
The teeth are sixteen to twenty-six on each half of each jaw ;
their crowns are compressed and lobed. The pterygoids do not
meet. The dorsal fin has a row of tubercles along its margin.

The Porpoise of our coasts, P. communis, is a smallish species
6 to 8 feet in length. There are two to four hairs present in
the young ; its colour is black, generally lighter on the belly.
The first six cervical vertebrae are fused. The ribs vary in
number from twelve to fourteen pairs. It is a gregarious Whale,
and will ascend rivers ; it has been seen for example in the
Seine at Paris. The name Porpoise is often written Porkpisce,
which of course shows its origin. Very . conveniently it was
regarded as a fish, and therefore allowed to be eaten in Lent.
The celebrated Dr. Caius, a gourmet as well as a physician and
the refounder of a college, invented a particular sauce wherewith
to dress this royal dish. Some time since Dr. Gray described a
Porpoise from Margate as a distinct species (see p. 342) on account
of the tubercles, which are now known to be a generic character.

Dr. Burmeister's P. spinipennis seems, however, to be really
distinct. It was captured near the mouth of the Kio de la Plata.
It is more tuberculated on the fin and back, and has fewer teeth
(sixteen as against twenty-six).

Mr. True's P. dallii of the Pacific (where the Common
Porpoise also occurs) is characterised chiefly by its very long
vertebral column, consisting of ninety-eight vertebrae ; there
are only sixty-eight in the other species. The Eastern genus
Neomeris is placed with Phocaena by Dr. Blanford. It practi-
cally only differs by the absence of a dorsal fin. It is only
about 4 feet long, and inhabits the seas of India, Cape of Good
Hope, and Japan. The one species is called N. phocaenoides.

The genus Globicepkalus is to be defined thus : — Teeth

xii GRAMPUS AND KILLER 375

seven to twelve on each side, confined to anterior end of
jaws. Skull raised into a prominence behind the blow-hole;
pterygoids large and in contact. Pectoral fin long and falcate ;
dorsal fin present. No beak. Vertebral formula C 7, D 11, L 11
to 14, Ca 27 to 29. Six pairs of the ribs are two-headed.

The best known species of the genus is the Ca'ing Whale,
G. melas.1 This animal reaches a length of 20 feet, and is thus
one of the largest of the Delphinidae. It is gregarious and was,
even is now, much hunted in the Faeroe Islands. Its sheep-
like habits (embodied in one scientific name deductor} enable
it to be easily driven on shore in herds, which are then har-
pooned. The foetus of this Whale has a few hairs ; the number
of phalanges in the two middle digits is very great, as many as
eleven to fourteen. G. scammoni, G. brachypterus, and G. indicus
are other reputed species of the genus allowed by True and
Blanford.

Grampus is a genus allied to the last. It has no teeth in
the upper jaw, and but three to seven in the lower jaw, near the
symphysis of the mandible. The pterygoids are in contact.
There is no beak, and the pectoral fin is long. There are twelve
pairs of ribs, of which six are two-headed. Apparently there
is but one species, G. griseus, known as "Kisso's Dolphin."
It is a Mediterranean and Atlantic form, and is not common.

The genus Orca has as characters : — Teeth ten to thirteen,
long and strong. Pterygoids not quite meeting. Vertebrae
C 7, D 11 to 12, L 10, Ca 23. The first two or three fused.
The. dorsal fin is long and pointed.

Of this genus there may be more than one species ; but the best
known is the Killer Whale, 0. gladiator (Fig. 180, p. 341), often
spoken of as the " Grampus." 2 It is marked with contrasting bands
of white or yellow upon a black body-colour. The animal grows to
a considerable length, as much as 30 feet. Orca is a powerful and
rapacious Whale; and Eschricht has stated that from the stomach
of one, thirteen Porpoises and fourteen Seals were extracted. They
will also combine to attack larger Whales, and Scammon has
related how he witnessed such an onslaught upon a California!!

1 See an essay on the hunting of this Whale, by S. H. C. Miiller, in Fish and
Fisheries, Edinburgh (Blackwood), 1883.

2 Grampus being a contraction of grand poisson is an obvious name to apply
to any Whale.

376 THE " HORRIBLE SEA-SATYRE " CHAP.

Grey Whale. " Belua truculenta dentibus," observed Olaus Magnus
of this Cetacean. The high dorsal fin has been much exaggerated
in old drawings ; it has been even represented as strong and
sharpened at the end, so as to be capable of ripping open the
belly of a Whale. The fact that it sometimes lies over a little
to one side is responsible for another anecdote : that an example
of this Whale was seen to retire with a. couple of Seals tucked
away under the flippers, another grasped by the dorsal fin, and a
fourth in the mouth ! " When an Orca pursues a whale," wrote
Dr. Frangius, " the latter makes a terrible bellowing like a bull
when bitten by a dog." It is probable, according to F. Cuvier,
that this Whale is the " Aries marinus " of the ancients, certain
bands of white upon the head giving an impression of curved
horns. It may also be the " horrible Sea-satyre " of Edmund
Spencer.

Allied to Orca, but distinguishable from it by some rather
minute peculiarities, is Pseudorca. It may be thus defined : —
Teeth eight to ten, much like those of Orca. Dorsal fin
rather small, falcate. Vertebral formula C 7, D 10, L 9, Ca 24.
Six or all the cervicals united. The curious fact about this
Whale, which embraces only a single species, P. crassidens, is that
it was first known in the fossil condition from remains discovered
in the fens of Lincolnshire. An important day for cetologists
was that on which a whole herd entered the Baltic and furnished
material for a better study of this Whale. It is not, any more
than its near ally Orca, confined to northern seas ; for several
examples, at first relegated to a distinct species (P. meridionalis),
have been obtained from the seas round Tasmania.

Orcella (which has been written Orcaella) has fourteen to
nineteen small sharp teeth in each half of each jaw. The
pterygoids are widely separate. The dorsal fin is small and
falcate. The vertebral formula is C 7, D 14, L 14, Ca 26.
Seven ribs are two-headed, and five of them reach the sternum.

This genus contains but a single species, D. brevirostris, which
is both marine and fresh-water in habit ; it occurs in the Indian
seas, and in the Irrawaddy even as far up as 900 miles from the
sea. Some regard the fresh-water individuals as a distinct form,
0. flumincdis.

Sagmatias is a genus known only from a skull, which is
remarkable for the elevation of the premaxillae into a crest ; the

xii STORIES OF THE DOLPHIN 377

pterygoids are short, and there are thirty-two teeth in each half
of each jaw.

Feresia is known from two skulls which are provided with
ten to twelve teeth in each half of each jaw. It is intermediate
between Globicephalus, Grampus, and Lagenorhynchus, according
to Sir W. Flower.

The genus Delphinus contains the Dolphin, D. delphis.1 The
genus may be characterised as follows: — Teeth small and
numerous, forty-seven to sixty-five. Vertebral formula C 7,
D 14 or 15, L 21 or 22, Ca 30 to 32. The atlas and axis are
fused, the rest free. The palatal border of the maxillae is
deeply grooved. The fins are falcate ; the beak long and distinct.

The Common Dolphin of the Mediterranean shows so many
variations of colour, slight differences in the proportions of the
bones of the skull, and in the number of the teeth, that it has
been divided up into at least seventeen " species." But M.
Fischer, who has studied many of these forms, does not admit
them, and most students of this group of mammals follow him in
the matter. The Dolphin is and has been the most familiar of
Cetaceans ; in consequence it has accumulated much anecdote of
a mythical character. The extreme intelligence and goodwill
towards man assigned to this creature by the ancients are possibly
due to the anomaly of a creature ostensibly a fish showing many
of the characters of higher animals. Its unfishlike intelligence
baffled the early observers, who at once endowed it with especi-
ally advanced attributes. Hence the stories of Arion and others.
The leaping of the Dolphin out of the water- is exemplified in
many Mediterranean coins and coats of arms ; the heraldic
dolphin is represented with an arched back as in leaping. The
Dolphin reaches a length of some 7 feet, and appears to be
world -wide in range. Possibly distinct is D. lonyirostris,
characterised, as the name denotes, by the very long beak ; it
has also more teeth and is a native of Malabar. D. roseiventris
again may be a third species of Delphinus. It comes from
Torres Straits, and has the under parts rosy in colour.

The genus Prodelphinus has, like Delphinus, a distinct beak ;
but it has not the grooved maxillaries. No other character of
importance appears to separate it from Delphinus.

1 See Actes Soc. Linn. Sordeaux, 1881 ; and for another figure, also coloured,
Flower, in Trans. Zool. Soc. xi. 1880, pi. i.

3/8 FRESH- WATER DOLPHINS CHAP.

The genus consists of some eight widely distributed species,
which are none of them large Dolphins.

Lagenorhynchus has the following assemblage of characters : —
Head with short, not very distinct beak. Dorsal and pectoral
fins falcate. Teeth small, twenty-two to forty-five in each half
jaw. Vertebrae ranging in number from se.venty-three to ninety-
two. Pterygoid bones either in contact or separate. There are
fifteen or sixteen pairs of ribs, of which six are two-headed.
Of this genus Mr. True allows eight species, which have been
increased by a ninth since the publication of his " Eevision." l

Two species of Lagenorhynchus are known from our coasts ;
the rest are mainly southern in range. The British species are,
firstly, L. albirostris, a Dolphin of some 9 feet in length. It has
a large number of vertebrae, ninety-two in number. L. albirostris
is a rare species, the first record of its occurrence on these shores
being in 1834. Since that date some eighteen individuals have
been shot or stranded on the shores of the British Isles. The
second British species, L. acutus, differs in colour from the first.
As in the last, the upper parts are black and the under parts
white; but in L. acutus there is also a stripe on the flanks,
brownish in colour. It has fewer vertebrae, not more than
eighty-two.

The next genus of Dolphins, Sotalia, is characterised by —
Teeth tolerably large, twenty-six to thirty-five. The vertebral
formula is C 7, D 11 or 12, L 10 to 14, Ca 22. The pterygoids
are not in contact in the middle line. It has a distinct beak.

Of this genus there are some six species (the exact number, as
in so many other genera, cannot be positively asserted), most of
which are fluviatile or estuarine in habit. They are also on the
whole characterised by their pale, if not actually white, coloration.
S. sinensis of the Amoy is white with pinkish fins. Sotalia
guianensis is American as its name denotes. It is figured by van
Beneden as of a pale brown colour. It is very abundant in the
Bay of Bio de Janeiro, and has the reputation of being a friend of
man like some other Dolphins. The natives hold that it will bring
to shore the bodies of drowned persons. The most singular species
of the genus is that recently described by Professor Kiikenthal
as S. teilszii.2 This animal is purely fresh-water, being found in

1 Bull U.S. Nat. Mus. No. 36, 1889.
- Zool. Jahrb. Syst. Theil, vi. 1892, p. 442.

xii STENO AND TUKSIOPS 3/9

the Oanmroon river, where it is extremely rare. The nostrils
(blow-hole) are prolonged into a snout-like process, a fact which is
of interest in connexion with the assertion that in Balaenoptera
the blow-hole is puffed out during spouting. What is temporary
in the Rorqual appears to be permanent in the Sotalia. More
remarkable still, perhaps, is the assertion that it is a vegetable-
feeding Dolphin. This is not a mere assertion except that it
may not apply universally ; for in the stomach of a specimen
nothing but vegetable debris was found. But in the stomachs of
other Whales (e.g. Rliachianectes) vegetable matter has also been
found, which may perhaps have been taken in accidentally \\h\\
the food.

Steno comes near Sotalia, and Dr. Blanford has transferred
to it (under the one name of Steno perniger) the two species,
Sotalia gadamu and Sotalia lentiginosa. It is, however, to be
distinguished from Sotalia by the following characters: — Teeth
large and few, twenty to twenty-seven on each side of each jaw,
with furrowed surfaces to crowns. Vertebrae C 7, D 12 cr 13,
L 15, Ca 30 to 32. Pterygoids in contact. There are but two
species apparently (not counting Dr. Blanford's).

Tursiops is not a very easily definable genus. These
are its chief features: — Teeth large, twenty -two to twenty-
six in number in each half of each jaw. Vertebral formula
C 7, D 12 or 13, L 16 or 1*7, Ca 27. Pterygoids in contact,
Beak distinct. Some five species are allowed ; but it seems
to be difficult to differentiate the others from Tursiops tursio.
This, the best-known form, is quite or nearly world-wide in
range, and occurs, though not abundantly, on our own coasts.
Mr. True has observed that the eyelids of this Whale, which
is largely hunted on the American coast, are as mobile as
those of a terrestrial mammal. The name " tursio " is derived
from Pliny. Belon would also derive from this word the
French vernacular "rnarsouin." The latter term is sometimes
regarded as a corruption of " Meerschwein," but it would seem
to be more probably derivable from " marinum suem," from the
Latin direct. T. tursio has the back black to lead-colour;
the under parts white. In the reputed species, T. abusalam,
from the Red Sea, the back is a dark sea-green. T. tursio reaches
a length of 1 2 feet, but is more usually smaller.

The o-enus Tursio must be carefully distinguished from

3 So THE SUSU CHAP.

Tursiops. It has no dorsal fin, the teeth are small and numerous
(forty-four), and the pterygoids are separate. There are two
species, T. borealis and T. peronii, the former being northern and
the latter more widely spread.

The genus Cephalorhynchus has for its chief characters the
following: — Teeth twenty-five to thirty-one, small and sharp.
Pterygoids widely separated. Dorsal fin not falcate, but tri-
angular or ovate in form. Beak not well marked off from the
head. The species of this genus are all southern in range ; four
are perhaps to be allowed.

Fam. 3. Platanistidae. — This family of Odontocetes may be
distinguished from the Dolphins by the following assemblage of
structural features: — Cervical vertebrae all free, and each one of
some length (for a Cetacean). Jaws long and narrow, with a
considerable length of symphysis. Teeth very numerous.

This very meagre series of differential characters is largely due
to Pontoporia on the Platanistid side, and to Monodon and
Delphinapterus upon the Delphinid side. Otherwise the family
Platanistidae would be extremely distinct. The two last-named
genera have separate cervical vertebrae, and in the Beluga at any
rate this is expressed externally by a quite distinct neck.
Moreover, as Mr. True has pointed out, the pterygoid bones have
not the involuted cavity below which characterises other
Dolphins ; and they have, what other Dolphins have not, an
articulation outwards with the roofing bones of the skull. Sir
W. Flower described the fact that in Inia (and the same occurs
in Pontoporia} the palatines are separated from each other by the
intervention of the vomer. In this feature they resemble certain
Ziphioids, Berardius, Oulodon ( = Mesoplodon) grayi, and Hypero-
odon. The true Dolphins also appear to show the same inter-
vention of the vomer in a few cases. There is nothing, therefore,
distinctive from the Delphinidae in this feature.

The existence of cartilaginous sternal ribs in Inia and
Platanista shows affinity between these two genera and the
Physeteridae. Pontoporia is Dolphin-like in this particular, as
it is also in the mode of articulation of the ribs with the verte-
bral column. But this last matter has already been dealt with.
The principal reason for placing Pontoporia with the other two
genera is the close resemblance which its skull bears to that of Inia.

The first genus of this family which will be noticed is Platanista.

xii PLATANISTA AND INI A 381

The following are its main characters : — Dorsal fin absent. Eyes
rudimentary. Pectoral fins large and truncated at the extremity.
Teeth, about twenty-nine in each half of each jaw. Scapula with
the acromion coinciding with its anterior edge. Skull with enormous
maxillary crests, and with the palatines entirely concealed by the
pterygoids. The length of the above definition will serve to
indicate how anomalous in many particulars is the structure of
this " Dolphin."

There is apparently but one species, P. gangetica. the " Susu."
The Indian vernacular name is derived from the sound that the
animal makes when spouting. It is an inhabitant of the Ganges
arid the Indus, together with their tributaries, and ascends very
high up its streams. It is also thought to be purely fluviatile
and never to desert the rivers for the sea. Platanista lives
chiefly by grubbing in the mud for prawns and fish. Grains
of rice have also been found in the stomach, but this would seem
to be accidental. The long snout of the Susu has been compared
to the long snout of the Gharial, a native of the same region.
This Whale grows to a length of over 9 feet, but this length is
exceptional. Its anatomy has been elaborately described by Dr.
Anderson.1

The next genus, Inia, is thus to be characterised : — Dorsal fin
rudimentary ; pectorals large and ovate. Teeth, as many as thirty-
two on each side, often with an additional tubercle. Skull without
large maxillary crests ; palatines not hidden by pterygoids, but
divided by vomer. The vertebrae of this genus are few in
number, only forty-one in all, which are thus distributed : C 7,
D 13, L 3, Ca 18. The peculiarities of the vertebral column
are several. In the first place, as has been mentioned in the
definition of the family, all the cervicals are separate and
individually of some length. Secondly, the axis has a better
trace of an odontoid process than in any other Whale except
Platanista, where it is even more obvious. The lumbar region is
remarkable on account of its restriction to three vertebrae. The
sternum, by what we must regard as convergence, is somewhat
like that of the Whalebone whales. It consists of one piece only,
of a roughly-oval form, to which apparently only two pairs of
(cartilaginous) sternal ribs are attached. In the fore-limb the
proportions between the humerus and the radius are more like

1 Anatomical Researches Yunnan Exp. 1878, p. 417.

382 THE AMAZONIAN DOLPHIN CHAP.

those of terrestrial mammals ; i.e. the humerus is distinctly the
longer, the converse usually obtaining among Whales. But
Plata nista again agrees with Inia. The teeth are remarkable
for the fact that the hindermost ones of the series have an
additional lobe ; they are not purely conical as are those of
Whales generally.

There is but one species, Inia yeoffrensis, which inhabits the
Amazons, and grows to a length of 8 feet. Its colour
variations are rather extraordinary, unless they can be set down
to sex, which has been denied. Some individuals are wholly
pink ; others are black above and pink beneath. This Whale is
believed by the Indians to attack a man in the water, and it is
added that the Sotalia of the same streams will defend him from
these attacks ! Naturally, therefore, superstitious reverence
attaches to this Dolphin, which is tiresome to the naturalist who
wants specimens, as Professor Louis Agassiz found.

In the genus Pontoporia l the dorsal fin is well developed
and falcate. The teeth are very numerous, 200 in all. The
ribs articulate as in Dolphins. The skull closely resembles that
of Inia, and the scapula is, as in that genus, " normal."

The proper name for Pontoporia is really Stenodelphis, which
name was first used by Gervais a month or two before Gray, who
separated it from the vague Delpkinus of its original discoverer,
Gervais himself. It has a longer snout than Inia, which, being
bent towards the extremity in a downward direction, curiously
suggests the skull of a Curlew. In details, however, the skull is
exceedingly like that of Inia. It is nearly symmetrical. The
vertebral formula appears to be the following: — C 7, D 10, L 5,
Ca 20 = 42, just one over the number of the vertebrae in
Inia. The sternum is in two pieces. Of the ten pairs of
ribs the first three are double-headed. These and the next have
sternal moieties joining the sternum, of which the first three are
ossified, the last being apparently merely a ligament.

There is a single species of the genus, P. Uainvillii. This
Whale is described by Mr. Lydekker as being of a clear brown
colour, harmonising with the waters of the estuary of the
Amazons and the La Plata which it inhabits. The same colour
characterises Sotalia pallida of those parts of the world, and

1 Flower, Trans. Zool. Soc. vi. 1867, p. 106 ; and Burmeister, Proc. Zool. Soc.
1867, p. 484.

FOSSIL WHALES 383

may be a colour adaptation. But the extant accounts of the
colour of this Dolphin vary- — quite possibly in accordance with
real variations, such as are exhibited by Inia already spoken of.
Pontoporia Uainvillii is a smallish Dolphin some 4 feet in length.

Fossil Odontocetes. — Several of the existing genera of
Dolphins are also known in a fossil condition, as well as
Ziphioid Whales closely related to existing forms. We shall
deal here only with a few genera of fossil Odontocetes which
depart in their structure from existing forms.

The genus Physodon is Miocene, and has been found in
Patagonia. It appears to be most nearly allied to the Physeteridae,
but should probably form a distinct family. Physodon was not so
large as Physeter, the skull measuring only some 10 feet. It
thus comes nearer in point of size to Kogia, and it is interesting
to note that its relatively-shorter snout is also suggestive of the
dwarf Cachalot. The general outline of the skull is, however,
more like that of Physeter, and there is the same deep cavity for
the lodgment of spermaceti. The main feature of interest in
the skull is the presence of teeth in both jaws, and the fact that
two or three are lodged in the premaxillae. This is precisely
what is found in the most ancient Whales, the Zeuglodonts.

Extinct Dolphins, apparently referable to the Platanistidae, are
the most numerous among the earlier forms of Cetaceans, and it
is significant that the earliest known forms of these go back to
the Eocene.

The genus Iniopsis of Mr. Lydekker,1 with one species, /.
caucasica, comes from rocks which seem to be of that age. The
back part of the skull of this animal, the only part of the skull
known, has the same squarish excavation of the maxillaries that
characterises Inia and Pontoporia. Its lower jaw was slender
and possessed numerous teeth.

The long snout and jaws of Platanistids, especially exaggerated
in Pontoporia among living forms, are constantly found in these
Tertiary Platanistids.

E'urhinodetyhis had a beak three and a half times the length
of the cranium, whereas in Pontoporia the proportions are as 2:1.
The teeth too were very numerous.

The genus Argyrocetus, from Patagonian Tertiary strata, was
an animal about as large as the existing Dolphin. It had the

1 Proc. Zool. Soc. 1892, p. 558.

384 ARCHAEOCETI CHAP.

slender rostrum and numerous teeth of the Platanistids and the
squared excavations of the maxillaries. Argyrocetus patagonicus
possessed also archaic characters, suggesting earlier affinities still.
The two condyles of the skull instead of being closely adpressed
to the skull stood out in a way more like that met with in
terrestrial mammals. The nasal bones instead of being abbreviated
rudiments are well developed as in the archaic Zeuglodonts.
The cervical vertebrae of this Whale are all perfectly free from
each other and individually long. The skull is on the whole
bilaterally symmetrical ; this again is a feature more pronounced
among the Platanistidae than among other Odontocetes. Accom-
panying these generalised Cetacean characters are some which
show that the animal was too specialised to be the direct ancestor
of any existing forms. The end of the mandible was upturned
and without teeth, its form being quite unique among Cetacea.
Other allied forms, such as Zarrhachis and Priscodelphinus,
showed the same length of the cervical vertebrae.

A very distinct family of extinct Whales is that of the
Squalodontidae. They to some extent bridge over the gap
between the existing Odontoceti and the Eocene Archaeoceti
(Zeuglodonts).

The skull of these Whales was on the whole Dolphin-like.
But they possessed teeth which were distinctly specialised into
incisors, canines, and molars. The molars have a coarsely-serrated
cutting edge as in the Zeuglodonts, and are also to some extent
two-rooted. But they are more numerous, and so far approximate
to the conditions which characterise the more typical modern
Odontocetes. Squalodon was a long-beaked form, and Prosqualodon
had a skull whose proportions are nearer those of Kogia.

SUB-ORDER 3. AECHAEOCETI.

This division of the Whale tribe embraces but a single family,
Zeuglodontidae, of which but a single genus, Zeuglodon, can with
certainty be discriminated.

Zeuglodon is an Eocene form of large size, with teeth which
are limited in number and disposed in three series as incisors,
canines, and molars. The molars are double-rooted, a fact which
has given to the genus its name. The nasal bones being long

xii AFFINITIES OF ZEUGLODON 385

instead of rudimentary like those of other Whales, the blow-
hole lies more in the middle of the face. The skull, too, is
not Whale -like in a number of other points. Thus the pre-
maxillaries take their fair share in the outline of the upper jaw ;
and, furthermore, bear the incisor teeth. The parietals meet
above in a crest and are not excluded from the roof of the
skull. The vertebrae of the neck are in no way shortened ;
neither are they fused together. The ribs are double-headed, and
the sternum is made up of several pieces. Some naturalists,
particularly Professor D'Arcy Thompson,1 have assigned a relation-
ship to the Seals to these ancient Cetacea ; but others2 have
disputed this view chiefly on the grounds that the characters
which appear to be Seal-like are simply characters which are
generalised and so far at most not Whale -like. Thus the long
neck and the serrated character of the teeth may be accepted as
Seal-like on the one hand ; but on the other, a simple serrated
tooth and a long neck are not by any means 'features of organisa-
tion which we should consider out of the way in an ancient form
of Cetacean which probably preyed upon fish. The humerus of
Zeuglodon, according to Mr. Lydekker, puts out of court any
possible near relationship to the Seals. But the matter under
dispute can be further studied by reference to the three memoirs
quoted below.

1 Thompson, Studies Mus. Dundee, i. 1890 ; and C. It. Congris de Zoologie, 1889,
p. 225.

2 Lydekker, Proc. Zool. Soc. 1892, p. 560.

VOL. X

CHAPTER XIII

CAKNIVOKA l FISSIPEDIA

Order VII. CARNIVORA

THIS order may be thus defined : — Small to large quadrupeds,
terrestrial, arboreal, or aquatic, of usually carnivorous habits.
The teeth have generally sharp and cutting edges, and the canines
are well developed ; the incisors are small, and four to six in
number. The number of toes is never less than four. There
are usually strong and sharp claws. The clavicles are incomplete
or absent. In the hand the scaphoid and lunar bones are always
united. The brain is well developed, and the hemispheres are
well convoluted. The stomach is always simple, while the
caecum, if present, is always small. The members of this
group have a deciduate and zonary placenta.

The fewness of the characters used in the above definition is
chiefly owing to the fact that the Seals and Sea-lions, although
they are referable without a doubt to this order, have undergone
in their metamorphosis into aquatic animals so many changes that
some of the main features in the structure of their terrestrial
relatives have been lost. This group will, however, be again
characterised. We shall deal at present with the land division
of the Carnivora, the CARNIVORA FISSIPEDIA as they are generally
termed. The name is of course given to them to distinguish
them from the corresponding division of the PINNIPEDIA. In
the latter group the feet and hands are modified into " fins " ; in
the other the fingers and toes are cleft, as with terrestrial beasts
generally.

1 For a general account of the osteology, see Flower, Proc. Zool. Soc. 1869, p. 4 ;
and for muscular anatomy, Windle and Parsons, Proc. Zool. Soc. 1897, p. 370, and
1898, p. 152.

CHAP. XIII

DOG'S SKULL

387

SUB-ORDER 1. FISSIPEDIA.

A very marked feature of the terrestrial Carnivora is to be
found in the structure of the teeth. The incisors are nearly
always six, and are somewhat feebly developed in many cases. The

FIG. 190.— Under surface of the
cranium of a Dog. x £. upf,
Anterior palatine foramen ; AS,
alisphenoid ; as, posterior open-
ing of alisphenoid canal ; BO,
basioccipital ; BS, basisphenoid ;
cf, condylar foramen ; earn, ex-
ternal auditory meatus ; ExO,
exoccipital ; Jim, Jtp, foramen
lacerum medium and posterius ;
./>«, foramen magnum ; fo, fora-
men ovale ; Frt frontal ; fr,
foramen rotundum ; gf, glenoid
fossa ; gp, post-glenoid process ;
Ma, malar ; MX, maxilla ; oc,
occipital condyle ; op, optic fora-
men ; Per, mnstoid portion of
periotic ; pgf, post-glenoid fora-
men ; PI, palatine ; PMx, pre-
niaxilla ; pp, paroccipital pro-
cess ; ppf, posterior palatine
foramen ; PS, presphenoid ; Pt,
pterygoid ; sf, sphenoidal fissure
or foramen lacerum auterius ;
sm, stylomastoid foramen ; SO,
supraoccipital ; Sq, zygomatic
process of squamosal ; Ty, tym-
panic bulla ; Vo, vomer. (From
Flower's Osteology.)

apf

canines are almost invariably very large strong teeth, and are
always present. In some of the extinct Cats they reached
enormous dimensions. The number of cheek teeth is not always
identical ; but the last premolar in the upper jaw and the first true
molar in the lower jaw, known as the " carnassial " or " sectorial "
teeth, mark a difference in structure between the anterior and
the posterior crushing teeth ; those in front of the carnassial
tooth have cutting edges, and are often merely small, conical
teeth ; those behind have broader crowns and are tuberculate ;
those of simpler forms often trituberculate ; those of others

388

OSTEOLOGY AND

with numerous tubercles. The carnassial tooth is often, but
by no means always, very much larger and especially longer
than the rest of the molar and premolar series. It is less
pronounced in some of the omnivorous Arctoidea. The skull of
the Carnivora is longer in the more primitive types, such as the
Canidae, and shorter in the more specialised Felidae. The orbit
is hardly ever completely shut off by bone, though the postorbital
process of the frontal sometimes approaches the corresponding
upward process of the zygomatic arch. The palate, which is
completely ossified, sometimes reaches back for some distance
behind the teeth ; it always extends as far as the last molar.
The tympanic bulla is often very inflated, and if flatter, as in the
Bears, is at any rate large and conspicuous. The lower jaw has

FIG. 191. — A, Atlas of Dog. Ventral view, x ^. B, Axis of Dog. Side view, x 4f.
o, Odontoid process ; pz, posterior zygapophysis ; s, spinous process ; sn, foramen
for first spinal nerve ; t, transverse process ; v, vertebrarterial canal. (From
Flower's Osteology.)

a high coronoid process, and the condyle is transversely elongated,
this part of the bone being rolled into an almost cylindrical
form ; it fits very closely into the glenoid cavity, and the
articulation is thereby very strict — an obvious advantage in a
creature with so great a need for power of jaw.

In the vertebral column the atlas always has large wing-like
processes ; the spine of the axis vertebra has a long antero-
posteriorly elongated form. The transverse processes of the
fourth to the sixth cervicals are, as a rule, double. These
features, however, though characteristic of the Carnivora are not
by any means distinctive. The true sacrum consists of but a
single vertebra to which the ilia are attached ; but at most two
other vertebrae are fused with this. The clavicle is always small
and sometimes quite rudimentary, or even absent. The spine of
the scapula is well developed, and almost equally divides the

BRAIN OF CARNIVORA

389

surface of that bone. The digits of the Carnivora are mostly
five, and are never less than four. The mode of progression may
be digitigrade or plantigrade, and the intermediate semidigitigrade

-S.ol

Rol
i

FIG 192.— Brain of Dog. A, Ventral : B, dorsal ; C, lateral aspe
lobe ; Cr.ce, crura cerebri ; Fi.p, great longitudinal fissure r;
lobes of cerebellum ; Hyp, hypophysis ; Med, spinal cord ; <™.
Po wms Varolii • Ftf, cerebral hemispheres ; Wu, middle lobe (vermis]
• I-%II cerebral nerves. (From Wiedersheim's Comparative Am

mode of walking also occurs. The brain in all Carnivora is large
and well convoluted. The arrangement of the convolutions
characteristic. There are three or four gyri disposed round each
other of which the lowest surrounds the Sylvian fissure,
stomach in these creatures is always simple in form., without

390 CLASSIFICATION OF CARNIVORA CHAP.

subdivisions. The caecum is never large, and may be, as in the
Bear tribe, completely absent.

The distribution of the Carnivora is world-wide, excluding only
the Australian region, if, as seems probable, the Dingo of that
region is an introduced species. The most striking features in
their distribution are perhaps the following : — There are no Bears
in the Ethiopian region or in Madagascar, and but a single species
in the Neotropical. The only Carnivora in Madagascar are the
Viverridae, and of the seven genera there found six are peculiar.
The Procyonidae are nearly entirely New World in range ; out of
sixteen genera of Mustelidae only five are New World, and only
two of those are peculiar to the American continent. The
Hyaenidae are limited to the Old World.

The classification of the Carnivora is a matter which is
difficult, and which has therefore been very variously effected.
It is unfortunate that the classification of Flower (based upon
the researches of H. N. Turner as well as his own, and accepted
by Mivart) should fail when applied to fossil forms. For it
separates with great clearness the existing genera into three
great divisions, the Cynoidea, Aeluroidea, and Arctoidea, definable
by visceral as well as by osteological characters. The apparent
anomaly, too, of a single supposed Viv.errine genus, to wit
Bassariscus, existing in America, while all the rest of its kin
are Old -World forms, was shown by his characters to be
neither an anomaly nor a fact. It will be better, therefore,
to divide the Carnivora into the families, Felidae, Machae-
rodontidae, Viverridae, Hyaenidae, Canidae, Ursidae, Procyonidae,
and Mustelidae,- indicating at the same time the reasons
for and against retaining the three divisions of Sir W.
Flower.

Fam. 1. Felidae.1 — This family includes only the Cats (i.e. Lions,
Tigers, " Cats," Hunting Leopard, etc.), and is to be distinguished
by the following characters : — In the skull the auditory bulla is
much inflated, and there is an internal septum ; the paroccipital
processes are flattened against the bullae. There is no ali-
sphenoidal canal. The dental formula is I 3, C 1, Pm 3 to 2,
M 1. The carnassial tooth of the upper jaw has three lobes to
the blade ; that of the lower jaw is without an inner cusp.

1 See St. G. Mivart "On the Aeluroidea," Proc. Zool. Soc. 1882, p. 135 : and
The Cat, London, J. Murray, 1881.

CLAWS OF CATS

391

The digits are five on the fore-feet, four on the hind. The
caecum is present and small. This family contains but two
genera, Felis and
Cynaeluriis.

The genus Felis is
very wide in its dis-
tribution, being com-
mon to both the Old
and the New Worlds.
Its distinctive charac-
ters, as opposed to
Cynaelurus, are mainly
the following : — The
claws are retractile,
and the retractility is
more markedly de-
veloped than in the

Cheetah.- The molar
is not so nearly in
a line with the other
over, has an inner tubercle.

FIG. 193. — Section of auditory bulla of Tiger, am, Audi-
tory meatus ; BO, basioccipital ; e, Eustachian canal ;
ic, oc, two chambers of bulla divided by s, septum ;
*, their aperture of communication ; Pt, periotic ;
Sq, squamosal ; t, tympanic ring. (From Flower's
Osteology.)

teeth ; the upper carnassial, more-
The legs are relatively shorter.

The complete retractility of
the claws is a very distinctive
feature of the true Cats. It is
brought about in this way : the
terminal joint of the toe, which
is clad with the claw, folds back
FIG. 194.— The phalanges of the middle into a sheath by the outer side

digit of the manus of the Lion (Felis Qf Qr above the middle phalanx.

leo). x \. a, The central portion _..,,. , . f, • •,

forming the internal support to the It IS held in this position by

horny claw ; b, the bony lamina re- ft sfcron(T ligament. The flexor

fleeted around the base of the claw ; °

ph\ proximal phalanx ; ph*, middle muscles straighten the phalanx

phalanx ; ph3, ungual phalanx. (From Wj1jcj1 Dears tne claw so that
Flower's Osteology.)

the natural position for the

animal is to be in a state of retracted claws, which of course
preserves them from friction ; when wanted for aggressive pur-
poses, they are pulled into sight by the action of the muscles
already mentioned.

Much has been written as to the shape of the pupil of the
Cat's eye. Some careful observations upon the matter have been

392 CATS' EYES CHAP.

made by Dr. Lindsay Johnson,1 who found that out of 180
Domestic Cats 111 had round pupils, and that in 19 the shape
was a pointed oval, intermediate conditions being offered by the
rest. These 180 comprised males and females of many varieties.
When the pupil of the Cat's eye contracts, it forms a vertical slit
with two pin holes, one at each end, through which alone light
appears to enter. In the Genet and the Civet the contraction
of the pupil is as in the Cat. In the Lion, Tiger — in fact
apparently in all the large Cats — the pupil retains its circular
shape even when contraction is fully effected. Dr. Johnson has,
furthermore,2 made some interesting experiments upon the Seal's
eye — a creature which has, of course, to exert its powers of
vision in two media, and from one to the other. This is effected
by dilatation of the pupil when in the water, and its contraction
to a vertical slit with parallel margins and rounded ends when
in the air, the contraction being to some extent at least under
the influence of the animal's will.

The coloration of these creatures is very varied : spots of
black, or bordered with black upon a more or less tawny ground-
colour, is the prevailing pattern. Stripes are also met with, as
in the Tiger, but these are usually cross stripes,3 while in the
related Viverridae there are many examples of longitudinal
stripes. Finally, many Cats, as for instance the Puma and the
Eyra, are " self-coloured " — have, that is to say, a uniform tint.
Just as the unstriped Horse sometimes shows traces of the
former existence of stripes, so the self-coloured Cats are occasion-
ally spotted when young ; this is markedly so in the case of the
Puma ; while the Lion is spotted as a cub, and in the adult —
particularly in the lioness — there are distinct indications of these
spots. It is evident, therefore, that there are grounds for re-
garding a spotted condition to be antecedent, at least in some
cases, to a uniform colour. There are divers explanations of
these hues and of these changes. It is held by many that the
coloration has a relation to the habits of the creature: the
spotted Cats, it is pointed out, are largely arboreal ; this is
eminently so with the Jaguar at any rate ; and in an arboreal

1 "On the Pupils of the Felidae," Proc. Zool. Soc. 1894, p. 481.

2 "Observations ... on the Seal's Eye," Proc. Zool. Soc. 1893, p. 719.

3 It is noteworthy that in the Tiger some of the stripes have pale centres and
are thus like spots pulled out, while there are also small black spots. .

COLOURS OF CATS 393

creature the spots, it is said, give the impression of flecks of sun-
light broken up by foliage. On the other hand, the self-coloured
Oats of a sandy to earthen hue assimilate in tint with a sandy or
stony soil. The stripes of the Tiger, it is thought, approximate
to the tall parallel stems of grasses and other plants in the dense
cover in which it lives. In favour of these views is undoubtedly
the fact that in other mammals and other animals belonging to
quite different groups the same four plans of coloration are met
with. Spots and cross stripes are found in the Marsupials ; the
young Tapir is spotted while the adult is self-coloured, and so
forth. This last fact, however, serves to illustrate another view
which has been put forward in explanation of these characteristic
markings of the Felidae. Eimer has come to the conclusion
that there is and has been a regular series of steps in the
evolution of these markings. The primitive condition was, he
thinks, a longitudinally striped one ; the stripes then broke up
into spots, and the spots rearranged themselves as transverse
stripes ; the self-coloured Puma and Lion are a final stage in this
gradual evolution. In support of this is the fact that spots
precede self-coloration in the individual growth of these animals.
The exact sequence of these markings is, however, contradicted
by Dr. Haacke's observations upon a certain Australian fish
which is cross striped when young and longitudinally striped
when adult, a precise reversal of what ought to occur on Eimer's
view.

The Felidae are almost universally distributed with the ex-
ception, of course, of Australia and a good deal of the Australian
region; the headquarters of the group are undoubtedly in the
tropics of the Old World.

The characteristics of a few species of the Cat tribe will now
be given. As there are at any rate forty-five species, this survey
will have to be somewhat incomplete.

The Lion, F. leo, differs from all other species by the mane of
the male. It is an inhabitant of Africa, India, and certain parts
of Western Asia. Within the historic period it ranged into
Europe. According to Sir Samuel Baker those of us who have
not seen the Lion in his native haunts have never seen a really
magnificent specimen of the brute ; but other travellers disagree,
and state that a captive Lion is often a finer animal — by reason,
of course, of good feeding. Unlike the majority of Cats, the Lion

LU LIBRA

394 THE LION AND THE TIGER CHAP.

cannot climb. His roar (which is so suggestive, towards its end,
of that animal who once dressed himself up in his skin) is
literally after his prey. The Lion, it is stated, does not roar
except upon a full stomach. The Lion is mainly nocturnal in its
habits, and is said to be not in the least dangerous if unprovoked
in the daytime ; but here again opinions differ. The tail of the
animal is provided at the extremity with a slight claw, but it
can hardly be sufficient for the animal to lash itself into a fury
with it. A Lion will live for thirty or forty years, and will breed
freely in captivity. The Gardens of the Zoological Society of
Dublin have been famed for their success in breeding Lions ; but
more surprising still, this has been successfully accomplished in
travelling menageries. The " desert " colour of the Lion is familiar
to all. It is stated that the likeness to the parched soil of
certain parts of Africa is greatly heightened by black patches in
the marie, for in certain regions of that continent the arid yellow
of the general environment is diversified by pieces of black lava.
It is apparently a popular delusion to speak of the Maneless Lion
of Guzerat. No doubt maneless Lions do come from there, but so
do young and maneless Lions from other places ; in short, it is
simply a question of age, and old Lions from the Asiatic continent
are as fully maned as those from Africa.

The Tiger, F. tigris, is an animal of about the same size as
the Lion, distinguished, of course, by the stripes. The skeletons
are much like those of other Cats ; but the skull of the
Tiger may be distinguished from that of the Lion by the fact
that the nasal bones reach back beyond the frontal processes of
the maxillae. The Tiger is an exclusively Asiatic beast, ranging
northward into icy Siberia. The northern individuals have a
closer fur, and have been quite unnecessarily separated as a
distinct variety. Nine feet six inches is the size of the average
full-grown Tiger ; but the skins will stretch, a fact of which the
sportsman will sometimes take advantage. A " man-eater " is a
Tiger which has discovered " that it is far easier to kill a native
than to hunt for the scarce jungle game." As with the Lion, the
accounts of travellers differ enormously, particularly with regard
to the strength of the creature. Some have said that a Tiger can
easily lift a full-grown bullock and leap with it in the mouth
over a considerable obstacle, a statement which is ridiculed by
Sir Samuel Baker. Unlike the Lion, the Tiger can climb trees ;

xm THE TIGER AND THE LEOPARD 395

it will also voluntarily enter the water, and can swim consider-
able rivers.

Mr. H. X. Kidley 1 observes that Tigers " habitually swim
over to Singapore across the Johore Strait, usually by way of the
intermediate islands of Pulau Ubin and Pulau Tekong. They
make the passage at night, landing in the early morning. As so
much of the coast is mangrove swamp, and the animals do not
risk going through the mud, they are only able to qross where
the shores are sandy, and thus they have regular starting- and
landing-places."

The Tiger is mainly nocturnal, but begins its depredations
towards five o'clock in the afternoon, before which it remains
sleeping in shady thickets. If the weather is rainy and windy
it becomes restless and wanders about earlier. Under the provo-
cation of extreme hunger it will hunt during the daytime.
Hunger, too, naturally produces extreme boldness. Mr. Kidley
relates a story of four Tigers who walked up the steps of a house
in search of the master of the house or his dog, and broke into
it, the inhabitants retiring in their favour. The Malays have
superstitions about Tigers, which are precisely paralleled by the
man-and-wolf stories of Europe. " Certain people are supposed
to have the power of turning into tigers for a short time, and
resuming their human form at pleasure. The transformation
commences tail first, and the human tiger is so completely
changed that not only has it all the actions and appearance of
the tiger, but on resuming its human form it is quite unconscious
of what it has been doing in the tiger state.", Mr. Kidley dis-
putes the common stories as to man-eaters. If a Tiger has once
tasted human flesh it does not always confine itself afterwards to
that article of diet, nor is it only aged and comparatively tooth-
less animals which hunt man. That they do take a large toll of
coolies is an undoubted fact, and many are the artifices to prevent
the rest from knowing the fate of one of their fellow-workmen, or
of becoming acquainted with the presence in the neighbourhood
of one of the dreaded beasts.

The Leopard or Panther, F. pardvs, is, like the Lion, African
and Asiatic in range. The animal is spotted with rosettes of
black spots surrounding a central field of the tawny colour of the
body generally. Some of the spots are solid and black. " The

1 Natural Science., vi. 1895, p. 89.

396 THE SNOW LEOPARD CHAP.

pantere like unto the smaragdyne " seems to be an inapt descrip-
tion of this Cat, unless indeed the eyes be referred to. The
ancients ascribed to it a most fragrant odour. As with the Tiger,
a northern variety of this Carnivore has a closer and longer fur.
There is a tendency towards melanism in this animal, the black
Leopard being comparatively common, particularly, it appears, in
high lands. Several other variations in colour are known. These
have receiyed different specific names ; but it seems that there is
in reality but one species of Leopard. The Leopard can climb
with the agility of any Cat. Sir S. Baker reserves the name
Panther for large Leopards, which reach a length of 7 feet
6 inches. But there is no valid distinction between any two
such varieties. The Leopard is as ferocious as the Tiger ; and
Sir Samuel Baker advises that the power of the human eye
be not experimented with when meeting unarmed one of these
brutes.

The Snow Leopard or Ounce, F. uncia, is a beautiful creature,

FIG. 195. — Snow Leopard. Fells uncia. XTJ^.

confined to the highlands of Central Asia. The ground-colour is
white, and the spots are larger than those of the ordinary Leopard.
Two examples of this rather rare Carnivore have been recently
on view in the Zoological Society's Gardens, London. The
Clouded Leopard, F. nebr-osa, is an animal of considerable size
(6 feet total length).

xin LYNXES 397

The Fishing Cat, F. viverrina, of India and China, is about
3 feet 6 inches including the tail. Its black spots upon a grey-
brown ground have a tendency to form longitudinal lines. It is
in fact, on Eimer's theory, a case of longitudinal stripes breaking
up into spots. It differs from the bulk of Cats by preying upon
fish, though it is not known how it catches them. It also feeds
upon the large snail Ampullaria. In addition to these there are
twenty-four species of Cats found in the Old World, mainly in the
Oriental region, of small to moderate size.

The European Lynx, F. lynx, has rather long legs, a short tail,

-rr — »---.>•••-

FIG. 196. — European Lynx. Fdis lynx, x

and tufted, pointed ears. It has only two premolars in the upper
iaw instead of the usual three. It seems to be doubtful whether
the Asiatic Lynx can be distinguished from the European, but
the Spanish form, F. pardina, does appear to be distinct.
Common Lynx, sometimes called F. canadensis, also ranges
America, where some other forms exist, known by the spec
names of F. nifa and F. bailey i.

In America there are altogether sixteen species of Cats i
we allow three species of Lynx, none of which, however, does
Dr. Mivart allow to be different from the European a
Lynx (F. lynx}.

398

HABITS OF THE JAGUAR

CHAP.

The largest of American Cats is the Jaguar, F. onca. This is
an arboreal creature with a long, heavy body and short limbs.
Its pelage is much like that of the Leopard, but the spots are
larger and more definitely arranged in groups. There are a
number of distinct rows of spots. The length of the body alone
is not greater than 4 feet. They prey very largely on the

FIG. 197. — Jaguar. Fdis onca. x -fa.

Capybara, and upon turtles, which they surprise upon the sand
when about to lay their eggs ; the reptiles are turned upon their
backs, so as to be incapable of escaping, and the Jaguar then
easily devours them. The Jaguar will even pursue the turtle
into the water, and will devour its eggs and the newly-hatched
young.

The Ocelot is another spotted American Cat. F. pardalis ]
ranges from Arkansas in North America southwards, its range
corresponding with that of the Jaguar. Although small for
one of the " larger cats," the Ocelot inspired with considerable
respect Captain Dampier, who remarked of it : " The Tigre-cat
is about the bigness of a bull-dog, with short truss, body shaped
much like a mastiff, but in all things else, its head, the colour
of its hair, the manner of its preying, much resembling the

1 Tor an account of this and of other mammals which occur in Central America,
see Alston in Messrs. Godman and Salvin's Biologia Centra li- Americana, 1879-
1882.

THE PUMA AND ITS YOUNG

399

tigre, only somewhat less. ... But I have wisht them farther
off when I have met them in the woods ; because their aspect
appears so very stately and fierce."

FIG. 198.— Ocelot. Felis pardalis. x^.

The Puma, F. con-color, the American Lion as it is called in
the north, is a rather smaller animal than the last, and of a
uniform tawny colour, tending to white on the abdomen and to
a dark stripe along the back. The young, as already mentioned,
are very distinctly spotted. Like the Tiger, the Puma can
endure extremes of heat and cold ; it is equally at home in
the snow of North America and among the tropical forests
and swamps of the south. It is a ferocious creature so far as
concerns Deer, Lamas, Eaccoons, even Skunks and Rheas, but,
according to Mr. W. H. Hudson, will not attack man, and will
even defend him against the Jaguar.1 In captivity the Puma
will purr like a Cat.

The Eyra, F. eyra, is another self-coloured American cat,
which has a curious likeness to the totally distinct Cryptoprocta
of Madagascar.

The Wild Cat of Europe, F. catus, is found over the greater
part of Europe, and also in Northern Asia. It was undoubtedly
common at one time in this country, though it appears never to
have extended its range into Ireland. But the real Wild Cat is
now rare in this island, and is confined to certain districts in
1 But Mr. Belt says that the "Tigre" never attacks man unless it be
provoked

400 THE WILD CAT

Scotland. Plenty of alleged wild Cats have been seen and even
shot ; but these are too frequently merely feral Cats, i.e. domestic
tabbies which have reverted to a hunting life. The real Wild
Cat differs from the domestic races by the proportionately longer
body and limbs, the shorter and thicker tail ; the pads of the
toes are not quite black. The period of the gestation of the
Wild Cat, according to Mr. Cocks, is a week or so longer than
that of any domestic Cat.

The Domestic Cat is in fact regarded as the descendant of the
Eastern F. caffra, or (perhaps and) the closely-allied F. maniculata.
It is highly probable, however, that after introduction into this
country as a domestic animal it has interbred with the Wild Cat.
Many allied species of Cats will interbreed, even two so far
apart as the Lion and the Tiger. There are interesting archaeo-
logical and linguistic reasons for regarding the Domestic Cat as
an importation. The legend of Dick Whittington's Cat points
to it being a rare and valuable animal, which a tamed F. catus
would not at that time have been. There was an enactment in
Wales of a penalty against him who should kill the king's Cat,
again suggestive of its rarity and consequent value. The very
name " Puss " is a hint of a foreign origin. Some would derive
it from Perse, and upon this is based the notion that the Cat is
from Persia. But it seems that Puss is the same as Pasht and
Bubastis, showing so far an Egyptian origin for the animal. The
ancestral Cats mentioned above are natives of Egypt.1

The genus Cynaelurus, which includes but a single species,
C. jubatus, the Cheetah or Hunting Leopard, is separated from
Felis by a number of characters. In the first place the claws
are non-retractile, or at least less retractile than those of the
true Cats. It is, moreover, longer legged. The molar is more in
a line with the other teeth of the jaw, and the upper carnassial
tooth has no inner tubercle. Messrs. Windle and Parsons have
lately pointed out many Dog-like features in the muscles. This
animal is about as large as a Leopard, but has plain black spots.
As its vernacular name implies it is used for sport, and is quite
easily tameable. It will purr like the Puma. The Cheetah
occurs in India, Persia, Turkestan, and also in Africa ; the latter

1 See E. Hamilton, The Wild Cat of Europe, London, Porter, 1896 ; and M. G.
Watkins, Gleanings from the Natural History of the Ancients, London, Elliot Stock,
1896.

xin SABRE-TOOTHED TIGERS 40 1

form is sometimes, though quite unnecessarily, separated as C.
lanigera. The genus occurs fossil in the Siwalik deposits of
India, the species being known as C. brachygnatha.

Fam. 2. Machaerodontidae. — This is a family of totally-
extinct Cats which range from the Eocene down to the Pleistocene.
Their general structure is like that of the Felidae ; but they
differ in a number of skeletal features. Thus there is an
alisphenoid canal, and, as in Bears, there is a postglenoid foramen.
There is also a distinct carotid foramen, which does not occur in
the true Cats. The teeth are often distinguished by the huge
size of the superior canines, which are " weapons for penetrating
wounds, without rival among carnivorous animals." These must
have been displayed at the sides of the chin when the mouth
was closed, and it has even been suggested that the animal
possessing these exaggerated canines could hardly have properly
closed its mouth. The lower canines were often on the contrary
much reduced, and in fact incisor-like. In tracing the series of
these Cats we find a gradual reduction of the teeth from a more
nearly complete number down to the specialised dentition of the
existing Cats. The genus Proaelurus, Miocene in range, had
four premolars in each jaw, and two molars in the lower and one
in the upper. This is the greatest number of teeth found in any
member of the group.

The resemblance of this genus to Cryptoprocta has been
insisted upon. Archaelurus has suffered a reduction, since one
premolar in the lower jaw has disappeared, its formula being thus
I | C \ Pm | M ^. The next stage is shown by Dinictis with
three premolars in both jaws. There are a good many species of
this genus which are all American and Miocene. This genus has
five toes upon the hind-feet, and was probably plantigrade. It
had retractile claws.

In the genus Nimravus the dental formula is still further
reduced. Another premolar of the lower jaw has gone, the
formula being thus I f C \ Pm f M £. Nimravus gpmphodus
was a Carnivore about the size of a Panther. It has no third
trochanter upon the femur, which process is present in the
corresponding bone of Dinictis. Pogonodon was an equally
large animal in which the premolars were three in each
jaw, but the molars have become reduced to one in the
lower, as they have in this and other genera in the upper

VOL. x 2 D

402 EOCENE CATS

jaw. Finally, Hoploplioneus has acquired the dentition of exist-
ing Cats.

The Machaerodons, however, show examples with a yet more
reduced dentition than that of the most reduced existing Cat, viz.
the Lynx, which has only two premolars in each jaw and one molar.
In Eusmilus the molar in both jaws is single, and there is but
one premolar in the lower jaw.

The genus Machaerodus itself, which appears to include
Smilodon, is referred by Cope to the true Cats, and not to the
Nimravidae, as he terms the family which we have called here
the Machaerodontidae. These creatures are known as "Sabre-
toothed Tigers," and were of very wide distribution, occurring in
South America as well as in Europe and North America. " As
nothing," remarks Professor Cope, " but the characters of the canine
teeth distinguished these from typical felines, it is to these that
we must look for the cause of their failure to continue. Professor
Flower's suggestion appears to be a good one, viz. that the
length of these teeth became an inconvenience and a hindrance
to their possessors. I think there can be no doubt that the
huge canines in the Srnilodons must have prevented the biting
off of flesh from large pieces, so as to greatly interfere with
feeding, and to keep the animals in poor condition. The size of
the canines is such as to prevent their use as cutting instruments
excepting with the mouth closed ; for the latter could not have
been opened sufficiently to allow any object to enter it from the
front. Even when it opens so far as to allow the mandible to
pass behind the apices of the canines, there would appear to be
some risk of the latter being caught on the point of one or the
other canine, and forced to remain open, causing early starvation.
Such may have been the fate of the fine individual of the
S. neogaeus, Lund, whose skull was found in Brazil by Lund, and
which is familiar to us through the figures of de Blainville."

Machaerodus is placed among the Felidae on account of the
fact that the condyloid and carotid foramina unite with the
foramen lacerum posterius. But as in at least one species,
M. palmidens, there is an alisphenoid canal, which, however, has
disappeared in the more recent American forms, it seems per-
missible to retain the genus in the family Machaerodontidae
though its existence reduces the differential character of that
family to a minimum. The genus goes back to the Eocene.

CIVETS AND GENETS

403

Fam. 3. Viverridae. — The Civets, Genets, and their kind
differ from the Cats in a number of points. They form, however,
by no means so uniform an assemblage as do the Cats ; so that the
difficulty is, as Dr. Mivart has remarked, not to divide them into
sub-families, but to avoid making too many. But before pro-
ceeding to subdivide the family we shall describe the characters
of the family and contrast them with those of the Felidae.

All the Yiverridae are comparatively small creatures. The
head and body are more elongate than in the Cats. The fingers and
toes are generally five ; but there are some (e.g. Cynictis) where
the formula of the toes is as in the Cats, i.e. four on the hind-foot.
In the Suricate the fingers are also reduced to four. The claws are
perhaps never completely retractile,1 and often are not at all so.
The dental formulae of the genera differ considerably ; but in the
majority there are more teeth than in the Felidae. The well-
known sharp-pointed, conical papillae of the Cat's tongue are not
present. The majority have a scent gland beneath the tail, from
which the perfume civet is derived. There are a number of
osteological characters which differentiate the two families ; thus
the alisphenoid canal is sometimes present. The bulla is divided,
as in the Cats, but is externally constricted.

It seems clear from some at any rate of the characters, i.e. the
more complete dentition, the five-fingered hands and feet, the
non-retractile claws, etc., that the Civets are on a lower level of
specialisation than are the Cats.

Sub-Fam. 1. Euplerinae. — The genus Eupleres is in many
ways the most aberrant type of Viverrid, and is placed in a
sub-family, Euplerinae. Its salient feature is the very peculiar
dentition : peculiar in the small size of the canines, the canine-like
character of the anterior premolars, and the resemblance of the pre-
molars to molars. In some of the characters of the teeth, Eupleres
is Insectivore-like, and was formerly grouped with that family.
There are four premolars and two molars in each jaw on each
side. It has five toes upon both fore- and hind-limbs; the skull
is very slender. It has no alisphenoid canal. The only species,
E. goudotii, is of an olive-grey colour, with dark bands across the
shoulders in the young. The nose and upper lip are grooved.
There are no scent glands. It appears to burrow in the ground,
and possibly contents itself with a diet of worms. Eupleres is a

1 The retractility is most marked in the Linsangs.

404 CRYPTOPROCTA

native of Madagascar, where all the most peculiar Viverridae
live.

Sub-Fam. 2. Galidictiinae. — Mivart has placed in this sub-
family the three Mascarene genera, Galidia, Hemigalidia, and
Galidictis. In them the orbit is not enclosed by bone ; there is
no alisphenoid canal, and there are five toes and fingers.

Galidia consists of but one species, G. elegans, of a chestnut
brown colour, with a tail ringed with black. The claws are not
retractile. The scent gland is absent. There are five digits
upon both hand and foot. There are three premolars and two
molars on each side of each jaw. The caecum is (for an Aeluroid)
long, and pointed at the apex ; it is quite twice the length of that
of Genetta.

Closely allied to Galidia is the genus Hemigalidia, of which
there are two species. It is distinguished from the last genus by
the non-annulated tail. It also differs in the dental formula,
which is for the molars Pm ^ M ^. This animal is termed by
Buffon the Vansire. He correctly enumerates its grinders, and
distinguishes it from the Ferret !

Galidictis is a third genus from Madagascar containing
two species, one of which has been unfortunately named G.
vittata, leading perhaps to some confusion with the totally
distinct Galictis vittata. As in the last two genera the digits
are five. The dental formula is that of Galidia. It is dis-
tinguished from the other two genera of its sub-family by the
longitudinal brown striping of the upper part of the greyish
body.

Sub-Fam. 3. Cryptoproctinae. — Cryptoprocta 1 represents a
special sub-family, Cryptoproctinae, and includes only a single
species, the Fossa (C. ferox) of Madagascar. It is the largest
Carnivore of Madagascar, being about twice the size of a Cat,
but with an elongated body; the colour is a tawny brown
with no striping. The animal is active and lithe in its
motions, and is said to be of almost unexampled ferocity in
disposition. Its exact systematic position has been much dis-
cussed. By Zittel it is placed in a sub-family (including the
extinct Proaelurus and Pseudadurus) of the Felidae. Mivart
and Lydekker, on the other hand, regard it as a genus of the
Viverridae. The dental formula of the molars, Pm£ M-k is

o J

1 Beddard in Proc. Zool. Soc. 1895, p. 430.

xin VIVERRA 405

more like that of the Felidae than of the Viverridae, and the
teeth are more Feline in structure. The claws of the feet are
retractile. As to internal structure the Fossa agrees largely
with the Viverridae, but then this family has no very marked
points of difference from the Felidae ; but where the anatomy
does diverge from that of the Felidae it approaches the Viverridae,
especially in the muscular system.

FIG. 199. —Fossa. Cryptoprocta ferox. xj.

The remaining and by far the larger number of genera of
Civets are grouped by Professor Mivart in two sub -families:
the VIVERRINAE, including the genera Viverra, Viverricula,
Fossa, Genetta, Prionodon, Poiana, Paradoxurus, Arctogale,
Hemigale, Arctictis, Nandinia, and Cynogcde ; and the HERPES-
TINAE, including the genera Herpestes, Helogale, Cynictis, and
probably Bdeogale and Rhynckogale. In the Viverrinae the
digits are always five, the claws are more or less retractile, the
prescrotal scent glands are usually present, and the anus does
not open into a sac. On the other hand, the Herpestinae are
characterised by the non-retractility of the claws, the absence of
the glands in question, and the fact that the anus does open into
a terminal sac.

Sub-Fam. 4. Viverrinae. — Viverra includes the true Civets.
The genus, save for one African species, is Oriental in range.
The molar formula is the complete one for the Viverridae, viz.

406 CIVET CAT CHAP.

Pm A M §. The secretion of the prescrotal gland of V. civctta
yields the civet of commerce.

The " Easse," genus Viverricula, has been separated generically
from the true Civets. It is, remarkably enough, common to both
Madagascar 1 and many parts of the Oriental region. It is,
moreover, capable of climbing trees, which its relatives are not.
It has no mane like Viverra and is of slighter build.

FIG. 200. — Civet Cat. Viverra civetta. x J.

Prionodon or Linsang differs from the last two genera in the
loss of an upper molar. It thus approaches the Cats, with which
it also agrees in the furry feet. It is a purely Oriental genus.
It also resembles the Cats in that the claws are apparently quite
retractile, a feature not common among the group. There are
three species of the genus. P. pardicolor has large black spots
and a ringed tail. Its body is some 15 inches in length.
Dr. Mivart has commented upon the particularly small caecum,
which, like that of Arctictis, seems to be on the verge of
disappearance.

Genetta, including the Genets, is almost purely African. It
has the full tooth formula of Viverra ; but is to be distinguished
by the absence of a scent pouch, and by a naked strip of skin
running up the metatarsus. These animals are all brownish
yellowish to greyish with darker spots. The Common Genet,
Gr. vulgaris, is South European, and just gets into Asia ; it is
also North African. The Genet, an animal " with an appetite
for petty carnage," is one of those smaller Carnivora which are
possibly to be regarded as meant by the word <ya\rj, and appear
to have " functioned " as Cats among the Greeks. So recently as

1 Where it has probably been introduced.

PALM CIVETS 407

the times of Belon we are told (by him) that Genets were
common and tame at Constantinople.

Poiana, containing a single African species, a spotted and
entirely Genet-like animal, has been separated as a distinct
genus. Dr. Mivart, however, holds it to be a Prionodon which
has acquired a Genet-like tarsus.

Arctictis, containing but one species, A. binturong, the
Biuturong, is in some ways an exceptional form. It is a black
arboreal creature of not very wide range in the Oriental region,
with a fully prehensile tail. This feature and its plantigrade
foot with naked sole have led to its being regarded as more allied
to the Arctoidea. It is, however, undoubtedly an ally of
Paradoxurus. The caecum is small, or may be quite absent.
The dentition is I f C 1 Pm f M f . The structure of the
animal has been investigated by Garrod.1

The genus Fossa is a Viverrine confined to Madagascar.
There is but one species, F. daubentoni, the " Fossane." It is
distinguished from Viverra by the presence of two bare spots on
the under surface of the metatarsus in the hind -limb, and by the
absence of a scent pouch. The animal is not much spotted and
striped, but the striping in the young is much more marked.

Of the genus Paradoxurus there are some ten or a dozen species,
belonging entirely to the Oriental region. The teeth are as in
Viverra, but occasionally the molars are reduced to one. The
pupils are vertical. The tail though long is not prehensile, " but
the animal appears to have the power of coiling it to some extent,
and in caged specimens the coiled condition not unfrequently
becomes confirmed and permanent " (Blanford). This fact
accounts for the name Paradoxurus; for a prehensile tail is
hardly to be expected in an animal of the zoological position of
the Palm Civets, and yet its occasional twisting led originally
to the view that it was so. The genus has scent glands. The
dentition is I f C \ Pm f M f . P. niger, the Indian Palm Civet,
is, like other species, not often to be seen in a wild condition. It
is arboreal, and, like other members of the genus, feeds upon a
mixed diet, consisting of all kinds of small Yertebrata and insects,
varied by fruit. Another species, P. grayi, is so distinctly vege-
tarian in its habits that it makes considerable havoc in pine-apple
beds in the Andaman Islands.

1 Proc. Zool. Soc. 1873, p. 196.

408

OTHER CIVETS

Arctogale is another Oriental genus with very small teeth, those
of the molar series being hardly in contact. The soles of the feet are
more naked than in the last genus, and the scent glands, if present,
appear to be small and ill developed. It has also a long tail, and
is arboreal in way of life. There is " nothing particular recorded "
as to its habits. The species are A. leucotis and A. stigmatica.

Closely allied to both the last genera is Hemigale, also
an Oriental genus. It is to be distinguished from Paradoxurus

FIG. 201. — Hardwicke's Civet Cat. Hemigale hardtmcki. x £. (From Mature.)

by having the soles of the feet much less naked, though they are
more so than in Viverra or Prionodon. The coloration of the
species, If. Jiardivicki (a Malayan animal), is very peculiar. The
body is banded with five or six broad transverse stripes, and the
basal portion of the tail is also ringed, an uncommon feature in
the group. A second species of this genus is H. hosei, from Borneo.
It is blackish in colour, but is not a melanic variety of the last.

Nandinia appears never to possess a caecum.1 It is also
peculiar among Carnivora in the non-ossification of the hinder

1 Flower, Proc. Zool. Soc. 1872, p. 683.

xni HERPESTES 409

part of the bulla. It is an African genus, containing two species
which are spotted. The tail is ringed.

Cynogale is at any rate a partially aquatic, short- tailed, web-
footed, reddish brown -coloured Civet, which lives upon fish and
Crustacea, and inhabits the Malay Peninsula, Sumatra, and
Borneo. It has long " moustaches," and is said to have a head
bearing a singular resemblance to the head of the Insectivorous
" Otter " Potamogale. The metatarsus is bald, and the pollex and
hallux are very well developed.

Sub-Fam. 5. Herpestinae. — There are over twenty species of
Herpestes (Mongooses) divided between the Ethiopian and Oriental
regions, one species, H. ichneumon, being also found in Europe.
The fur has a " pepper and salt " appearance ; the feet are
plantigrade. There are five fingers and toes. The pollex and
hallux are small ; the tail is long. The tarsus and meta-
tarsus are usually naked. The Egyptian species " has been
injudiciously denominated the Cat of Pharaoh." It is perhaps
better known as Pharaoh's Mouse. The beast is so far Cat-like
that it will destroy Eats and Mice ; and it has been exported to
sugar plantations for that very purpose. More famous are its
combats with venomous serpents. According to Aristotle and
Pliny the Ichneumon first coats its body with a coating of mud,
in which it wallows, and then with this armour can defy
the serpent. Topsell tells the tale better. The Ichneumon
burrows in the sand, and " when the aspe espyeth her threaten-
ing rage, presently turning about her taile, provoketh the
ichneumon to combate, and with an open mouth and lofty head
doth enter the list, to her owne perdition. For the ichneumon
being nothing afraid of this great bravado, receiveth the en-
counter, and taking the head of the aspe in his mouth biteth
that off to prevent the casting out of her poison." In the West
Indies the animal has been described as fearlessly attacking the
deadly Per de Lance and receiving its bites with impunity ; it is
also added that it will eat the leaves of a particular plant as an
antidote ! The real explanation of the result of these encounters
is of course the agility of the Ichneumon * — fort cauteleuse beste, as
Belon says.

Another species, H. allicauda, is distinguished, as the name
denotes, by its white tail. A species of this genus, H. urva,

1 See also vol. viii. p. 591.

410 AFRICAN MUNGOOSES

sometimes raised to generic rank as Urva, is partly aquatic in
habit ; it feeds upon crabs and frogs, but is quite willing to take
to poultry and their eggs.

Helogale is a genus whose validity appears doubtful (to Dr.
Mivart). It is African, and contains two species.

FIG. 202. — White-tailed Ichneumon. Herpestes albicauda. x i.

Cynictis is an African genus, with five digits on the fore-limbs
and four on the hind. As in Herpestes, the orbit is completely
encircled by bone. There is but a single species, C. penicillata,
which is of a reddish colour and has a bushy tail.

Bdeogale, also African, has the toes still further reduced ;
there are only four on both limbs. The tarsus is hairy and the
tail bushy. They are " very rare animals, and nothing is known
of their habits." It is known, however, that they will kill
poisonous snakes, for Dr. Peters took a Ehinoceros Viper out of
the stomach of one.

Rhynchogale l differs from all other genera of Viverridae, except
Crossarchus and Suricata, in having no groove upon the muzzle.
There are five digits. There is the full Viverrine dentition, with
five premolars in the upper jaw ; but this may be an abnormality.2

Crossarchus differs from the last in only having three pre-
molars on each side of each jaw. It is also African, and there
are several species.

Suricata is the last genus of Viverridae ; it is also African,
and contains a single species, S'uricata tetradactyla, the " Meerkat "
of the Cape. The Suricate has but four toes on each foot ; the
tarsus and the metatarsus are naked below. The body is banded
posteriorly. There are fifteen dorsal vertebrae, and the orbit is

1 The original name was Rhinogale.
2 That it is an abnormality has been recently stated.

HYAENAS

411

closed by bone. The Suricate lives in caves and rock crevices,
and will dig burrows. It is distinctly a diurnal animal, and sits
upon its hind-legs after the fashion of a Marmot. As Buffori
noticed in a tame specimen (thought by him to be a native of
Surinam), the animal barks like a dog. The Suricate is largely
vegetarian, living upon roots.

FIG. 203. — Suricate. Suricata tetradactyla. x £.

Fam. 4. Hyaenidae. — Unlike though the Hyaenas appear to
be to the last family — mainly perhaps on account of size — they
are, nevertheless, very nearly akin to them, more so than to the Cat
tribe. It will be remembered that the striping and spotting of
the Hyaenas is very Genet- and Suricate-like.

There are admittedly two genera among the Hyaenidae,
Hyaena itself with three species,1 and the Aard Wolf, Proteles, with
but one. But Dr. Mivart considers that the Spotted Hyaena should
form a genus apart, Crocuta — a proceeding which was initiated
by the late Dr. Gray of the British Museum. The Hyaenidae
are to be distinguished by the following characters : — There are
generally four toes, always so in the hind-foot. The claws are
non-retractile. The nose and upper lip are grooved. The molar
formula is Pm f M \. The soles of the feet are covered with
hairs upon the tarsus and metatarsus. No scent glands.
Tail short. Dorsal vertebrae more numerous than in other
Aeluroids, i.e. fifteen. The bulla is divided by a rudimentary
septum only.

1 For the anatomy of Hyaenas see- Morrison Watson in Proc. Zool. Soc. 1877,
p. 369 ; 1878, p. 416 ; and 1879, p. 79.

412

CHARACTERS OF HYAENAS

The genera Hyaena and Crocuta, the Striped and Spotted

FIG. 204. — Spotted Hyaena. Crocuta maculafa. x fo,~

Hyaena respectively, are African and Asiatic in range, Crocuta being
limited to South Africa. There is neither hallux nor pollex.
The Hyaenas, stigmatised by Sir Samuel Baker as "low-

FIG. 205. — Striped Hyaena. Hyaena striata. x TV

caste creatures," are mainly carrion feeders. Much Arab supersti-
tion is associated with them. Certain peculiarities in the structure

THE AARD WOLF

413

of the organs of reproduction have led to the belief that a Hyaena
changes its sex every year. Its almost human-sounding howls
are supposed to be a deliberate trap for the unwary traveller.
There is also a legend that in the eye of the Hyaena is a stone
which if placed under the tongue of a man endows him with the
gift of prophecy.

Proteles presents many resemblances to the Hyaenas, but also
certain differences ; by many it is placed in a separate family.
There is but one species, P. cristata, the Aard Wolf of South
Africa. In outward aspect it is very Hyaena-like, the coat being
striped, and the ears, though longer, resembling those of a Hyaena.
There is also a mane. There are, however, five toes on the fore-
feet. The teeth are feebler, particularly the molars, which
are also reduced in number. The skull, as in Hyaena, has no
alisphenoid canal, but the bulla tympani is divided by a
septum. The animal seems to feed largely upon insects,
particularly Termites, and also upon carrion.1

Of extinct Hyaenoids Ictitherium seems to be transitional
between them and the Viverridae. Its dentition, |, \, |, ^, is
that of a Yiverrid, and the feet are five -toed. The upper
carnassial tooth, however, is like that of Hyaena in having a
strong inner cusp. Other extinct genera of Hyaenas are Lycyaena
and Hyaenictis. The genus Hyaena itself goes back as far as
to the Miocene, and occurred in Europe until the Pleistocene.
The Cave Hyaena of this country seems to be indistinguishable
from Crocuta maculata, though it has received the name of H.
spelaea.

Fam. 5. Canidae.' — This family cannot be .divided into more
than five genera, and is universally distributed with the exception
of New Zealand. The auditory bulla is smooth and rounded, and
has internally a very incomplete septum, extending through about
one-fourth or one-third of the cavity. The meatus has a fairly
prominent under lip. The paroccipital process is long and promi-
nent. The mastoid is distinct, though but slightly developed.
The glenoid foramen is large; the condyloid foramen is con-
spicuous, and the carotid canal is deep within the foramen
lacerum posterius. The last three characters are Bear-like ; the

1 Flower, Proc. Zool. Soc. 1869, p. 457.

2 For a general account of the Canidae see Mivart, A Monograph of the Canidae,
London, 1890.

414 DOGS AND WOLVES

form of the bulk is Aeluroid. The teeth vary somewhat in
number, and the following table will serve to indicate the gradual
reduction observable in the number of molars : —

Otocyon I f C \ Pm A M £-°f ±

Canis generally I £ C 1 Pm £ M J-£- *

C^ow IfCi-PmAMf

Icticyon I -»- C £ Pm £ M ^r-l

All the Dogs have a caecum1 of simple cylindrical form. In
C. cancrivorus, C. jubatus, and Nyctercutes procyonides this organ
is straight or only very faintly curved ; in other Dogs it is
coiled into an S-like form, sometimes with an additional twist.
The Dogs have, as a rule, five toes, one being dropped in Lycaon.
The tail is fairly long and distinctly bushy. There is in a
number of species a gland at the root of the tail, the presence of
which can frequently be detected by the wet appearance due to
the oozing secretion. The great majority of existing Canidae
belong to the genus Canis. But certainly three, and more
doubtfully four, other genera can be distinguished.

The genus Icticyon contains but one recent species, the Bush
Dog (/. venaticus, Lund) of British Guiana. The animal has a.
somewhat Paradoxure-like, at any rate a distinctly un-dog-like,
aspect, being longish in the body (some 2 feet long), shortish in
the legs, and big-headed. It is blackish in colour, verging
towards golden brown on the head and back. Sir W. Flower, to
whom we owe our chief knowledge of its structure, characterises it
as like a young Fox, and with the playful manners of a puppy.
The animal appears to hunt in packs and by scent, and has a
reputation for ferocity. Icticyon differs from Canis and agrees
with the Indian Cuon in having but forty teeth, the last molar
having disappeared from the upper and lower jaws. The
caecum, unlike that of the majority of Canidae, is only slightly
curved. The brain, oddly enough, shows a Cat-like peculiarity.
It has been pointed out that in their long bodies and short legs
the genera Cuon and Icticyon resemble the primitive dogs.2

A genus Nyctereutes is usually separated from Canis for the

inclusion of N. procyonides only. The separation is based upon
*

1 Flower, Proc. Zool. Soc. 1879, p. 766.
- Proc. Zool. Soc. 1880, p. 70.

xiii DENTITION OF OTOCYON 415

the strikingly unusual coloration of this Dog. It is a small
animal, with numerous long white hairs dorsally. The face,
chest, and much of the belly are black. Its aspect distinctly
recalls that of a Eaccoon,1 especially in the black patches below

FIG. 206. — Raccoon-like Dog. Nyctereutes procyonides. xj.

the eyes, whence of course the scientific name and the pseudo-
vernacular " Eaccoon-like Dog." It inhabits China and Japan.
As to structure, there is hardly anything that justifies its ex-
clusion from the genus Cams. Garrod, however, mentions the
unusually large size of the Spigelian lobe of the liver.

Wortman and Malkens2 have instituted a genus Nothocyon for
Dr. Mivart's species C. urostictuss and C. parvidens, which are
both South American forms.

The genus Otocyon contains but one species, 0. megalotis, an
African species, ranging pretty widely in that continent (from
the Cape to Somaliland, in sandy districts), and sometimes con-
fused with the Fennec on account of its long ears. Its prin-
cipal structural difference from other Dogs is that there is an
additional molar in each jaw, the molar formula being thus M |-
or even A. Moreover the carnassial teeth are not so pro-
nounced, and Professor Huxley laid especial stress upon the

1 The relationship between the Canidae and the Procyonidae must not be lost
sight of in considering this point of external likeness.

2 Bull. Amer. Mus. Nat. Hist. xii. 1900, p. 109.

3 Proc. Zool. Soc. 1890, p. 98.

416 CAPE HUNTING DOG CHAP.

likeness of some of the cheek teeth to those of the more primitive
Arctoids. The angle of the lower jaw is inflected, a character,
however, which seems to be more general than is usually allowed
among animals not referable to the Marsupials. It is possible
that Otocyon is a persistent Creodont-like form which has
developed in a direction curiously, and in a most detailed
fashion, parallel to the Dogs. If, however, we may assume the
addition of the molar, then this anomalous but not necessarily
untenable conclusion is obviated.

The genus Cuon, or Cyon, has been instituted for the two or
three species of Eastern Dogs (C. primaevus, C. dukkunensis, etc.)
which agree with each other in the constant loss of a molar in
the lower jaw, or, it should be said, almost constant loss, for
the missing tooth is occasionally represented. The latter of the
two species mentioned, the Dhole, is, like its congeners, an
animal which hunts in packs ; it is said to hunt even the
ferocious Tiger, and to be thus one of the few animals which can
face the largest and fiercest of the Carnivora.

The genus Lycaon is a very distinct type, being differentiated
from other Dogs by the possession of only four toes on both
fore- and hind-limbs, and by the dental formula, which is Pm ^
M §. The one species is L. pictus, the Cape Hunting Dog. It is
singularly like a Hyaena1 in general appearance ; the ochraceous
grey ground-colour with black markings and the long ears pro-
duce this likeness. The animal has got its vernacular name
from the habit of hunting in packs. Its range is over a good
part of Africa. The occurrence of this species (or at least genus,
for the name L. anglicus has been used) in caves in Glamorgan-
shire seems to show that it is a comparatively recent immigrant
into Africa. As to its visceral structures, Lycaon^ does not differ
widely from other Dogs. It has, however, no lytta beneath the
tongue. The intestines are thus divided : large, 9 feet 1 inch ;
small, 1 foot 3 inches. This contrasts with the proportions
observable in some other Dogs. While other Dogs have but a
cartilaginous rudiment of the clavicle, Lycaon has a considerably
larger representative of this bone.

The bulk of the Dogs, Wolves, Foxes, and Jackals are thus
left over for inclusion in the genus Oanis. But the numerous

1 Temminck, its original describer, placed it in the genus Hyaena.
2 See Garrod, Proc. Zool. Soc. 1878, p. 373.

ALOPECOIDS AND THOOIDS

417

members of this genus can, according to Professor Huxley, be
sorted into two series by certain cranial characters. The two

FIG. 207. — Fennec Fox. Canis zerda. x £.

series he termed the " Alopecoid " or Fox-like, and the " Thooid "
or Wolf-like. It was suggested that the generic name Vulpes be

FIG. 208. — Prairie Wolf or Coyote. Canis latrans. x

used for the former, and Canis for the second. The characters
which will be dealt with immediately are also to be noted among
VOL. x 2 E

418

CHARACTERS OF FOXES

the Dogs belonging to genera that have already been separated
off. Thus Lycaon is distinctly Thooid. The characters in
question are these: — In the Fox series, the frontal air-sinus of
the Thooids is absent ; the cranial cavity is pear-shaped, without
an abrupt angle coinciding with the supra-orbital sulcus, such as
exists in the other group ; the coronoid process of the mandible
is rather higher and more turned back in the Foxes, while the
depth of the mandible at the level of the first molar is greater.

FIG. 209. — Japanese Wolf. Canis Iwdophylax. x J. (From Nature.)

To the Fox series belong among others the species C. lagopus
(Arctic Fox), C. zerda (the Fennec), C. cha'ma (the Silver-backed
Fox of Africa), C. virginianus (the Virginian Fox), C. velox (the
Kit Fox), and of course the Common Fox of this country. On
the other hand, the Dogs proper (such as C. dingo), the Wolves
(C. lupus, C. pallipes, C. niger), the Japanese Wolf (C. hodophylax),
the Eed Wolf of America (C. jubatus\ the Jackals (C. aureus, G.
anthus, etc.), the Prairie Wolf (C. latrans), and a number of
American forms, such as C. azarae, its close ally C. cancrivorus
( = C. rudis), C. antarcticus, C. magellanicus, etc., are decidedly
Wolves rather than Foxes.

THE BLUE FOX 419

The Arctic Fox, Canis lagopus, is known by its bluish summer
and pure white winter dress as " Blue Fox " and " White Fox "
respectively. It is an inhabitant of the Arctic north ; but in
former days, as its remains show, it descended to such southern
latitudes as Germany and this country. The most southern
point which it now inhabits is Iceland. This small Fox is well
known as being one of the few animals which change their dress
to a complete white in. winter. This change is, however, not
absolutely universal ; and M. Trouessart has even stated that
the supposed change does not exist, but that the colours are a
question of age and sex. This Fox feeds on birds and cast-up
carcases of Whales and Seals ; it is also said to devour shell-fish,
and actually to store up food when abundant for seasons of
scarcity. A Fox has been observed to " carry off eggs in his
mouth from an eider duck's nest, one at a time, until the whole
were removed " ; and in winter to " scratch a hole down through
very deep snow to a cache of eggs beneath." These anecdotes
are told by Sir Leopold M'Clintock ; but others have also
asserted the storing habits of this Fox, which really has only
a short time of the year in which it can catch suitable living
food.

Canis vulpes, the Fox, is not only a native of England, but
extends as far to the east as Egypt, the so-called C. aegyptiacus
being at most a mere variety. Varieties indeed occur in these
islands ; the English Fox being redder, the Scotch greyer. Not
only is the Fox a truly indigenous English beast, but its remains
go back a very long way into past time. Its bones occur in the
Eed Crag, a deposit of Pliocene times. Its prevalence now is no
doubt due to its preservation as a beast of chase. It lives in
burrows, either excavating them itself or taking possession of
those of some other animal ; the Badger suffers in this way, and
is said to be vanquished not by the teeth of the burglarious Fox,
but by its far fouler habits ! It is curious that the expression
"foxing" is not so suitable to this animal as to many others.
The habit of " shamming death " is a widely-spread one in the
animal world, but at least not common with our Fox. The
sagacity of the Fox appears to be a little more proverbial than
actual ; literature teems with its accomplishments. The worthy
Archbishop of Upsala, Olaus Magnus, figured Foxes dipping their
tails in the streams, and then pulling out inquisitive crayfishes

42O JACKALS CHAP.

which had seized upon them. " It is a crafty, lively, and libidi-
nous creature," observed a writer of the last century.

Of Jackals there are many species, both African and Oriental.
Mr. de Winton allows the following list of African species l : —
C. anthus, C. variegatus, C. mesomelas, 0. lateralis. C. mesomelas
is distinguished by the broad black patch in the middle of the
back. These animals do not appear to go in packs as so many
Canidae do ; they live upon carrion, but also rob hen-roosts, and
commit other depredations upon the live stock of farmers. The
" Quaha," C. lateralis, is distinguished from the last by its sharp
bark, and by the obvious side stripe which has given to it its
name. It is curious that it should live in apparent amity with
C. mesomelas, since the habits of the two are identical and would
lead, one might suppose, to a severe struggle for existence, in
which one of the two would disappear. Of Indian Jackals
C. aureus is the most familiar type.

The European Wolf, Canis lupus, was once, but is no longer,

FIG. 210. — Wolf. Canis lupus, x |.

an inhabitant of the British Islands. Their former prevalence
is indicated by many names of towns and villages, such as Ulceby
and Usselby in Lincolnshire, the town of Wolverton, and Woolmer
Forest. In Saxon times Wolves were very abundant ; and even
so recently as the reign of Elizabeth they were to be seen on

1 Proc. Zool. Soc. 1899, p. 533.

xiii HISTORY OF THE WOLF 421

Dartmoor and in the Forest of Dean. In the New Forest they
were hunted in the twelfth century. It would seem that the
last English Wolf was slain some time during the reign of Henry
VII. In Scotland, however, they persisted very much longer.
So recently as 1743 was the last killed. But before this period
they had begun to get exceedingly scarce, for the price of a skin
in 1620 is quoted at £6:13:4. In Ireland Wolves lingered yet
longer; about 1770 is believed to be the date of their final
extinction in that island. The Wolf nowadays is distributed
over the greater part of Europe, Northern Asia, and North
America, the American form not being considered to be distinct
from its European ally. Much legend has collected round this
fierce Carnivore. Aristotle, usually accurate in the main, still
" states more of wolves than experience warranted." Pliny,
unable to sift truth from falsehood, was in this matter " an eager
listener to all old women's tales." Aelian added to his marvels
and asserted that the Wolf cannot bend its head back ; if it
should happen to tread on the flower of the squill it at once
becomes torpid. So the wily fox, fearing his more powerful
enemy, takes care to strew his path with squills ! The conversion
of men into Wolves was a well-known superstition, dating from
Grecian and Roman times ; it formed the basis of much of the
witchcraft persecutions of the Middle Ages and onwards, and has
left its mark in folklore, e.g. the Wolf in " Red Riding Hood."

The Indian Wolves, C. pcdlipes, C. chanco, and C. laniger, are
hardly, if at all, different from C. lupus. Professor Huxley has
remarked upon the likeness of C. pcdlipes to a Jackal, thus bridging
over the very inconsiderable gap that may be held to divide
Jackals and Wolves.

The Dingo, Canis dingo, is an interesting and somewhat
mysterious species of Dog or Wolf. As is well known, it is an
Australian species ; but it does not seem to be certain whether it
was tamed and brought over to Australia by the native races, or
is a true and indigenous Australian species.

The colour of this species varies, but is usually of a reddish
brown ; it is, however, often grey and indeed almost black.
Whether indigenous or introduced, the Dingo is a plague to
Australian settlers, devouring Sheep, which it generally destroys
by tearing out the paunch. It does not as a rule hunt in packs.
The Dingo is stated to feign death with so much persistence that

422

THE DINGO DOG

CHAP.

an individual has been known to be partly flayed before moving.
Dingo remains have been found in river-gravels in Australia

o o

where no human remains have been detected. This argues for its
indigeneity ; but, on the other hand, it has been pointed out that
man himself in the Australian continent goes back a very long

FIG. 211. — Dingo. Canis dingo, x J.

way into time, and may thus still have imported this companion
with him. Anyhow it is quite a wild creature now. Dr. JSTehring,
an expert investigator into the subject of domestic animals, has
stated that the skeleton of the Dingo does not suggest a feral
animal at all but a purely wild race.

The Domestic Dog is usually spoken of as Canis familiaris ;
but to remains in bone caverns the name of C. ferus or C. mikii
has been given. There seems to be no doubt that the Dog was
the " friend of man " in very early times. Its remains have been
met with in Danish kitchen-middens, in the lake-dwellings of
the Swiss lakes, and during the Bronze Age in Europe generally.
But " there are few more vexed questions in the archaeology of
natural history than the origin of the dog." Its remains already
referred to may in many cases have argued its use as food. But
in a Neolithic barrow a Dog was found buried with a woman, the

ANCIENT DOGS 423

skeletons of both being in situ ; this animal was about the size of
a Shepherd Dog. The actual Dog of to-day is divisible into more
than 180 different breeds ; but in a work upon " Natural History "
it would seem out of place to enumerate and characterise these
artificial products. Authors vary in their opinion as to what
stock gave rise to the domestic races of the past and of to-day.
The Jackal, the Bunasu (C. primaevus), the Indian Wolf (C.
pallipes), have been proposed as likely ancestors. It is more
probable that there is much admixture, and that various wild
types have been selected by man in various countries.

Extinct Canidae. — Many of the existing species of Canidae
are also to be found in Pleistocene deposits of the countries which
they now inhabit. A few show a wider range in the immediate
past than in the present. Thus Lycaon (L. anglicus) has been
met with in caves in Glamorganshire, while Icticyon of South
America appears to be congeneric with Speotlios of the Brazilian
caves. The African Otocyon seems to occur in deposits in India.
There are also numerous extinct species belonging to the genus
Canis, which extend as far back as the Pliocene.

The earlier types of Dogs have been placed in different genera.
Cynodictis is an Eocene form from European strata. The skull is
decidedly Civet-like, with a short snout. The fore- and hind-feet
were five -toed, with well -developed pollex and hallux. The
dentition was that of modern Dogs, the molars being two in the
upper and three in the lower jaw. The general aspect of the
creature and the form of the skeleton was much like that of the
Viverrine genus Paradoxurus, of which, as well as of the Dogs,
Cynodictis might have been an ancestor. .

Simocyon of the Upper Miocene serves as the type of a separate
sub-family of Dogs, Siinocyoninae. The skull is short, broad, and
high ; the shortening of the skull affecting the jaws has reduced
the teeth greatly ; the first three premolars are very small, fall
out soon, and are thus often deficient. There are only two
molars in each jaw. This type is of course nowhere near the
ancestral Dog. It is a much -specialised branch of an early type.
Cephalogale is less specialised; there are the usual four pre-
molars. Enhydrocyon is an intermediate form; it has lost one
premolar in each jaw.

Amphicyon, forming the type of another sub-family, Amphi-
cyoninae, though usually placed among the Dogs, presents us with

424 THE BEAR TRIBE

many Bear-like features in its organisation. The feet, for instance,
were plantigrade and five-toed. The ulna and the radius are
specially compared with the same bones in the Bear tribe. The
skull on the other hand is as distinctly Dog-like in form. The
molars are large, broad, and crushing, and Bear-like. The largest
known species, A. gigantcus, is of about the size of the Brown
Bear. Amphicyon is a Miocene genus. Eocene and allied to
it is Pseudamphicyon. This genus has, like Amphicyon, the
complete dentition of forty-four teeth. In the Amphicyoninae
generally the feet are five -toed, the humerus has an entepi-
condylar foramen and the femur a third trochanter. The upper
molars are large.

The closely allied and American genus Daphaenus has also
plantigrade feet, and has in its structure many reminiscences of
the Creodonts. So, too, has the Eocene Uintacyon.

Cynodesmus is closely allied to Cynodictis. It has ancient
features combined with quite modern ones. The skull is
described as being Creodont-like, but the dentition is that of the
rnicrodont modern Dogs. In accordance with its age the cerebral
convolutions of this Dog are much simpler than in existing Dogs,
and the hemispheres do not cover the cerebellum so much.

The Bear-like Carnivora or Arctoidea. — That division of
the Carnivora which is typically represented by the Bears em-
braces three recent families, which are united by a number of
characters. These Carnivora are always plantigrade or nearly so.
They have nearly always five toes. The claws are not retractile,
or at most semi-retractile as in the Panda. In the skull the
tympanic bulla is often depressed, and is not so globular and
obvious as in the Cats. Its cavity is not divided by a septum.
The paroccipital processes are not applied to it. The carnassial
tooth is less emphasised in this group than in the Cats.

These characters, however, have to be used with caution, as
they are hardly universally applicable. A fairly typical Arctoid
bulla is seen in such a form as Cercoleptes. The bulla itself is a
little more swollen than in Ursus, but it is flattened off in the
same way towards the bony meatus. The paroccipital processes,
slightly developed, are at a distance of 1-inch from the posterior
margin of the bulla. In the Kaccoon the bullae are much more
swollen, and the paroccipital processes are closer to them. In
the Marbled Polecat, Putorius sarmaticus. the bullae are fairly

CHARACTERS OF ARCTOIDEA

425

swollen, and there is but little flattening towards the meatus :
the paroccipital processes, though slight, are in contact with the
bullae basally, though their free tips are turned away from them.
Finally, in Ictonyx the bullae are much swollen ; there is but little
flattening towards the meatus, and the paroccipital processes, them-
selves much swollen, are pressed closely against the bullae. The
Mustelidae, therefore, in this as in other characters, approach the
Aeluroids.

There is no caecum, a feature which marks off the Arctoidea
from all Carnivora except the Viverrids Nandinia and Arctictis
(occasionally). The brain is characterised by the possession of

Cur-

FIG. 212. — Section of the left auditory bulla and surrounding bones of a Bear (Ursus
ferox). am, External auditory meatus ; BO, basioccipital ; Car, carotid canal ;
e, Eust'achian canal ; g, glenoid canal ; Sq, squamosal ; T, tympanic ; t, tympanic
(From Flower, Proc. Zool. Soc. 1869.)

ring.

what Dr. Mivart has described as the " ursine lozenge," a tract
about the middle of the hemispheres, defined posteriorly by the
crucial sulcus, and formed by the emergence on to the surface of
the brain of the hippocampal gyrus.

The Arctoidea are very widely distributed. But there are
some curious exceptions. Thus there are no representatives of
the group (as might be expected) in the Australian region ; they
are completely absent from Madagascar; while the true Bears
(family Ursidae) are totally absent from Ethiopian Africa, and
are only represented by a single species, Ursus ornatus, in the
Neotropical region.

It is noteworthy that the Arctoidea never show spots or

426 PROCYONIDAE

cross stripes (save rings on the tail), which are so common a
feature of the coloration of the Cat-like forms.

In bracketing together the three families which are described
in the following pages, emphasis is laid upon a number of un-
doubtedly common features. Palaeontology seems, however, to
suggest that the Mustelidae come nearer to the Viverridae. That
the Bears and Dogs are connected by extinct annectent genera
does not interfere with their present distinctness.

The systematic arrangement of these Carnivora is not easy.
It may be useful, however, to give a method of arrangement for
the convenient placing of the genera.

The most primitive group is perhaps that of the true Bears,
family Ursidae ; for in them the molars are two above and
three below, and have thus not become diminished in number
as in some of the other members of the order. Moreover, the
Bears have lobate kidneys, which character, often occurring in
the young of animals which when adult have smooth kidneys,
may be looked upon as a primitive character. The feet further-
more are completely plantigrade. This family will contain only
three genera, Ursus, Melursus, and Aeluropus.

Next comes the family Procyonidae, in several members of
which one molar is lost below, though in others the more
archaic formula is retained. The kidneys are simple. This
family contains the American genera Procyon, Nasua, Bassariscus,
JBassaricyon, Cercoleptes, and the Old- World form Aelurus.

The third family, Mustelidae, has the molar formula reduced to
^ or ^-. The kidneys are simple except in the Otters. To this
family are assigned the following genera : — Arctonyx, Conepatus,
Meles, Mephitis, 2'axidea, Mydaus, Mellivora, Helictis, Ictonyx,
Mustela, Galictis, Grisonia, Putorius, Gulo, and the aquatic Lutra,
Enhydris, and Aonyx.

Fam. 6. Procyonidae. — This family is mainly American in
range, the genus Aelurus alone being a native of the Old "World.
But Zittel would include with the genera of this family the
Viverrine and Oriental genus Arctictis, a proceeding which is
perhaps hardly admissible, though the occasional absence of a
caecum in that animal is so far in favour of such an alliance.
The largely vegetable nature of its food and its arboreal habits
cause a certain amount of likeness to some of the members of
the present group of Carnivores. The Procyonidae have two

THE RACCOONS 427

molars in either half of each jaw. The carnassial teeth are not
typically developed, and the molars are broad and tuberculate.
The tail is long, often prehensile, and often ringed in the dis-
position of its colour pattern. The alisphenoid canal is absent
save in the aberrant Adurus. Both condyloid and postglenoid
foramina are present. The members of this family are planti-
grade.

The genus Procyon includes at least two species of Raccoon,
the northern form, P. lotor, and the South American, P. cancri-
vorus. To these may possibly be added a third, P. nigripes.
This genus is characterised by the length and the mobility of

the fingers, and indeed it uses its hands greatly. It has no
median groove upon the muzzle, which is found in many other
Arctoids ; the ears are moderately large ; the tail is not long,
being about one-third of the entire length of the animal, in-
cluding the tail. The soles of the feet are naked. Its limbs are
very long (for an Arctoid), and this gives to the animal a
bunched-up appearance when walking. There are four premolars
and two molars on each side of each jaw. There are fourteen
pairs of ribs, of which ten pairs reach the sternum. The latter
is composed of nine pieces.

The first-named species has received its name from the fact-
of which there is abundant proof— that it dips its food into
water. As a matter of fact, the animal frequents the margins of
streams, and hunts in the shallow water beneath stones for cray-

428 " MIMICRY " OF BASSARICYON CHAP.

fish, and it also captures fish. Not only is this animal partially
aquatic, but it can climb well — " they make their homes in trees,
but carry on their business elsewhere." The animal can be
readily tamed, but is a tiresome pet on account of its insatiable
curiosity and its skill in the use of its hands, which enables it
to unlatch doors and generally to pry about everywhere. The
Raccoons are mostly nocturnal creatures.

The genus Bassaricyon l includes two species, both American,
B. alleni being from Ecuador, and B. gabbii from Costa Rica.
They have so much the aspect of a Kinkajou that a specimen,
which arrived at the Zoological Gardens, was presented and
entered as one of those animals. Nevertheless there are many
differences between the two genera. The tail of Bassaricyon is

FIG. 214. — Bassaricyon. hassancyon al'eni. x

riot prehensile, and the animal, as will be seen from Fig. 214,
has a sharper snout ; the brain is more like that of Bassariscus.
The likeness to Cercoleptes can hardly perhaps be regarded as an
example of " mimicry " since the forms are so nearly related, and
the advantage of such an imitation remains to be proved. The
muzzle of Bassaricyon is grooved ; the ears are fairly large ; the
soles of the feet are naked ; there is but a single pair of teats.
There are two molars and four premolars to each half jaw.

The dorsal vertebrae are thirteen in number; nine of the
ribs reach the sternum. The slenderness and convexity of the
lower margin of the lower jaw, as well as the feeble angular
process, distinguish this genus from its undoubtedly near ally
Cercoleptes. The dental formula also is different.

Bassariscus has a ringed tail like a Raccoon, and is also
American in range ; it furthermore agrees with the Raccoon in
1 See Beddard, Proc. Zool. Soc. 1900, p. 661, for anatomy.

xin THE KINKAJOU 429

being nocturnal and mainly arboreal in habit. There are appar-
ently three species, of which B. astutus is the best known, having
been on several occasions exhibited at the Zoological Society's
Gardens, the last examples so lately as 1900. The animal was
for a long time believed to be allied to the Oriental Paradoxures,
and its occurrence in America was therefore puzzling. The real
affinities of the creature were, however, definitely set at rest by
Sir W. Flower, and later accounts of its anatomv have confirmed

FIG. 215. — Cunning Bassarisc. Bassariscus astutus. x \. (From Nature. )

this opinion.1 The vertebrae are more numerous than in Pro-
cyon, and the teeth are slightly different ; otherwise it presents
many likenesses to its nearest ally. The ears are long ; the nose
is grooved ; and the palms and soles are naked.

The Kinkajou, Cercoleptes, is likewise an American Arctoid.
It ranges from Central Mexico down to the Eio Negro in Brazil.
It was at one time confounded, and, considering its external
appearance, not unnaturally, with the Lemurs. Sir E. Owen
dispelled this view by a careful dissection of the creature. Never-
theless, there are certain anatomical features in which it differs

1 Beddard, Proe. Zool. Soc. 1898, p. 129.

430

POTTO AND POTOS

from Carnivora and resembles Lernuroids.1 It has been pointed
out that the form of the lower jaw " much resembles that of the
Lemuroid Microrhynchus." There is, however, no doubt that it.
is rightly placed in the present group. The tail is very pre-
hensile, and the animal is therefore, as might be supposed from this
circumstance, purely arboreal. It has some twenty-eight verte-

3==!

,r:;;-i

FIG. 216. — Kinkajou. Cercoleptes caudivolvulus. x

brae. This genus has a median groove upon the nose. The claws
are long and sharply pointed, and the palms and soles of the
feet are naked. The premolars are three, the molars two. There
are fourteen dorsal vertebrae, of which nine are united to the
nine -jointed sternum by ribs. There is but one species, C.
caudivolvulus, of a uniform yellowish-brown colour.

Nasua, the Coati, ranges from Texas to Paraguay, and has

FIG. 217. — Coati. Nasua rufa. x J.

two species. In Guatemala it reaches a height of 9000 feet on
the mountains. The nose is produced into a short and very

1 It is a curious fact that a native name for the creature is "Pottos" (of. of
course Potto) ; and indeed the generic name Potos seems to have the priority over
Cercoleptes.

xiii AELURUS AND AELUROPUS 431

mobile proboscis, hence its name. The native Mexican name for
the creature is " Quanhpecotl."

The Coati is largely arboreal, and hunts iguanas in large
bands, some of them being on the trees and some on the ground
beneath. It also grubs up worms and larvae, for which purpose
its long snout is suited. The molars of the genus resemble
those of Procyon.

There is not a median groove upon the nose. The palms
and soles are naked. Six teats occur. There are thirteen dorsal
vertebrae. Nasua nasica l and JV. rufa are the best known and
perhaps the only species. The colour of the fur varies a good
deal, and has led to the use of other names for supposed species.

Aelurus, the Panda, is a largish animal found in the south-
eastern Himalayas up to a height of 12,000 feet. It has a
glossy fur of a reddish colour, and a "white somewhat cat-like
face." The molar formula which distinguishes it from the New-
World Arctoids belonging to the Procyonidae, as well as from
its possible ally Aeluropus, is Pm % M |-. The anatomy of the
animal has been described by Sir W. Flower.2 Dr. Mivart has
pointed out that the muzzle though short is upturned in a way
distinctly recalling that of Nasua. The animal inhabits forests,
and feeds almost entirely upon vegetable food. It eats eggs,
however, and insects. Though living to a great extent upon
the ground, it is also arboreal, and has sharp semi-retractile claws.
It is said to be dull of sight, hearing, and smell, and yet with
these disadvantages is also unprovided with cunning or ferocity.
Its habits have been compared with those of a Kinkajou.

Fossil Procyonidae. — In addition to several of the existing
genera, the remains are known of various extinct forms of Pro-
cyonidae. Leptarctus, with one species, L. primaevus, is of Plio-
cene age, but is known only by one ramus of the lower jaw. It
appears to " offer a number of transitional characters between
the more typical Procyonidae and the aberrant Cercoleptes." 3

Fam. 7. Mustelidae. — Contrary to what has been stated with
regard to the habits of the Procyonidae, the Mustelidae are for
the most part "bloodthirsty robbers," and are spread over the

1 " Narica " is generally written, after Linnaeus. But this was, according to
Mr. Alston, probably an error for nasica.

2 Proc. Zool. Soc. 1870, p. 752.

3 See Wortman, Bull. Amer. Mus. Nat. Hist, vi. 1894, p. 229.

432 THE PIG-LIKE BADGER CHAP.

whole surface of the world, with the exception of Australia and
Madagascar. The molar teeth are generally reduced to one in the
upper jaw, and sometimes to one in the lower jaw, which thus
gives "a sort of prima facie resemblance to the feline dentition."
There is no alisphenoid canal; postglenoid and condyloid fora-
mina are found.

Sub-Fam. 1. Melinae. — Of this sub-family there are repre-
sentatives both in the Old and New Worlds.

Meles, the Badger, is exclusively Palaearctic in range.1 Dr.
Mivart says that Meles has a relatively longer dorsal region than
any other Carnivore, and that it is most nearly approached by

FIG. 218. — Badger. Meles taxus. x J.

its allies Ictonyx and Conepatus. The molar formula is, as in
Arctonijx, Mydaus, and Helictis, Pm ^ M ^. The molars differ
from those of any other Carnivore in the much greater size of
the first molars than of the last premolars. The nose is not
grooved ; the soles of the feet are naked. The claws of the fore-
feet are much longer than those of the hind-feet.

The genus Arctonyx is a " pig-like badger " from Hindostan,
Assam, and North China. The epithet " pig-like " is derived from
the long and mobile snout, which is truncated and has terminal
nostrils. It is remarkable for having a part of the palate formed
by the pterygoids, as in Whales and certain Edentata (e.g. Myrme-
copkaga). There are sixteen dorsal vertebrae. A. collaris lives in

1 As a small point of likeness between this Mnstelid and the Procyonidae may
be mentioned the colours of the face. M. anakuma is particularly Raccoon-like.

xin SECRETIONS OF SKUNKS 433

fissures of rocks, or in holes dug by itself. It is a purely nocturnal
beast.

The singular genus Mydaus, containing the species M. meli-
ceps, the Teledu or Javanese Skunk, is an inhabitant of Java and
Sumatra. It frequents the mountains of these islands, into the
soil of which it burrows in search of worms and larvae. There
is but one species, which is " like a miniature badger, of rather
eccentric colours." It is blackish brown, with a yellowish -white
top to its head, and a stripe of the same colour down the back.
It may be distinguished by its elongated snout, obliquely trun-
cated, and with inferiorly- placed nostrils. As to osteological
characters, it has a more oblique symphysis of the mandible
than in any other Carnivore. The secretion of the anal glands
is said to rival that of the Skunk in offensiveness and in the
distance to which it can be propelled.

Sub-Fam. 2. Mustelinae. — Representatives occur in both the
Old and JSfew Worlds ; but the genera and even the species are in
one or two cases common to both.

Galictis Barbara,1 the Tayra, is a brown, elongated, and Weasel-

FlG. 219.— Tayra. Galictis barbara. x f .

like animal from Mexico and South America. As is the case with
the Weasel, it is sometimes gregarious, a herd of twenty having
been observed. The soles of the feet are naked, and the molar
formula is Pm •§. M ^. In these characters the Grison (G. vittata)
agrees with G. barbara ; but it has been referred to a different
genus, Grisonia.

The Grison, " this savage and diabolical-looking weasel," as

1 See Trans. Zool. Soc. ii. 1841, p. 201.
VOL. X 2 F

434 CHARACTERS OF GRISONS

Mr. Apliri terms it,1 is known also as the " Huron." It almost
rivals the Skunk in the power of the odour which it can emit
when enraged. A trapped specimen was placed in a cage 50
yards or so from the house, and even at this distance it was
disagreeably easy to tell when any one visited the animal — at
least when the wind set in the right direction. It is greyish
yellow above and blackish beneath, presenting, as has been

FIG. 220. — Grisou. Grisonia vittata. x A.

remarked, a curious similarity to the Eatel. The nose of this
animal is destitute of a median groove, which is present in the
Tayra ; the soles of the feet, however, are naked as in that animal,
and it is nearly plantigrade in walk. It differs also from Galictis
in having sixteen 2 instead of fourteen dorsal vertebrae. Eleven
of the ribs reach the sternum. Considering the differences that
exist between some other genera of Arctoids, it may be fairly
allowed that a genus Grisonia is tenable.

G. allamandi is darker coloured than the Orison, with a white
band from the forehead to the neck. Mr. T. Bell described a
tame individual as eating eggs, frogs, and even a young
alligator.

A third genus of this group has recently been founded by Mr.
Oldfield Thomas 3 for a small African animal, which is Grison-like
in its coloration. The name given to the genus, Galerisciis,
is intended to suggest its likeness to the Grison (Galera or
Grisonia'). The chief distinctive feature of this genus, whose
skeleton is not yet known, is the presence of only four digits on
each limb ; the pollex and the hallux being entirely absent. The
ears of this Grison are short.

1 Proc. Zool. Soc. 1894, p. 306.
'- I found fifteen. 3 Ann. Nat. Hist. (6) xiii. 1893, p. 522.

xin FEEDING OF THE GLUTTON 435

The genus Mustela includes the Martens and Sables, which are
distinguished from the following genus by the molar formula,
which is Pm£ M-|. The same character separates them from
Galictis, and also the generally hairy under surface of the feet.
En more southern latitudes, however, the palms are sometimes
naked. The nose is grooved, and the ears ace short and broad.
The genus is widely distributed, being common to the Old and
Xew Worlds. In the Old World it extends from Europe to Java,
Sumatra, and Borneo. The largest species of the genus is the
American Pekan, an animal which may be 46 inches in length,
including the tail. There are two species of Sable, one European
(J/[ zibellina), the other American.

The only British species of the genus is the Pine Marten,
M. rnartes. It is dark brown, with a brownish-yellow throat, and
reaches a length of some 1*7 inches, with an eight-inch tail. It
is getting rare, but is still fairly common in the Lake country.
The animal is largely arboreal in habit, whence the vernacular
name. It is also called Marten Cat. The allied 3f. foiiia, the
Beech Marten, has been stated to be, but apparently is not, an
inhabitant of these islands. The colour of the animal is a rich
brown. It has small eyes and ears and a short tail. The palms
of the hands and the soles of the feet are hairy ; the muzzle is
naked, and has a groove as in Cercoleptes, etc.

The Glutton, G-ulo, is a well-marked genus, containing but
one species, which is circumpolar in range. The dentition is
Pm ^ M ^. The ferocity but not the voracity of this animal
appears to have been exaggerated. It mainly feeds on carcases,
and is not really a successful hunter. As to the carcases, Olaus
Magnus tells in straightforward language the way in which the
animal dilates in size during a meal, and presently, after follow-
ing the practice of the ancient Eomans, returns to the banquet :
" Creditur a natura creatum ad ruborem hominum qui vorando
bibendoque vomunt redeuntque ad mensam " !

This is one of the few land animals which ranges completely
round the pole. There is no difference to be noted between the
Old- World and the New-World specimens. It is now an entirely
northern form, but in Pleistocene times it reached as far south as
this country. The fossil species seems to be Gulo luscus, and to be
quite indistinguishable from the living forms.

Putorius, the genus which embraces the Weasel tribe, contains

436 BRITISH ARCTOIDEA

many species known popularly as Weasels, Ermines, Stoats, Ferrets,
Polecats, Minks, and Vison. Not only is the genus common to
both Old and New Worlds, but in a few cases the species (e.g.
P. erminea) range from Asia to America. The molar formula is

~

FIG. 221. — Polecat. Mustela putorius. xJ.

Pm § M ^. The form of the body is an exaggerated one, the
length of the trunk to the limbs being very great. The feet
are more or less hairy beneath, and the animals are digitigrade.
The nose is grooved. The dorsal vertebrae vary from thirteen
to sixteen.

There are four British representatives of this genus : —
The Polecat, P. foetidus, is a dark brown-coloured animal. Its
total length is about 2 feet, of which the tail occupies some 7
inches. It is a species banned by the gamekeeper, and hence is
approaching extinction in this country. It is excessively blood-
thirsty, as are apparently all the members of this genus, and kills
out of mere wantonness. The Ferret is simply a domesticated
variety of the Polecat.

The Stoat or Ermine, P. erminea, is reddish brown above,
white beneath. In winter, in certain localities, it becomes white
with the exception of the black tip of the tail. This colour-
change bears some relation to the degree of latitude. It is
universal in the north of Scotland, rare in the south of England.
As is the case with some other animals that generally change

THE IRISH STOAT 437

their colour in the winter, there are individuals which seem to
have lost the power of change, and others which change in an
apparently capricious manner, not influenced by season or cold.
Like so many other animals, the Stoat appears at times to
migrate, which it does in large parties. Such parties are
said to be dangerous, and will attack a man who crosses
their path.

The Weasel, P. vulgaris, has much the same colour as the
Stoat, but is a smaller animal ; it differs also by undergoing
no seasonal change. It is equally agile and ferocious, and
ought to be encouraged, as it vents its ferocity largely upon
Voles and Moles, which it can pursue underground. Like other
species of Putorius, it seems to kill its prey by biting through
the brain-case.

The fourth British species is the recently -described Irish
Stoat, P. hibernicus. It is somewhat intermediate between the
last two.

Poecilogale is a genus recently instituted by Mr. Thomas for
a small South African Weasel, P. albinucha, coloured like the
Zorilla, i.e. with whitish stripes upon black, but differing in its
reduced molar formula, which is Pm § M ^ or -|.

Lyncodon1 is thought to be more doubtful; it is South
American (Patagonian), with the same molar formula as the most
reduced forms of the last genus, i.e. Pm f M ^. The ears are
short and almost invisible ; the claws of the anterior limbs are
long, those of the hind limbs short. It is not quite certain that
it is not "an aberrant southern form of Putorius brasiliensis."
That its distinction is justifiable appears to be shown by the
discovery in the same region of a fossil species, L. luganensis.
Matschie places it near Galictis.

The Katel, Mellivora, is common to India and West and South
Africa. It is a black animal with a grey back and grey on the
top of the head, the contrast of colour suggesting a dorsal
carapace. It runs with a swift trot. The animal lives much on
the ground, but can climb trees. It is exclusively nocturnal in
its habits. It has the reputation in India of feeding upon dead
bodies, a view which has probably no foundation in fact save
that it can burrow. The molar formula is Pm f M ^. There are
fourteen dorsal vertebrae. The African and Indian species are

1 See Matschie, SB. Ges. Naturf. Berlin, 1895, p. 171.

433

HELICTIS AND 1CTONYX

hardly to be distinguished from each other. The ears are very
minute. The tail is short. The muzzle is rather pointed, and
the soles and palms are naked.

FIG. 222. — Ratel. Mellivora capensis. x J.

The structure of Helictis has been described by the late
Professor Garrod,1 as well as by Sir W. Flower in his general
account of the Carnivorous skeleton. The animal, which is a
native of East Asia, is sometimes gaily coloured. If. subaurantiaca,
the species dissected and figured by Garrod, is a varied black and
orange. The genus is arboreal, and the tail may be moderately
long and bushy. The ears are small ; the nose is grooved ; the
palms are naked, but the soles of the feet are hairy. There are
fourteen dorsal vertebrae. The molar formula is Pm £ M }.

The Zorilla, Ictonyx, is the last of the Old-World genera of
Melinae. It is African, ranging from the tropical parts of the
continent to the Cape. " In colour and markings," remarks Dr.
Mivart, " as well as in the odour of the secretion of its anal
glands, the one or two species which form this genus resemble
the skunks ; so much so that did they inhabit the same region,
and were they devoid of an offensive secretion, they would
certainly be said to mimic the skunks." The molar formula of
the genus is Pm | M |. There are fifteen dorsal vertebrae. The
nose is grooved and the soles partly hairy.

The American Badger, Taxidea, is a burrower of omnivorous
tastes, and correlated with the former habit are the immense
1 Proc. Zool. Hoc. 1879, p. 305.

xiii SKUNKS 439

claws of the fore-paws. It is North American, but gets into
Mexico. The molar formula is as in the American genera
Mephitis and Conepatus, and as in the Old-World Ictonyx, and
it thus differs from that of Meles. Besides the great size of the
claws upon the hand, which are larger relatively than those of
any other Carnivore, the genus Taxidea is to be distinguished
from all Arctoids (indeed, from all Carnivora) except Mydaus, by
the fact that the pelvic limb is of the same length as the
pectoral. The muzzle is furry except at the very extremity ; this
is grooved. The animal is carnivorous, subsisting upon the
following very varied kinds of food — " Spermophiles, Arvicolas,
birds' eggs, and snails, also honey-comb, wax, and bees."

The Skunk, Mephitis, is an American animal with several
species, which range from North to Central America. The black-
and-white colour distinguishes the genus, which is furthermore
marked by the fact that the third digit of the hand is relatively
longer than in any other Carnivore except Taxidea. The soles are
partly hairy. It is a terrestrial fossorial animal with well-known
powers of protecting itself from aggression. But nevertheless the
Skunk has its enemies, and is not quite so unmolested as is some-
times popularly supposed. The Puma, Harpy Eagle, and the
Great Horned Owl will at least occasionally attack and devour
it. The molar formula is Pm § M ^. There are sixteen dorsal
vertebrae.

Gonepatxs is a more southern form of Skunk, extending down
into South America. Its dentition is like that of Mephitis save
for the loss of an upper premolar. This genus, which has been
further subdivided, differs from Mephitis in the fact that the
soles of the feet are wholly naked, whereas in Mephitis those of
the hind -limbs are partially hairy. It has no groove on the
nose. Its tail is shorter than that of Mephitis. This Skunk
has the same habits as the last. In certain parts of South
America the animals are so abundant and their odour so powerful
that in the evening there is generally a recognisable smell about.
This is said to be good for the headache !

Sub-Fam. 3. Lutrinae. — Of this sub-family there are at least
two genera. Enliydris (Latax)? the Sea-Otter, is confined to the
shores of the North Pacific. It is more purely aquatic than are

1 Lydekker, " Note on the Structure and Habits of the Sea-Otter (Latax lutris),"
Proc. Zool. Soc. 1895, p. 421 ; and -ibid. 1896, p. 235.

44O OTTERS CHAP.

other Otters. Specimens have been seen swimming fifteen miles
from land. The gait of the creature when on land is suggestive
of a marine animal ; the webbed hind-feet are doubled back upon
the knuckles during progression upon land, and locomotion is
effected by a series of short springs from these feet; the Otter
does not walk " in ordinary acceptance of the term." The tail
is flattened, being twice as broad as it is thick, and ends in a
bluntish point. Enhydris feeds mainly upon crabs and sea-urchins,
but also upon fish. Its dental formula is peculiar by reason
chiefly of the reduction of the lower incisors. The formula runs
as follows : I f C \ Pm f M £.

The molar teeth of this creature, in accordance with its diet,
have lost the sharp points of the Mustelidae in general ; the
crowns are flattened, and the tubercles very blunt. In this it
contrasts with Lutra, and presents some resemblance to the
Crab-eating Eaccoon, Procyon cancrivorus ; but the teeth are still
further blunted. Enhydris feeds largely upon sea-urchins and
shell-fish, and needs blunt teeth for the crushing of the hard
shells of its prey. It is interesting to notice that the habits of
this animal have been altered by the interference of man. The
creature has been hotly pursued for a long time on account
of its valuable fur. Instead of feeding and breeding upon
the shore in places readily accessible to its pursuers, the Sea-
Otter has now taken to the open sea in a greater degree. It
utilises masses of floating seaweed for those purposes, and hunts
for its food in the deeper water at a greater distance from the
shore. In conjunction with the increasing rarity of the Sea-
Otter the price of its skin has enormously increased : whereas in
1888 the average price per skin was £21 : 10s., the value of a
fine skin now is at least £100, and as much as £200 and even
£250 has been given. The animal is captured by netting and
by clubbing and spearing.1 From the Miocene Siwalik beds re-
mains of an allied form, Enhydridon, have been obtained, whose
teeth are somewhat intermediate in their crowns between Lutra
and Enhydris.

Lutra, including the Otters, is widely distributed. Both
manus and pes are webbed. The ears are small and hairy.
The nose is not grooved, and the naked part is very circumscribed ;

1 See an article by Mr. Lydekker in Knowledge, April 1898, from which many
of the above facts have been taken.

RANGE OF OTTERS

441

the claws upon the hind-feet are flattened and somewhat nail-
like. There are about ten species, but of course, as is so
universally the case, a great many more names have been given.
The molar formula is like that of Enliydris save that there is an
extra premolar in the upper jaw. There are fourteen pairs of ribs,
of which eleven pairs reach the ten-jointed sternum. The caudals
are twenty-three. The Cape Otter, the " clawless " Otter, has been
separated as a genus Aonyx. So too has the South American
Pteronura brasilien&is. But in neither case is the separation allowed
by Mr. Thomas in a recent revision of the genus.1 The latter
species has the reputation of being very fierce, and is known in

FIG. 223. — Otter. Lutra vitlyaris. x J.

Uruguay by the name of " Lobo de pecho bianco." The British
species, L. vulyaris, reaches a length of 2 feet or so, with a tail of
1 6 inches ; it ranges over the whole of Europe and a large
portion of Asia. This Otter often burrows in the banks of the
streams which it frequents ; and in the burrow in March or April
the female brings forth her young, three to five in number. It
will also frequent the sea-coast.

Fossil Mustelidae. — Besides a number of the existing
genera there are fossil members of this family which cannot be
referred to existing genera. These latter extend back into time
as far as the Eocene. Stenoplesictis, one of these Eocene forms
referable to the sub-family Mustelinae, is to be distinguished

1 " Preliminary Notes on the Characters and Synonymy of the different Species
of Otter," Proc. Zool. Soc. 1889, p. 190.

442 BEARS CHAP.

from living Mustelines by its comparatively long legs. In this
genus as in several others there are two upper molars.

Fam. 8. Ursidae. — This family is nearly universal in dis-
tribution, and consists of but three genera, Ursus, Melursus, and
Aeluropus.

Ursus has the palms and soles naked except in the Polar
Bear, which needs a furry sole to walk with ease upon ice
surfaces. The ears are fairly large, and the nose may or may
not be traversed by a median groove.1 The molar formula2 is
Pm ^ M -|. The brain is naturally (because of the size of the
animals of this genus) richly convoluted. The lobate kidneys
have already been mentioned in defining this family (see p. 426).

A very large number of species of Bears have been described.

-

FIG. 224. — Himalayan Bear. Ursiis tibetanus. x T\.

But it is the opinion of Mr. Lydekker 3 and of others that many
of these are really to be referred to the European Brown Bear ;
in this event the Grizzly of North America, the Isabelline Bear,
the Syrian Bear, a Bear from Algeria, the Kamschatkan and
Japanese Bears, besides the extinct Ursus fossilis of Pleistocene
caves, are to be regarded as slight modifications of Ursus
arctos. On the other hand, the great Cave Bear, U. spelaeus,

1 Even apparently in the same species.
- The number of premolars is reduced in the Polar Bear.
"The Blue Bear of Thibet," etc., Proc. Zool Soc. 1897, p. 412.

SLOTH BEAR

443

aud the Thibetan Blue Bear ( U. pruinosus) are distinct species,
not to be confounded with U. arctos. Neither, of course, are the
Peruvian U. ornatus and the Sun Bear, U. malayanus.

The Polar Bear has even been placed in a separate genus,
Thalassarctos, a proceeding which is quite unnecessary. The
white colour of this Bear tends to become browner with age. It
is one of the few mammals which extend right round the pole ;
the Polar Bear is of course a purely Arctic animal. The chief food
of the Polar Bear is Seal. Out of thirty Bears examined, Mr.

FIG. 225. — Malayan Bear. Ursus malayanus. x ^\.

Koettlitz found that only fifteen had animal remains in their
stomachs, and these remains were invariably Seal. The animal
apparently hunts by scent rather than by sight or hearing, both of
which senses seem to be somewhat dull. The males and females
wander separately, except of course during the breeding season.
The Bears dig holes in which they may remain for some time,
but there is no hibernation. In Pleistocene times, the Polar
Bear extended as far south as Hamburg. The female has four
mammae, pectoral in position.

Mdursus includes only M. labiatus, the Sloth Bear of India.
This animal has an upturned snout, which .is described as closely
resembling that of My dans, the Teledu. The snout has no groove.

444 PERE DAVID'S BEAR CHAP.

All Bears are largely vegetarian and insect feeders ; but this Bear
is especially so. It delights in the nests of Termites, and its
energy in destroying these hills for the sake of their inhabitants
is so great that the name of " sloth " appeared to Sir Samuel
Baker to be an entire misnomer.

Aeiuropus, a rare Carnivore with but one species, A. melano-
leucus, is not inferior in size to the Brown Bear, and is dis-
tinguished by its largely white coloration. It was discovered in
the mountains of East Thibet by Pere David, and described by
Milne -Edwards1 as a distinct and new genus, the discoverer
himself having named it as a species of Ursus. It is a vegetable-

FIG. 226. — Aelnropus inelanohucus. J*.

feeding creature and bulky in form, with a rudimentary tail and
a short broad head ; in fact, more like a Bear than a Procyonid
(with which group it is placed by some). The width of the
head, however, is greater than in any other Carnivore ; it is
most closely approached in this by Aelurus and by Hyaena.
The molar formula is Pin | M f . The soles are hairy. There
is no alisphenoid canal. The molars are especially large and
multicuspid.

Fossil Ursidae. — The genus Ursus itself goes back to Plio-
cene times. The well-known Cave Bear, Ursus spelaeus of

1 Nouv. Arch. Mus. vii. 1872, full. p. 92; and Jtecherches pour servir & I'histoire
naturelle des Mammiferes, 1868-1874, p. 321. This genus has quite recently
(Lankester, Trans. Linn. Soc. viii. 1901, p. 163) been definitely referred to the
Procyonidae.

FOSSIL BEARS 445

Pleistocene times, was one of the commonest of Carnivorous
creatures during the very early times of the present era. It
was as huge as a Polar Bear or a Grizzly. The skull is remark-
able for the fact that the first three premolars, which are small
in all Bears, dropped out early in life. An immense number of
names have been given to what are in all probability the same
species as this Cave Bear of remote antiquity.

Hyaenarctos is the oldest genus of true Ursidae. It goes
back into Middle Miocene times, and ranged over Europe and
North Africa.

Arctotherium is an American genus of Pleistocene times.
The likeness of some of the extinct Canidae to Bears has been
already commented upon.

CHAPTER XIV

CARNIVORA (CONTINUED} — PJNNIPEDIA (SEALS AND WALRUSES)—

CREODONTA

SUB-ORDER 2. PINNIPEDIA

Tins group includes the Seals, Sea -Lions, and Walruses,1 all
aquatic and, for the larger part, marine creatures. Being aquatic
they have to some extent acquired a fish-like form, though not
so completely as have the Whales and even the Sirenia. This
is most complete so far as the group is concerned in the Seals,
where the hind-limbs have become soldered to the tail and are
inefficient as walking legs, where the external ears have vanished,
and where the general shape of the body is tapering and thus
fish-like. The Walruses and Sea-Lions are less modified in this
direction ; in the latter (not in the former) the external ear,
though small, is persistent, and the hind-limbs are capable of
being used as organs of progression upon dry land. The general
characters applicable to the Carnivora, given upon a previous
page, apply to the Pinnipedia.

The characters confined to the Pinnipedia as a whole are
mainly these : — The greater part of the limbs are enclosed
within 'the skin, the hands and feet are fully webbed, and
there is a tendency for the nails to disappear, and for the
^, phalanges to increase in number — characters which are clearly
• not diagnostic of the order but correlated with an aquatic
life, since they reappear, and are indeed exaggerated, in the
Cetacea. The teeth are peculiar in that the milk dentition is
feeble and is early shed. This, as it were, undue emphasis upon
one of the two sets of teeth is another likeness to the Whales,

1 For the genera of Pinnipedia see Mivart, Proc. Zool. Soc. 1885, p. 484.

CHAP. XIV

SKKLETON OF SEAL

447

where, however, it is the
milk dentition that is
most pronounced, the
" permanent " being
feeble and very early
shed. But the dentition
of the Pimiipedes pre-
sents other likenesses to
the Cetacea, which are,
it must be remembered,
regarded by some as a
modification of the Carni-
vorous stock, in which
case, of course, the like-
nesses may be genetic
rather than due to
adaptation in the two
cases. There is a dis-
tinct tendency towards a
homodont series, the
grinding teeth being
often very simple, and
the very distinct carnas-
sial tooth of many terres-
trial Carnivores being
absent. Finally, the
number of the back teeth
shows some signs of being
on the increase ; and
Professor Klikenthal has
found that this increase
is due to the division of
existing teeth. Here is
a point of likeness to the
many teeth of the typical
Toothed Whales. Dr.
Xehring found in several
examples of Halichoerus
grypus the normal five
back teeth increased to

448 TEETH OF SEALS CHAP.

six, and the additional molar was at the end of the series, thus
suggesting a lengthening of jaw coupled with an increase in
number of teeth.

The incisor teeth of the Pinnipedia differ from those of the
land Carnivora in that there are nearly always fewer than ^, at
least in the adult animal. In possessing lobulated kidneys the
Pinnipedia differ from all terrestrial Carnivores except the Otters
and Bears — a significant fact.

In the characters of the skeleton the Pinnipedia show many
peculiarities. The cranial part of the skull is proportionately
to the facial part greater than in terrestrial Carnivora ; there is
no lachrymal bone, and the orbit is to some extent defective in
ossification. The alisphenoid canal, so important a feature in
the Carnivora, may be present or absent. It is present, for
example, in Otaria jubata.1 This genus also has the more

FIG. 228. — Patagonian Sea-Lion. Otaria jubata. x Js.

primitive small and rugged tympanic bullae, which are inflated
and more Cat-like in others. The vertebrae show an interesting
Creodont peculiarity in the complex interlocking arrangements of
the zygapophyses of the dorsal vertebrae. The ossicula auditus
differ from those of their terrestrial allies in their large size and
massive growth. In this they have come to be like those of the
Whales and Sirenians.

There is no doubt about their close resemblance £o the

1 Murie, Trans. Zool. Soc. viii. 1874, p. 501.

xiv ORIGIN OF MARINE CARNIVORA 449

terrestrial Carnivora, but the question is, to which group of
Garni vora have they the most likeness. The semiaquatic Otter,
and the still more thoroughly aquatic (marine) Enhydris,
suggest an affinity in that direction. The long body and short
legs of the Otter, which is more thoroughly at home pursuing
fish in the streams than in waddling clumsily upon the banks
of the streams, seem to require but little external change
to convert it into a small Seal, while, the long and completely
webbed hind digits of Enhydris are even more like those of a
Pinniped. The Sea-Lions, in which the external ear has been
preserved, and in which the limbs have not become so entirely
useless for progression on the land as they have in the Seals,
seem to be the intermediate step in the evolution of the latter.
This, however, is not the opinion of Dr. Mivart, who, without
definitely committing himself on the point, presents some evidence
for the assumption that the marine Carnivora are diphyletic.
This double origin, however, is not from two groups of the
terrestrial Carnivora. Dr. Mivart, in common with many others,
holds that the Pinnipedia as a whole are undoubtedly nearer to
the Arctoidea than to either of the two remaining sections of
the sub-order. One of the most striking structural characters
in which they show this resemblance is the brain ; the peculiar
Ursine lozenge, already treated of as so distinctive a character of
the Arctoidea, is repeated in the Pinnipedia.

There are, however, other points of likeness which seem rather
to point to a Creodont origin. Patriofelis is a genus that from
more than one side may be looked upon as a possible ancestor of
these animals. The Creodont peculiarity of the vertebrae has
already been referred to. It may be added that the facial part
of the skull is small in Patriofelis, which appears, moreover, to
have had an alisphenoid canal. A very remarkable resemblance
lies in the structure of the astragalus. This is not deeply
grooved on the tibial facet as it is in Fissiped Carnivora. This
might be held to be an instance of degeneration in the aquatic Seals,
which do not use their limbs as walking organs. But Professor
Wortman * has pointed out that in the Sea-Otter, which is entirely
aquatic, the groove exists and is plain. The likeness offered to
the Seals by the spreading feet of Patriofelis is noticed under the
description of that genus.2

1 Bull. Amer. Mus. Nat. Hist. vi. 1894, p. 129. 2 P. 456 Iwlow.

VOL. X 2 G

450

CHARACTERS OF SEA-LIONS

Fam. 1. Otariidae. — The family Otariidae l is no doubt the
least modified of the aquatic Carnivora. It is rational, therefore,
to commence the survey of the group with this family. They
have preserved, as already noted, the independence of the hind-
limbs ; the external ear is present, though small ; there is an
obvious neck, and the nostrils are at the end of the snout, as in

i v - "< o.

"^-•^ 'V->

FIG. 229. — Cape Sea-Lion. Otaria pusilla. x^.

terrestrial creatures generally. The nails are small and rudi-
mentary, save those upon the three middle digits of the foot. It
is a singular fact that among the Otaries the angle of the lower
jaw is " inflected as much as in any Marsupial." The literature
relating to this family is great, and it seems difficult to reconcile
the very varying opinions as to how many genera ought to be
admitted. Mr. Allen arranged the nine species which he allowed

1 See especially Allen, North American Pinnipedes, 1880.

xiv FUR SEALS AND HAIR SEALS 451

in six genera ; but more generic names have been proposed. At
the other extreme stands Dr. Mivart, who speaks of only one
genus, Otaria ; of this genus the number of species is by no
means agreed upon. There can, however, be no doubt of the
distinctness of the Northern Fur Seal, 0. ursina (the " Seal " of
commerce and the cause of international complications), of the
Patagonian Maned Sea-Lion, 0. jubata} of 0. pusilla of the Cape,
of the California!! 0. gillespiei, of 0. hookeri from the Auckland
Islands, and of four or five others. The range of the genus is wide,
but is mainly Antarctic. It is usual to speak of " Hair Seals "
and " Fur Seals," the latter being the species which produce the
" sealskin " of commerce. The difference is that in the Fur Seals
there is a dense, soft under-fur, which is wanting in the other group.
It is, however, impossible to make this character the basis of a
generic subdivision. There is a Fur Seal, 0. .nigrescens, in South
America as well as the more widely-known northern form.

Fam. 2. Trichechidae. — This family contains but one genus,
Trichechus, the Walrus or Morse, or Odobaenus, as the more
correct term seems to be. It is a tiresome result of accurate
conformity with the rules of priority in nomenclature that the
name Trichechus should be applied to the Manatee. There is but
one species of Walrus, though it has been attempted to show that
the Pacific and Eastern forms are different. The animal is Arctic
and circumpolar. The Walrus is characterised by the enormous
canines of its upper jaw, which form the well-known tusks and
reach a length of 30 inches. The animal can progress on land
like the Sea-Lions ; but, as in the Seals, there are no external
ears, though there is a slight protuberance above the meatus
auditorius. The strong bristles upon the upper lip are as thick as
crow quills. The pectoral limb has nails, but these are small, as
in the Sea-Lions. The under surface of the manus has a warty pad,
which cannot but assist 2 in maintaining a foothold upon slippery
ice. The hind-limbs have longer nails, which are still diminutive
and subequal in size. There is no free tail. The liver of this
animal is much furrowed, but not so much so as in Otaria, though
more so than in Phoca. The kidneys are of course lobulate, as in
the other aquatic Carnivores. The milk dental formula appears to
be I f C 1 Pm + M f . In the adult the formula3 is I £ C £ M f .

1 Murie, Trans. Zool. Soc. vii. 1894, p. 411. 2 Cf. the Dugong, p. 336.

3 Kiikenthal, Jen. Zeitschr. xxviii. 1894, p. 76.

452

SEALS IN THE CASPIAN

Fam. 3. Phocidae. — The true Seals have no external ears,
and the nostrils are quite dorsal in position as in other aquatic
animals, such as the Crocodile. There is obviously an approach
to the conditions characteristic of the Whales. The hind-limbs
are useless for locomotion on land. They are bound up with the
tail, and form functionally merely a part of the tail. In this
family there are, at any rate, eight genera.

Phoca and Halichoerus are not very wide apart from each
other. In both there are five well-developed claws on feet and
hands. They are British, and generally Arctic and temperate
in range. For some reason or other the late Dr. Gray placed
Halichoerus in the same sub-family with the Walrus ! Phoca is

FIG. 230. — Common Seal. Phoca vitulina. x$-. (From Parker and Harwell's Zoology.)

not only marine, but is found in the Caspian and in Lake
Baikal. Their existence in those inland seas is believed to be a
vestige of a former connexion with the sea. Halichoerus grypus
is a large seal 8 feet in length when full grown. Its colour is
yellowish grey, with darker grey spots and blotches. It is not
uncommon on the shores of our islands, particularly of the Hebrides
and Argyllshire. The commonest Seal is Phoca vitulina, not more
than 4 to 5 feet long, and of the same spotted coloration as the
last. This Seal has, however, a much wider distribution, being
Arctic as well as British, American, and North Pacific. A curious
fact about this Seal is that it is not impatient of fresh water ;
not only will it ascend rivers, but it will live in inland lakes. It
is said to be especially sensitive to musical sounds. P. hispida
is British, but a rare visitor to our islands. It is essentially an
Arctic species. The Harp Seal, P. groenlandica, is so called on
account of a harp-shaped black bar in the males, which starts at

GENERA OF SEALS 453

the shoulders and extends to the thighs. Like the other Seals
mentioned, the young are white when first born. As may be
inferred from its scientific name this species is also Arctic in
range. It is also a rare visitor to these shores.

The genus Cystophora is the only other genus of which there
is a British representative. It is called the Hooded Seal on
account of an inflatable sac upon the face, with which it is said
to attempt to terrify its enemies. The genus has an incisor less
in each half of each jaw than Phoca and Haliclioerus. Its
formula is 1 1- while these genera are both -|. C. cristata is a
large species reaching a length of 10 feet. The colour of the
back is dark grey with deeper coloured spots. A few individuals
only have been recorded from our coasts.

Stenorhynchus ( = Ogmorhinus) is an Antarctic genus. The
hind -feet are clawless. The incisors are §. The molars have
an additional cusp, i.e. three in all.

The genus Leptonyx with but one species, L. weddelli, is purely
Antarctic in range. Like the last genus it has two incisors, and
has but rudimentary claws upon the hind-feet ; the first and fifth
toes moreover are the longest. The genus chiefly differs from
the last in the simple conical crowns of the molars, which have
not the additional cusps of Stenorhynchus.

OmmatopJioca is another Antarctic genus with but a single
species, 0. rossi. In this genus the hind-feet have no claws, and
the first and fifth toes are longer than the others. The claws of
the fore-feet are rudimentary. The immense size of the orbits
gives the name to the genus. There are two incisors, and the
molars are all very small.

Monachus is a northern genus inhabiting the Mediterranean
and the Atlantic in the vicinity of Madeira and the Canary
Islands. It has rudimentary nails upon both pairs of feet.
The first and fifth toe of the hind-feet are longer than the
others. As with the preceding genera, the incisors are two in
each jaw. The species are M. albiventer, the Monk-Seal, and
M. tropicalis, the Jamaica Seal.

Allied to Cystophora is the genus Macrorhinus, with (possibly)
two species, of which one is Antarctic, the other frequents or
frequented the coast of California. The incisors are two in
the upper jaw, and but one in the lower. The premolars are
four and the molar one ; all the teeth are small and simple, but

454 THE ELEPHANT SEAL CHAP.

have long roots. The nose of the male has a dilatable proboscis.
The southern Elephant Seal is M. leoninus, and reaches a
length of some 20 feet. It occurs on the shores of Kerguelen
and some other more or less remote islands. Its habits have
been studied and described by several observers, beginning with
Anson in the last century. The late Professor Moseley gave a good
account of this marine monster in his book on the voyage of
the " Challenger." When the animal is enraged, the end of the
snout is dilated ; but when this happens there is no long and
hanging proboscis such as has sometimes been described. The
inflation affects the skin on the top of the snout, which thus rises
rather upwards during inflation. The inflated region, according
to Mr. Vallentin, quoted by Mr. J. T. Cunningham, is about
1 foot long in an individual of 17 feet. It has been stated that
this proboscis is a temporary structure, only appearing in the
breeding season ; but recent observations have shown that this
statement is inaccurate ; it persists all the year round. The
males fight greatly during the breeding season, and produce a
roar which has been compared to the " noise made by a man
when gargling." The females and the young males bellow like a
bull. The males fight of course with their teeth, literally falling
upon one another with their whole weight. Mr. Cunningham
thinks that the use of the proboscis is to protect the nose from
injury; or that it may be merely the result of "emotional
excitement." In any case the Bladder-nosed Seal, Cystophora, is
undoubtedly protected from injury by the possession of a corre-
sponding hood. The nose is the most vulnerable place, and the
existence of this hood would stave off the effects of a blow in
that region. Moseley, however, has said of Macrorhinus that it
cannot be stunned by blows on the nose as other Seals can ; but
he attributes this, not to the dilated snout, but to the bony
crest on the skull, and to the strength of the bones about the
nose. This Seal crawls with difficulty on the land, and as the
animals move " the vast body trembles like a great bag of jelly,
owing to the mass of blubber by which the whole animal is
invested, and which is as thick as it is in a whale."1 When
lying on the shore, these animals scrape sand and throw it
over themselves, apparently to prevent themselves from being

1 Cunningham, "Sexual Dimorphism in the Animal Kingdom," London, 1900 ;
see also Flower, Proc. Zool. Soc. 1881, p. 145.

xiv ANCIENT CARNIVORES CREODONTA 455

incommoded by the direct rays of the sun, to the effects of

which they are very susceptible. The Elephant Seal is mild

and inoffensive, unless enraged, and, of course, during the
breeding season.

Order VIII. CREODONTA.

This entirely extinct group of Mammalia may be thus character-
ised : — Small to large carnivorous mammals, with skull on the
whole like that of the Carnivora and with trenchant teeth ; digits
with unguiculate phalanges ; tail long ; extremities usually with
five, sometimes with four digits. In the carpus a centrale is
present, and the scaphoid and lunar are separate. Interlocking
of posterior dorsal and lumbar zygapophyses very perfect. Brain
small but convoluted.

This group, which corresponds with the CARNIVORA PRIMI-
GENIA of Mr. Lydekker, is not easy to separate absolutely from the
existing and more especially from some of the extinct members
of the CARNIVORA VERA. They also come exceedingly near the
Condylarthra, the presumed ancestors of the Ungulata, and like
them begin in the earliest Tertiary deposits. Their likeness to
the carnivorous Marsupials has also been insisted upon ; but it
would seem that the succession of teeth in the Creodonta is
typically Eutherian.

The characteristics of the group may be exemplified by an
account of the genus Hyaenodon, after which some of the more
important deviations in structure shown by other genera will be
referred to.

Hyaenodon is both American and European, and ranges
through the Eocene and the Upper Miocene. It is a much-
specialised Creodont, and therefore exhibits well the distinctive
characters of the group. About a dozen species .have been
described. One of the best-known is the American H. cruentus,
and the following description refers to it. The back part
of the skull is low and broad, and is compared by Professor
Scott (who has described this and other species) as being
"somewhat like that of an opossum."1 The whole skull is

1 Journ. Ac. Sri. Philadelphia, ix. 1886, p. 175.

456 HYAENODON AND PATRIOFELIS CHAP.

long, and the top has a great sagittal crest. The paroccipital
processes are short and are closely applied to the mastoid processes.
The mesethmoid is larger than in the carnivorous Marsupials, and
the frontals are very large. The palate has a peculiar structure ;
in most species the hinder ends of the palatines are separated
by a narrow fissure which broadens gradually, thus forming the
posterior nares. In H. leptocephalus the posterior nares are brought
very far back by the meeting of the alisphenoids. The pre-
sphenoid, contrary to what we find in the Dog, for example, is
chiefly concealed by the vomer, which covers it. The mandible
has a long and strong symphysis, and its angle is not inflected.
The fore-limb is described as being " weak when compared with the
modern Carnivora." The scaphoid and lunar are separate, and
there is a centrale. The teeth present us with nearly the typical
formula. There is only one molar missing in the upper jaw. The
canines are enlarged. It has been suggested from a consideration
of its palate that Hyaenodon was a semiaquatic animal ; the
deep cleaving at the extremities of the phalanges seems to point
in the same direction, since they resemble in this the genus
Patriofelis, which there are other reasons to regard as aquatic.
This latter genus has a fore-limb which is very like that of the
Pinnipedia, the digits are much spread out, and would seem
to have supported a kind of paddle. In any case it certainly
fed upon aquatic tortoises, for their remains have been found in
its coprolites. The name Limnofelis, also applied to what appear
to have been members of this genus, is suggestive of their habits.
Patriofelis, at least one species, seems to have been of about the
size of a Lion.

Mesonyx has a brain case which is actually smaller than that
of the Marsupial Thylacinus. The lachrymal bone is very large,
and extends a little way over the face, as is also the case with
Hyaenodon ; this condition is also found in Insectivora and in
Thylacinus. The axis vertebra has a curiously-shaped spine,
which is very different from the hatchet-shaped process of that
vertebra usual in the Carnivora, but is not unlike what exists
in the Arctoid genera Meles and Mydaus. The limbs show
much disparity in length, and seem to argue a much -arched
back when the creature progressed. The carpus is stated to
be strikingly like that of the Insectivora. There is as in
other Creodonts a separation between the scaphoid and lunar;

xiv CREODONTA MESON YX 457

the centrale appears to be present. The pelvis " is most
like that of the bear," the metacarpals and the tibia, and some
other bones, resemble those of the Hyaena. In fact this animal
shows those combined characters which are common in archaic
forms.

CHAPTER XV

RODENTIA TILLODONTIA

Order IX. RODENTIA1

SMALL to moderately large animals, furry, sometimes with spines.
Toes with nails of a claw-like character, or sometimes approach-
ing hoofs. Usually plantigrade, and only occasionally and
partly carnivorous. Canine teeth absent ; incisors long and
strong, growing from persistent pulps, and with enamel only or
chiefly on the anterior face, producing a chisel -shaped edge;
molars few (two to six), separated from the incisors by a wide
diastema. Caecum (nearly always present) very large, and often
complicated in structure. Brain, if not smooth, with few furrows,
the hemispheres not overlapping the cerebellum. Surface of skull
rather flat ; orbits not separated from temporal fossae ; malar
bone in middle of zygomatic arch ; palate very narrow, with
elongated incisive foramina ; articular surface for lower jaw
antero-posteriorly elongated. Clavicles generally present. Testes
generally abdominal. Placenta deciduate, and discoidal in form.
The Rodents are a very large assemblage of usually small,
sometimes quite minute, creatures, embracing an enormous
number of living generic types. They are distributed all over
the world, including the Australian region, and, being small and
often nocturnal, and by no means particular in their diet, have
managed to thrive and multiply to a greater extent than any
other group of living mammals. They are chiefly terrestrial
creatures, and often burrow or live in ready-made burrows.

1 See especially Tullberg, " Ueber das System der Nagethiere," Act. Ak. Ujysala,
1899 ; and Alston, Proc. Zool. Soc. 1875, p. 61 ; and for nomenclature, Thomas, Proc.
Zool. Soc. 1896, p. 1012 ; and Palmer, Proc. Biol. Soc. Washington, xi. 1897, p. 241.

CHAP. XV

CHARACTERS OF RODENTS

459

Some, however, such as the Voles, are aquatic ; others, e.g. the
Squirrels, are arboreal, and there are " flying " Eodents exemplified
by the genus Anomalurus. Their range of habitat is in fact as
wide as that of any other Order of mammals, and wider than
that of most.

The most distinct anatomical characteristic of the Eodents
concerns the teeth. They are without exception entirely deprived
of canines. Thus there is a long diastema between the incisors
and the molars. Another peculiarity is, that in many cases the
dentition is absolutely
monophyodoiit. In
such forms as the
Muridae there seems to
be no milk dentition
at all. In that family
there are only three
molars ; but in other
types where there are
four, five, or six molars,
the first one, two, or

three, as the case may pic. 231.— Side view of skull of Cape Jumping Hur

Na

Per

(Pedetes ca/er). x f. AS, Alisphenoid ; Ex.O,
exoccipital ; FT, frontal ; L, lachrymal ; Ma,
malar ; MX, maxilla ; Na, nasal ; OS, orbito-
sphenoid; Pa, parietal; Per points to the large
supratympanic or mastoid bulla ; PMx, pre-
maxilla ; Sq, squamosal ; Ty, tympanic. (From
Flower's Osteology.)

be, have milk pre-
decessors, and may thus
be termed premolars.
This has been definitely
proved to be the case
in the common Eabbit, which has the unusually large number of six
grinding teeth in each half of the upper jaw when adult. The
first three of these have milk forerunners. On the other hand the
existence of four molars does nou apparently always argue that
the first is a premolar ; for Sir W. Flower found that in Hydro-
ckoerus,1 none of the teeth had any forerunners, at any rate so
far as could be detected from the examination of a very young
animal. The Eabbit appears to be also exceptional, in that the
second incisor of the upper jaw and the incisor of the lower jaw
have milk forerunners. In any case the tendency towards mono-
phyodontism is peculiarly well-marked in this group of mammals.
The incisors of Eodents are as a rule in each jaw a single pair of
long and strong teeth, which grow from persistent pulps, and

1 Proc. Zool. Soc. 1884, p. 252.

460

INCISOR AND MOLAR TEETH

"row to a very great length, extending back within the jaw to
near the hinder part of the skull. These teeth are reinforced in
the upper jaw by a small second pair in the Lagomorpha only.
The incisors are chisel-shaped, and often brown or yellow upon
the outer face, as is the case also with some Insectivores. This
peculiar shape, and their strength, renders them especially capable
of the gnawing action which characterises the Kodents. It has
been pointed out that where the incisors are wider than thick,
the gnawing powers are feebly developed ; and that on the con-
trary, where these teeth are thicker than wide, the animals are
good gnawers. The incisors have often an anterior groove, or it
may be grooves.

Fin. 232. — Molar teeth of Rodents. A, of Capybara (Hydrochoerus) ; B, of Squirrel
(Sciurus) ; C, of Ctenodactylus. (After Tullberg. )

The cheek teeth vary in number from two (Hydromys) to six
(Rabbit) on each side of the two jaws. Four is the prevailing
number outside the large division of the Bat-like Kodents.
They are often set at an angle to the horizontal plane of the jaw,
looking outwards and obliquely to its longitudinal axis ; the in-
dividual teeth too are not unfrequeiitly bowed in form, remind-
ing us of those of Toxodon. This of course only occurs in those
genera which have hypselodont teeth. The pattern of the teeth
varies much, and the different forms recall the teeth of more
than one other group of mammals. They are either bunodont
or lophodont. In many cases the tooth is encircled with a ridge
of enamel, which is either almost simple or has a more com-
plicated contour ; such teeth are by no means unsuggestive of the
Toxodonts. Some of the lophodont molars are by no means
unlike those of the Proboscidea. In Sciurus vidgaris the

xv SKULL OF RODENTS 461

encircling ridge is broken up into tubercles, which gives to the
tooth a striking likeness to those of Ornithorliynchus. Other
genera have teeth like those of many Ungulates. It has been
shown by Sir J. Tomes1 that the minute structure of the enamel
differs in different groups of Eodents.

The skull shows certain primitive characters. In the first
place there is no distinction between the orbital and the temporal
fossa.2 The sutures between the bones retain their distinctness
for very long. Other characteristic features are the following : —
The nasals are large, and so are the paroccipital processes. The
palate in front of the molars is not distinct from the sides of the
skull, its edge gradually becoming rounded off above. It is also
very narrow. The premaxillae are large in relation to the great
incisors. There is often a very much enlarged infra-orbital foramen
through which passes a part of the masseter muscle. The jugal
bone lies in the middle of the zygomatic arch, which is complete
and enormously enlarged in the Spotted Cavy (Codogenys paca).
As in many Marsupials, the jugal bone sometimes extends back-
wards to the glenoid cavity, where the lower jaw articulates. It is
usually said with an absolute want of accuracy that the cerebral
hemispheres of the Eodents are smooth and without convolutions.
This error has been repeated again and again in text-books. As
a matter of fact the cerebral hemispheres of many forms are quite
well convoluted,3 the degree of furrowing corresponding, as in so
many groups of mammals, with the size of the animal. This at
any rate is generally true, though the large Beaver with its scant
convolutions is an exception. The smaller forms, such as Mus,
Sciurus, Dipus, and Cricetus are quite smooth -brained. The best
furrowed brain of any Eodent which has been examined is that of
the huge Hydrochoerus. The Sylvian fissure is very generally
not pronounced ; but is particularly well-marked in Lagostomus.
In all, or in most, Eodents the hemispheres are separated by an
interval from the cerebellum, the optic lobes being visible between
the two.

The mouth cavity of this group of mammals is divided into
two chambers by a hairy ingrowth behind the incisors; this
arrangement is useful for animals which use their strong incisors

1 Phil. Trans. 1850, pt. ii. p. 529. 2 Seen, however, in Chaetomys.

3 See Beddard, Proc. Zool. Soc. 1892, p. 596, and Gervais, Journ. Zool. i. 1872,
p. 450.

462 THE ANOMALURES

as gnawing and excavating tools as well as for the purposes of
alimentation ; for it allows of substances being gnawed away
without the products of the chisel-like action being taken into
the hinder cavity of the mouth. The Rodents have for the most
part a simple stomach of normal form ; but in a few this is
complicated by a marked constriction, which divides the cardiac
from the pyloric portions. The Hamster, for example, is thus
characterised. In all the members of the order, with the excep-
tion of the Dormice and some allied forms, the caecum is large
and often sacculated. In some forms (e.g. Arvicola, Myodes,
Cuniculus) the large intestine is coiled upon itself in a spiral way
— a state of affairs strongly suggestive of Ruminants.

The Rodents are primarily divisible into two great groups, the
Simplicidentata and the Duplicidentata, characterised mainly by
the upper incisor teeth. In the former there is but one pair of
these teeth ; in the latter a second smaller pair lie behind the
former.

' SUB-OEDEE 1. SIMPLICIDENTATA.
SECTION 1. SCIUEOMOEPHA.

The Anomaluri are separated by Thomas and others from this
section as an equal section, while by Tullberg they are grouped
with Pedetes.

Fam. 1. Anomaluridae. — The genus Anomalurus suggests at
first sight the Flying Squirrels of Asia, Pteromys. It is, how-
ever, an entirely African genus, and is to be distinguished from
the Asiatic Rodents by a series of scales at the root of the tail,
imbricated, keeled, and forming possibly a " climbing organ."
This character serves also to distinguish the present genus from
Sciwropterua. The cartilage, moreover, which supports the
patagium springs from the elbow. There are four molars in each
half of each jaw. The eyes and ears are large. There are five
fingers and toes, but the thumb is small, though provided with
a nail. The sternum has seven joints, and nine ribs reach it.
The clavicle is strong. Huet, who has recently monographed the
genus,1 allows six species. The species vary in size.

"Observations sur le genre Anomalurus," JVouv. Arch. Mus. (2), vi. 1883,
]). 277.

HABITS OF ANOMALURUS 463

Anomalurus peli appears, according to Mr. W. H. Adams,1 to
be a common species on the Gold Coast ; it is coloured black and
white, but in spite of the warning which this colour should
convey, is considered by the perhaps rather omnivorous native as
" the greatest delicacy." The animal is nocturnal, but affects only
bright moonlight nights. Their " flying " consists of a jump from
a high branch to a lower one, after which they reascend the tree
to a point of vantage for another jump. They are said to feed
upon nuts ; but Tullberg only found the remains of leaves in the
stomach.

Idiurus is a lately-described genus allied to Anomalurus.
There are at any rate two species, /. zenkeri and I. macrotis.
The thumb is more reduced than in Anomalurus, and the fibula,
contrary to what is found in that genus and in most Sciuromorphs,
is fused with the tibia below.

A third genus, very recently described and allied to both the
foregoing, is Aethurus. It is a native of the French Congo,2 and
differs by the absence of flying membranes. It has, however, the
pad of large scales. There is but one species, A. glirinus. It
has a black bushy tail. The postorbital processes of the frontals
are totally wanting — there are not even the traces to be seen in
Anomalurus. The thumb has vanished. If we are to compare
Anomalurus with the Squirrels then, thinks Mr. de Winton, the
present genus is probably diurnal by reason of the want of flying
membranes.

Fam. 2. Sciuridae. — The Squirrels, genus Sciums, are world-
wide in range, the Australian region and the island of Madagascar
being alone excepted.

The eyes and the ears are large ; the tail is of course long
and bushy. The fore-feet have an inconspicuous thumb ; the
hind-feet have four toes. The soles of the fore-feet are
naked or furry, those of the hind-feet are hairy. There are
twelve or thirteen dorsal vertebrae, and in correspondence seven
or six lumbars. The caudal vertebrae may be as many as twenty-
five. In the skull the frontals are broad, and there are long

1 "On the Habits of the Flying Squirrels of the genus Anomalurus," Proe. Zool.
Soc. 1894, p. 243.

2 W. E. de Winton, " On a New Genus and Species of Rodents," etc., Proc. Zool.
Soc. 1898, p. 450. Apparently just at the time of the publication of this paper
Matschie described the same animal as Zenkerella.

464 SQUIRRELS AND GROUND SQUIRRELS . CHAP.

postorbital processes. The infra-orbital foramen is, as a rule, not
large, but is increased in size in a few forms. The number of
separate pieces of bone in the sternum is five. The molars of the
upper jaw are five, but the first is very small and soon drops out.

The Squirrels are often rather brilliantly coloured. The
Chinese Sc. castaneiventris has grey fur with a rich chestnut-
coloured under surface. The Malabar Squirrel, Sc. maximus, as
its name implies, a large animal, has a deep reddish or chestnut-
coloured fur above, which becomes yellow below. The " Common
Squirrel," " the lytill squerell full of besynesse," which is the
Squirrel of this country, is brownish red on the upper parts and
white below. It ranges from this country as far east as Japan.
Like many other Eodents the Squirrel likes animal food and will
eat both eggs and young birds. " Camel's hair " brushes are
made from this animal. The genus Tamias, almost exclusively
North American in range, is included by Dr. Forsyth Major l
in this genus, which then consists of considerably over one
hundred species.

The Ethiopian Ground Squirrels, genus Xerus, have a more
elongated skull than Sciurus, and the postorbital processes are
shorter. The feet are not hairy.

Nannosciurus forms a perfectly distinct genus of Squirrels.
These " Pygmy Squirrels " differ in possessing a very elongated
" face " and in the very broad frontal region. The teeth are
unlike those of Sciurus in certain features, and have been
especially compared by Forsyth Major to those of the Dormice.
Four species of this genus are Malayan ; one is West African.

The Bornean Rheithrosciurus macro tis is the only species of
its genus. The genus may be distinguished by the exceedingly
brachyodont molars, this feature being more marked in this genus
than in all other Squirrels. It is called the " Groove-toothed
Squirrel," from the " seven to ten minute parallel vertical grooves
running down the front face of its incisors." 2

The genus Spermophilus includes a large number (forty or so)
of Palaearctic and Nearctic animals known as Sousliks. The ears
are small ; there are cheek pouches as in Tamias. The general
aspect of the animal is like that of a Marmot, and they bridge over
the exceedingly narrow gap which separates the Marmots from
the true Squirrels. Anatomically the skull is like that of

1 Proc. Zool. Soc. 1893, p. 179. 2 Flower and Lydekker.

xv PRAIRIE DOGS 465

Arctomys ; the molars are five in the upper and four in the lower
jaw. The caecum is relatively speaking very small ; the measure-
ments in a specimen of S. tredecimlineatus, dissected by Dr.
Tullberg, were: small intestine, 580 mm.; large intestine, 170
mm. ; and caecum, 27 mm. In Tamias also the caecum is not
greatly developed. These animals are burrowing in habit.

The Prairie-dogs, genus Cynomys, of which the best-known
species is perhaps C. ludovicianus, are very like the Squirrels, but
they are not arboreal creatures ; they live in burrows on the
ground, as their vernacular name denotes. The genus is entirely
North American, and four species have been differentiated.

The Prairie-dos or Prairie-marmot is some 10 inches to one

FIG. 233. — Long-tailed Marmot. Arctomys caudatus. x f .

foot in length. The tail is no more than 2 inches. The ears
are very small ; the thumb is fully developed and bears a claw.
The measurements of the various sections of the intestine are the
following: — Small intestine, 860 mm.; large intestine, 690 mm.;
caecum, 75 mm. Thus the caecum is not large comparatively
speaking. These animals dig burrows on grassy plains which
they share with the Ground Owl (Speotyto cunicularis} and with
Eattlesnakes, all three species appearing to live in perfect amity.
Probably the Owls use the conveniently-constructed burrows, and
the Eattlesnakes come there to look after the young of both.

Closely allied to the last are the Marmots, genus Arctomys.

They differ in the rudimentary character of the thumb and in the

longer tail. The' eyes and ears are small. The distribution of

the genus is Nearctic and Palaearctic. There are ten species of

VOL. x 2 H

466

MARMOTS AND FLYING SQUIRRELS

the genus. The Alpine Marmot, A. marmotta, is familiar to most
persons. The animal lives high up in the Alps, and when
danger threatens it gives vent to a shrill whistle. It hibernates
in the winter, and as many as ten to fifteen animals may be found
closely packed together in a single, carefully-lined burrow.

The only other European species is A. lobac, the Siberian
Marmot, which occurs in the extreme east of Europe, and is
also Asiatic. There are four North American species, including
the Quebec Marmot, A. monax.

The genus Pteromys (of which the proper name, antedating

FIG. 234. — Flying Squirrel. Pteromys alborufiis. xi.

Pteromys by five years, appears to be Petaurista) is confined to the
Oriental region, where there are a dozen species or so. The
limbs are united by a parachute extending to the toes, and sup-
ported anteriorly by a cartilage attached to the wrists. There
are also membranes anteriorly uniting the fore-limbs to the neck,
and posteriorly uniting the hind-limbs to the root of the tail and
a trifle beyond. The skull and the dental formula are as in
Sciurus, but the pattern of the molars, which is much complicated,
seems to argue a different mode of nutrition. There are twelve
pairs of ribs. The large intestine (in P. petaurista) is very nearly
as long as the small, and the caecum is also " colossal " ; the
measurements in an individual of the species named were : small
intestine, 670 mm.; caecum, 320 mm.; large intestine, 650 mm.

THE BEAVER 467

The caecum is disposed in a spiral. The teats are three pairs,
non-inguinal in position.

The size of these squirrels is 16 to 18 inches exclusive of
the tail, which is longer. These animals can make an exceedingly
long jump with the help of their flying membrane. Nearly eighty
yards is the longest distance given for these aerial excursions. It
is stated that they are able to steer themselves to some extent
while in the air. As appears to be the case with so many
Rodents, these animals feed largely upon beetles and other insects,
besides bark, nuts, etc.

The allied genus Sciuropterus has a much wider range. It
extends into the Palaearctic region and into North America,
besides being found in India. There is here no membrane reach-
ing to the tail. The palms and soles are furry. The caecum is
very much shorter, and so is the large intestine. The latter, in
8. volucella,' is not more than one-third of the length of the small
intestine. In other features there are no remarkable differences
in structure, save that the mammae, always three pairs, may be
inguinal.

Of the genus Eupetaurus1 but a single species is known, which
is limited to high altitudes at Gilgit and perhaps in Thibet. Its
principal difference from the other genera of Flying Squirrels is that
the molars are hypselodont instead of brachyodout. The inter-
femoral membrane is rudimentary or wanting. The one species
is E. cinereus. It is thought to live " on rocks, perhaps among
precipices." Dr. Tullberg attributes the hypselodont teeth to the
fact that the mosses upon which it is believed to feed may have
much sand and earth intermingled, which would naturally lead to
a more rapid wearing away of the teeth, and hence a need for a
good supply of dental tissue to meet this destruction.

Fam. 3. Castoridae. — This, the third family of the Sciuro-
morpha, contains but one genus, Castor, the Beaver, with at most
two species, one North American, the other European. This large
Rodent has small eyes and ears, as befits an aquatic animal, and the
tail is exceedingly broad and covered with scales ; the transverse
processes of the caudal vertebrae, in order better to support the
thick tissues lying outside them, are divided in the middle of
the series into two. The hind-feet are much larger than the fore-
feet, and are more webbed than in any other aquatic Rodent.

1 Thomas, J. Asiat. Soe. Bengal, Ivii. 1888, p. 256.

468 HISTORY OF THE BEAVER CHAP.

In the skull the iiifraorbital foramen is small as in Squirrels.
The postorbital process has practically vanished. The four
molars stand out laterally from the jaws. The incisors, as might
be surmised from the habits, are particularly strong. The stomach
has near the entrance of the oesophagus a glandular patch,
which seems to be like that of the Wombat (see p. 144).
In both sexes the cloaca is very distinct and comparatively
deep.

The two species of the genus are C. canadensis and C. fiber.
The latter is of course the European species, which is now found
in several of the large rivers of Europe, such as the Danube
and the Rhone. But it is everywhere getting scarce, and limited
to quite small and isolated colonies.

In this country it is absolutely extinct and has been since
before the historic period. There is apparently no documentary
evidence of its survival down to this period. But the numerous
names of places which are called from this animal illustrate its
former prevalence. Examples of such names are Beverley in
Yorkshire, and Barbourne or Beaverbourne in Worcestershire.
In Wales, however, Beavers seem to have persisted longer. But
they were rare in the Principality for a hundred years or so before
the Norman Conquest. The king Howel Dda, who died in
948 A.D., fixed the price of a Beaver skin at 120 pence, the
skins of Stag, Wolf, and Fox being worth only 8 pence apiece.
The Beaver was called by the Welsh "Llost-llyddan," which
means "broad -tail." Its existence in the country is handed
down in the name of Llyn-ar-afange, which means Beaver lake.
The last positive record of the Beaver in Wales seems to be the
statement of Giraldus Cambrensis that in 1188 the animal was
still to be found in the river Teivy in Cardiganshire. In
Scotland the Beaver is said to have continued down to a later
date. Ireland it never reached. The remains of this animal
by their abundance show the former prevalence of C. fiber
in this country. It is known from the fens of Cambridgeshire,
and from superficial deposits elsewhere. The Thames formerly
had its Beavers, and apparently it was widely spread through the
country generally.

The Beaver not only furnishes collars and cuffs for coats ;
it was used, as every one knows, to provide hats. But the useful-
ness of the animal by no means ended here in the eyes of our

xv CASTOREIN AND ITS USES 469

forefathers. The Rev. Edward Topsell observed that " for giving
great ease unto the gowt the skinnes of beavers burned with drie
oynions " are excellent. Castorein as a drug, if not in actual use,
has quite recently been a part of the pharmacopoeia. It is
derived from the anal glands common to this and other Eodents,
and indeed many other mammals.

A large extinct form of Beaver is Trogontherium,1 found in the
" Forest-bed " of Cromer. The skull is about one-fourth longer
than that of Castor. It has a less inflated bulla, and slightly more
pronounced postorbital processes than Castor. The third molar
(fourth grinding tooth) is relatively larger than in Castor, and has
a rather more folded crown. The foramen magnum is more
triangular.

Fam. 4. Haplodontidae. — A separate family seems to be re-
quired for the genus Haplodon, whose characters will therefore be
merged with those of its family. It is to be distinguished from
most other Squirrel-like creatures by the fact that there is no post-
orbital process to the frontal. The molar teeth are five in the
upper and four in the lower jaw. The Sewellel, H. rv/us, like
the other species of the genus (ff. major}, is found in North
America west of the Eocky Mountains. It has the habit of the
Prairie-marmot, and has a short tail, only moderately long ears,
and five-toed feet. Tullberg is of opinion that this animal nearly
represents the ancestral form of the Squirrel tribe.

SECTION 2. MYOMOEPHA.

This subdivision of the Eodents contains, according to Mr.
Thomas's recent estimate,2 no less than 102 genera. It is there-
fore obviously impossible to do more than refer to some of the
more interesting of these. This group is again divided into the
following families : —

(1) Gliridae, including the Dormice.

(2) Muridae, the Eats, Mice, Gerbilles, Australian Water-rats,
Hamster.

(3) Bathyergidae, Cape Mole, etc.

(4) Spalacidae, Bamboo Eats.

1 E. T. Newton, Trans. Zool. Soc. xiii. 1892, p. 165.
2 Proc. Zool. Soc. 1896, p. 1016.

4/0 SLEEP OF THE DORMOUSE CHAP.

(5) Geomyidae, Pouched Eats.

(6) Heteromyidae, Kangaroo Eats.

(7) Dipodidae, Jerboas.

(8) Pedetidae.

The G-liridae have no caecum, so usual in the Eodentia. It
is true that all the genera have not been dissected, but it is
known that in the true Dormice, as well as in the genus
Platacanthomys, a caecum is absent.

Apart from these few exceptions the Mouse-like Eodents all
possess a caecum, though it is often not very large. They are
all smallish animals, and are modified to a great variety of habit
and habitat. There are burrowing, swimming, and climbing
forms. The group is universal in range, even including the
Australian region, in which they are the only Eodents.

Fam. 1. Gliridae. — This family, also called Myoxidae,1 includes
the Dormice, and is entirely an Old- World family, absent only
from the Malagasy region. Its most important differential
character is the total absence of the caecum and of any sharp
boundary between the small and large intestine. The molars are
usually four. The eyes and ears are well developed.

The genus Muscardinus includes only the Common Dormouse,
M. avellanarius. This small creature, 3 inches long with a tail
of 2^ inches, is, of course, a well-known inhabitant of this
country. It is also found all over Europe. It is not particularly
abundant in this country, and a good specimen is said to be
worth half a guinea. As the specific name denotes, it lives
largely on hazel nuts ; but it will also suck eggs and devour
insects. The animal makes a " nest " in the form of a hollow
ball. Its hibernation is well known, and has also given rise to
the German name (" Schlafer ") of the group. It was well known
to Aristotle, who gave or adopted the name 'EXeto? for the
animal. Its winter sleep — suggestive of death — and its revivifica-
tion in the spring gave the Bishop of Salamis, Epiphanius, an
argument for the resurrection of man. The fur was reckoned
in Pliny's time a remedy for paralysis and also for disease of
the ears.

The genus Myoxus includes also but a single species, M. glis,
the so-called "Fat Dormouse" of the Continent. It has no

1 Reuvens, "Die Myoxidae oder Schlafer," Leyden, 1890, allows but one genus,
Myoxus, the other genera adopted here being termed subgenera.

xv THE RATS AND MICE 471

glandular swelling at the base of the oesophagus, such as occurs
in the last genus and in Graphivrus. Of Graphiurus there are
thirteen species, all African in range. The genus does not differ
widely from the last. There is, however, a glandular region of
the oesophagus. Eliomys is the last genus of typical Dormice.
It is Palaearctic in range.

Platacantkomys, of a Dormouse -like form, has like other
Dormice a long tail, on which the long coarse hairs are arranged
in two rows on opposite sides towards the tip ; it is represented
by a single species, P. lasiurus, from the Malabar coast. It is
arboreal in habit. The fur is mingled with flattened spines.
The molars are reduced to three on each side of each jaw. This
form has been bandied about between the " Mice " and the
" Dormice " ; but Mr. Thomas's discovery of the absence of the
caecum argues strongly in favour of its correct location among
the Gliridae. Typlilomys is an allied genus, also from the
Oriental region. This and the last are placed in a special
sub-family of the Gliridae, Platacanthomyinae, by Mr. Thomas.

Fam. 2. Muridae. — This family, that of the Eats and Mice
in a wide sense, is the most extensive family of Eodents. In it
Mr. Thomas includes no less than seventy -six genera. The
molars are generally three. The tail is fairly long, or very long,
and the soles of the feet are naked.

Sub-Fam. 1. Murinae. — The true Eats and Mice may be
considered to form a sub -family, Murinae. The genus Mus,
including the Eats and Mice in the limited sense of the
word, contains about 130 species. They are exclusively Old
World in range, being only absent from the Island of
Madagascar. In the New World there are no species of
the restricted genus Mus. The eyes and the ears are large ;
the pollex is rudimentary, and bears a nail instead of a claw.
The tail is largely scaly. All the members of the genus are
small animals, some quite minute. In this country there are
five species1 of the genus, viz. the Harvest Mouse, M. minutus,
which has a body only 2^ inches long with an equally long tail.
It is the smallest of British quadrupeds with the exception of
the Lesser Shrew. The Wood Mouse, M. sylvaticus, is about
twice the size ; it differs also from the last species in that it

1 To which a sixth, the "Yellow-necked Mouse," Mus flavicollis, may perhaps
be added.

472 THE BROWN RAT AND THE BLACK RAT CHAP.

frequents barns, and is thus sometimes mistaken for the Common
Mouse, from which, however, it is to be distinguished by its
coloration and longer ears. The latter, M. musculus, is too
familiar to need much description. A curious variety of it has
occurred. This has a thickened and a folded skin like that of
a, Ehinoceros, and the hair has disappeared. The Black Rat,
M. rattus, is like a large Mouse, and is smaller and blacker in
colour than the " Hanoverian Rat." It is sometimes called the
" Old English Rat," but seems nevertheless to be not a truly
indigenous Rodent. It has been so defeated by competition with
the Hanoverian Rat that it is now not a common species in this
country.

The Hanoverian or Brown Rat, M. decumanus, is a larger and
a browner animal than the last. It is very widely distributed
through the globe, no doubt largely on account of the fact
that it is readily transported by man. The same is the case
with the Common Mouse, whose real origin must be a matter of
doubt. The original home of the Brown Rat is thought by
Dr. Blanford to be Mongolia. There is so far a justification
for the name " Hanoverian Rat " that the animal seems to
have reached this country about the year 1728. But there
is no reason for calling it, as is sometimes done, the Norway
Rat.

Some members of this genus, whose fur is interspersed with
spines, or which are quite spiny, have been separated as a genus,
Leggada, which, however, is not generally allowed.

Closely allied again is Chiruromys, which has a strongly
prehensile tail, a feature which is not common among the
Myomorpha, though Dendromys, a tree -frequenting form, and
Mus minutus, already spoken of, show the same character.
Many Mice seem to have prehensile tails, which they can curl
round branches ; but it is not so fully developed as in the species
just named.

A number of other genera are referable to the true Mice, the
sub-family Murinae of Thomas's classification. The Syrian and
African Acomys has very spiny fur, so much so that "when
it has its spines erected it is almost indistinguishable at
the first glance from a diminutive hedgehog." The genera
Cricetomys, Malacomys, Lophuromys, Saccostomus, Dasymys are
restricted to the Ethiopian region. Nesokia is Oriental, reaching

BARBARY MICE 473

also the Palaearctic region. Vandeleuria, Chiropodomys, Batomys,
Carpomys are Oriental, the last two being confined to the
Philippines.

Another peculiar Philippine genus is Phlaeomys, of large size,
and allied to it is Crateromys, originally confounded with it.
Batomys granti is also confined to Luzon. Its molars are three,
like those of the also restricted and Philippine Carpomys melan-
urus, which is an arboreal form. There is a second species,
C. phaeurus.

Phlaeomys is placed, however, by Mr. Thomas in a distinct
sub-family of its own, Phlaeomyinae, and is removed from the
Murinae.

Rapalomys, with but one species, is Burmese. Pithecochirus is

FIG. 235. — Spiny Mouse. Acomys cahirinus. x

Javanese and Sumatran. Conilurus (also known as Hapalotis) is
a genus containing species which are termed Jerboa Rats, on
account of their mode of progression. They are desert and
Australian forms. There are sixteen species.

Mastacomys, with one species, is limited to Tasmania. Uromys,
with some eight species, is from Queensland, and inhabits also the
Am Islands and the Solomon Islands. The Celebesian Echiotlirix,
or Craurothrix as it should apparently properly be called, is another
genus containing but a single species. Golunda is both Oriental
and Ethiopian, one species occurring in each region. The beauti-
ful little striped Barbary Mice, ArviacantUs (or Isomys}, are
African, north as well as tropical.

The genus Saccostomus resembles the Hamsters in the presence
of cheek pouches. Its teeth, however, are Murine. It agrees with
Steatomys in the comparatively short tail. The caecum is rather
long.

474 PHILIPPINE RODENTS CHAP.

Sub-Fam. 2. Hydromyinae. — The genus Hydromys,1 of which
there are several species, the best known being If. chrysogaster,
is an exclusively Australian form, and is aquatic in habit. It
is a foot or so in length, and has a fairly long tail. The fore-
and hind-limbs are webbed, in correspondence with its habits.
The Australian Water- Eat is black, with an admixture of golden-
coloured hairs dorsally and golden colour below, with a lighter
median stripe. The thumb is small, and the webbing of the
hands is not so marked as is that of the feet. The molars are
only two in each half of each jaw. The caecum is rather small,
the measurements of the alimentary canal being : small intestine,
895 mm. ; large intestine, 278 mm. ; caecum, 70 mm. Allied to
the last is Xeromys, a genus which is also Australian, but limited
to Queensland. It has been established by Mr. Thomas,2 who
discovered that it has the same reduced formula as Hydromys.
Xeromys, however, is not an aquatic animal, and has unwebbed
feet.

In the Luzon highlands Mr. Whitehead has discovered, and
Mr. Thomas quite recently described,3 a number of peculiar
Eodents. Of these the genera Chrotomys, Celaenomys, and Cru-
nomys are allied to the Australian and New Guinea Hydromys.

Chrotomys whiteheadi is unusual among Muridae, in its
coloration being marked by a pale stripe down the back. The
creature is the size of the Black Eat (Mus rattus}. It is terres-
trial not aquatic in habit, in spite of its likeness to Hydromys.
The molars, however, are |.

Crunomys fcdlax is more like Hydromys. It has, however,
three molars, as in the last genus. But the skull has the
flattened form characteristic of Hydromys as opposed to Mus.

Like Batomys, Celaenomys silaceus is also somewhat inter-
mediate between Hydromys and Mus. It is described as very
Shrew-like in appearance, and has a very pointed muzzle. Its
habits Mr. Whitehead is " quite unable even to guess at." Like
Hydromys and Xeromys this Eodent has but two molars.

Sub-Fam. 3. Rhynchomyinae. — The genus EliyncJiomys, con-
taining but one species, Eh. soricioides (of Thomas), is also, as both
its generic and specific names imply, a somewhat Shrew-like form in
external aspect. The skull, too, is Insectivore-like in its elonga-

1 For anatomy see Windle, Proc. Zool. Soc. 1887, p. 53.
2 Proc. Zool. Soc. 1889, p. 247. 3 Trans. Zool. Soc. xiv. 1898, p. 377.

THE GERBILLES

475

tion, and the lower incisors are worn to needle-like points. The
two molars are excessively minute, and thus the always large gap
in the jaws is greatly exaggerated. It is suggested that this Eat
is an insect-eater, but nothing positive is known.

Sub-Fam. 4. Gerbillinae. — The Gerbilles form another sub-
family, Gerbillinae, of the Muridae, or a family, according to some.
The best-known genus is Gerbillus, including the Gerbilles proper.
These animals are Old World in range, belonging to the three
regions of that part of the world. There are a large number of
species in the genus, over thirty. They have a Jerboa-like form,
with rather long hind-limbs and a long and hairy tail. But the
hind- as well as the fore-feet are five-toed. The molar teeth have
no trace of tubercles, but only transverse lamellae of enamel.

FIG. 236. — Gerbille. Gerbillus ae.gyptius. x \.

The incisors are orange ; they are white in Dipus. Gerbillus
pyramidum is 90 mm. long, with a tail of 125 mm. The ears
are long, 13 mm. The tail has longer hairs at the tip.

Psammomys is in some respects different. The tail is shorter
than in Gerbillus; its length in an individual of 165 mm. was
130 mm. As in Gerbillus there are four pairs of teats, two
pectoral and two inguinal. This genus is exclusively Palaearctic
in range. Meriones has a range co- extensive with that of
Gerbillus.

Pachyuromys is an Ethiopian genus with a short tail. As
the generic name denotes, the tail is not only short but thick
and fleshy.

Sub-Fam. 5. Otomyinae. — The allied genera, Otomys and
Oreinomys, are Ethiopian. Otomys unisulcatus has a tail shorter
than the body, the measurements of a female of this species being
137 mm. with a tail of 87 mm. The ear is long, whence the
name; it measured in this specimen 20 mm.

476 ANATOMY OF THE CRESTED RAT CHAP.

Sub-Fam. 6. Dendromyinae. — The genus Deomys is an Afri-
can form, consisting of only one species from the Congo region.
D. ferrugineus has a reddish colour as its name implies ; the soles
are quite naked and the tail is long and slender. It is con-
siderably longer than the body, measuring (minus a fragment of
the tip) 172 mm., while the body is 125 mm. long. The
characters of the molar teeth, which are three, are intermediate
in their form between those of the true Rats and those of the
Hamsters.

Dendromys is also Ethiopian in range. There are several
species. D. mesoinelas is a smallish creature, 60 mm. long, with
a tail of 90 mm.

Steatomys is another African genus, allied to the last. Its
tail, however, is only half the length of the body. The two
remaining genera are Malacothrix and Limacomys. Their range is
African.

Sub-Fam. 7. Lophiomyinae. — Allied to the Hamsters is the
singular East African genus Lophiomys, with only one species,
L. imliausi, of Milne-Edwards.1 The size is between that of a
Rabbit and of a Guinea-pig. The stomach is curved and somewhat
intestiniform. It has been termed the Crested Rat on account of
the " prominent crest of stiff hair running down the back." The
fingers and toes are five, and the very long tail is clad with hair
longer than that upon the body generally. The pollex is rudi-
mentary, and the hallux is opposable.

The most remarkable structural feature in this genus con-
cerns the skull, and on account of this it has been regarded as
the type of a separate family. The temporal fossa behind the
eye is covered over by a complete bony plate, formed by a down-
growth of the parietal, meeting an upgrowth from the malar ;
this singular arrangement of the bones recalls the conditions
which obtain in turtles. The whole skull, moreover, is covered
with symmetrically disposed granulations, such as are found in
no other mammal ; it suggests rather the skull of certain fish.
It is believed that the bony plate already referred to is not really
a portion of the bones of which it appears to be a prolongation,
but merely an ossification of fasciae in this region. The atlas
is granulated like the skull ; there are sixteen pairs of ribs and
a feeble clavicle. The molars are three, and of a peculiar form.
Nouv. Arch. Mus. iii. 1867, p. 81.

xv A PLAGUE OF VOLES 477

They have, in the case of the first three, transverse ridges, from
which stand up two sharp and long tubercles. The other teeth
have two ridges. The incisors are pale yellow. The shape of
the teeth and the smallness of the caecum suggest that this
Eodent is not so purely a vegetarian as others, and that it
nourishes itself largely upon insects.

Sub-Fam. 8. Microtinae. — The Voles or Water-Eats form a
distinct group of Murine animals, to which the sub -family
name of Microtinae has been applied from the genus Microtus
(more generally known as Arvicola), a genus which includes
the Water-Eat and Field- Voles of this country. This genus
has short ears, and a short and hairy tail. Its build is
stouter and clumsier than that of the Eats. The genus is
confined to the Palaearctic and the Nearctic regions. In this
country there are three species. The best known is the Water-
Vole or Water-Eat, M. amphibius, which has been seen by
most people, and which frequents streams, ponds, and canals.
The feet, curiously enough, are not webbed, which seems to
argue the recent adoption of an aquatic life. Mr. Trevor-
Battye has remarked that this animal, when swimming at
leisure, uses its hind-limbs only, carrying the fore pair at
the sides like a Seal. The Bank-Vole, M. glareolus, is rather a
local species in this country. It is a terrestrial Vole, and
burrows. The Field- Vole, M. agrestis, has become notorious on
account of the "plagues," to which its immense numbers have
on occasions given rise. It is the smallest species, and has a
greyish-brown fur like the Water- Vole, the Bank- Vole being
redder. To give an idea of the cost of the depredations of this
animal, Mr. Scherren quotes * a farmer who gave evidence before
the Agricultural Commission to the effect that, putting the
damage of one Vole at two pence, the amount of loss suffered on
a farm of 6500 acres in two years would be £50,000 !

The genus Fiber comes very near the last. It is a North
American genus. The hind-feet are slightly webbed ; the tail is
a trifle shorter than the body, and is compressed and scaly, with
scattered hairs. The thumb is short, but with a fully-developed
claw. As in the last genus, the small and large intestines are
roughly of the same length, and the caecum is about one-fourth
of the length of either. It is known as the " Musquash."
1 Popular Natural History of Animals, London, 1898.

478 THE SCANDINAVIAN LEMMING CHAP.

Of Fiber zibethicus, or rather a closely-allied form, F. osoyoos-
oisis, from Lake Osoyoos near the Rockies, Mr. Lord writes l that
it constructs for itself a house of bulrushes built up from the
bottom in 3 or 4 feet of water. It is dome-shaped, and rises
about a foot out of the water. " If a dead or badly-wounded
duck be left on the pool, it is at once seized on, towed into the
house, and doomed." Thus it appears that this Rodent, like so
many others, is largely carnivorous. It has also been asserted
that it eats fish.

Neofiber is an allied genus, North American in range. The
species, N. alleni, is compared, as regards outward form, with the
Water- Vole, M. amphibius. It has, however, a shorter tail.

Another very well-known member of this sub-family is the
Lemming. The name, however, applies to two quite distinct
genera. The genus Cuniculus, including the Banded Lemming,
C. torquatus, is an inhabitant of North America, Siberia, and
Greenland. The tail is short, its length being 12 mm. as against
a body length of 101 mm. The feet are furred beneath, a not
unusual state of affairs in Arctic mammals. The ears are very
slight. The thumb is well developed, and bears a claw.

In Myodes, on the other hand, which is not so markedly
an Arctic animal, though occurring in both Palaearctic and
Nearctic regions, the ears are rather bigger, though still smaller
than those of Microtus. The under surfaces of the feet are
similarly furred. The tail is also short. It is commonly said
that the two genera are to be distinguished by the furred feet
of Cuniculus, and by the absence of fur in the present genus.
That, according to Tullberg, does not appear to be the case. The
differences are thus so much reduced that it seems almost un-
necessary to retain the two genera. The best known species of
Ihjodes is of course the Scandinavian Lemming, M. lemmus.
This animal used to occur in this country in Pleistocene times (as
did also C. torquatus), and recently Dr. Gadow has found remains
with skins attached in caves in Portugal It may still survive
in the mountains of the Peninsula.

The actual habitat of the Lemming in Scandinavia is the

great tablelands, 3000 feet high in the centre. The migrations

do not take place with regularity ; even twenty years may elapse

before the appearance in cultivated lands of those countless

1 Proc. Zool. Soc. 1863, p. 95.

xv HAMSTERS AND COTTON RATS ' 479

hordes so familiar (as far as their description is concerned) to
everybody. The Lemmings do not return from their exodus.
They die from various causes, including combats with one
another. Their chief foes, however, are Wolves and Gluttons,
Buzzards and Eavens, Owls and Skuas, which batten on the
migrant hordes. Their sudden increase in numbers recalls the
similar increase at times of the Field- Vole, to which reference
has already been made.

Ellobiiis is an Old-World genus, which leads a " Talpine " life,
and has in consequence rudimentary external ears and very
small eyes. The tail is short. Contrary to what might be
expected from its mode of life, the claws upon the digits are
not strong.

The remaining genera of Vole-like Murines are Phenacomys
and Synaptomys from North America, and Siphneus from Palae-
arctic Asia. Evotomys is one of those genera which are common
to both the Palaearctic and the Nearctic regions, but the bulk of
the species are North American.

Sub-Fam. 9. Sigmodontinae. — This is the name given to
another sub-family of Murine Eodents, a group which includes
the Hamsters in the Old World as well as a large number of
South American genera of Eat-like animals. Of these latter
there are a very large number, the bulk of the group being
American.

The Hamsters, genus Cricetus, as it is usually called, although
apparently the correct name is Hamster, are Old-World forms of
Pouched Eats. The Common Hamster, C. frumentarius, is about
210 mm. long, with a tail of 58 mm. It has cheek pouches.
The small and the large intestines are not very unequal in
length, and the caecum is fairly large, being about one-sixth to
one-seventh of the length of either. It is a purely vegetable-
feeding creature, and in Germany where it occurs (and from
which language its vernacular name is derived), hibernates during
the winter in its burrow, having previously surrounded itself
with a great accumulation of food carried thither.

To North America are peculiar the genera Onychomys, Sig-
modon, and Peromyscus. The genus Sigmodon, the Cotton Eats,
reaches Central America, and even gets a little farther south.
The other two genera, though mainly North American, also
extend their range to the south. Onychomys has hairy foot-

480 A FISH-EATING RAT

pads, a state of affairs which characterises a number of these
Rodents.

The genera Megalomys, Chilomys, Reithrodontomys, Eligmo-
dontia, Nectomys, Rhipidomys, Tylomys, Holochilus, Reithrodon,
Phyllotis, Scapteromys, Acodon, Oxymycterus, Ichthyomys, Blari-
nomys, Notiomys are South American forms. Oryzomys and
Rheithrodontomys are common to both parts of the New World.

The genus Ichthyomys is remarkable on account of its un-
Rodent-like habits and of certain associated structural changes.
/. stolzmanni was obtained from Mount Chanchamays in Peru
at an altitude of 3000 feet ; it is a habitual fish-eater, and
lives in streams. Another species, /. hydrobates, was formerly
referred to Habrothrix. The skull shows likenesses to that of
the Australian Hydromys ; but the most marked characters of
adaptation are those of the teeth and caecum. The cutting
edges of the upper incisors form a reversed V of obvious use
in holding a slippery fish. The caecum is much reduced, short,
and narrow. The general Otter-like shape of the creature is
largely due to its flattened head, though its " size and general
proportions are much as in the common Black Eat." *

This sub-family contains a number of genera from Madagascar,
viz. Br achy tar somys, Nesomys, Hallomys, Brachyuromys, Hypo-
geomys, Gymnuromys, and Eliurus.

Sub-Fam. 10. Neotominae. — The last sub -family of the
Muridae is that of the Neotominae, containing the North Ameri-
can genera Neotoma, Xenomys, Hodomys, and Nelsonia.

Fam. 3. Bathyergldae. — This family contains several genera
which consist of subterranean forms. All these Rodents agree
in a number of characters, of which the principal are as
follows : — r

The eyes are very small, and the external ears are reduced to
the merest fringe of skin round the aural aperture. The legs are
short, as is the tail ; the hair-covering is reduced — a reduction
which finds its culmination in the nearly nude Heterocephalus^
Being burrowing creatures, a number of other modifications in
accordance with this mode of life are to be seen in their
structure. The upper incisors stand out in front of the closed
lips, and prevent the entrance of earth. For the same reason

1 See 0. Thomas, "On some Mammals from Central Peru," Proc. Zool. Soc.
1893, p. 333.

xv THE CAPE MOLE-RAT 481

the nostrils are small, and the forehead but little expanded
between them.

The genus Bathyergus contains but a single species, the Cape
Mole-Eat, which is found in Southern Africa ; it is of moderate
size, not exceeding a small Eabbit in dimensions. On the
fore-limbs are exceedingly long claws, of which that borne by
the second finger is the longest, and the claw of the thumb the
shortest. The hind-feet have by no means such long claws.
The scratching and burrowing is naturally chiefly effected by the
fore-limbs. The small and large intestines are of equal length,
and each is rather more than six times the length of the caecum ;
in these measurements the present genus differs from the next.

Georhyclms. — Of this African genus there are about ten
species. The claws are not so long as in the last genus, but
there are, as in Bathyergus, four molar teeth on each half of
each jaw. The intestinal measurements in an example ' of G.
capensis were : small intestine, 25 inches ; caecum, 4 inches ;
large intestine, 15 inches.

The genus Myoscalops or Ifeliphobius (also with an African
range) has six back teeth on each side. A number of species
sometimes referred to the last genus are placed here by Mr.
Thomas. The claws are small.

One of the most remarkable genera of this family is the little
Heterocephalus from Abyssinia and Somaliland. As Mr. Thomas
justly remarks,1 it " is a peculiar-looking little creature, about the
size of the Common Mouse, but looking almost more like a tiny
hairless puppy 011 account of its nearly naked skin, small eyes,
and peculiar physiognomy." Though apparently naked, there are
numerous scattered hairs over the entire body, and the toes are
fringed with stiffish hairs, which must be advantageous to a
burrowing animal. There are two species, H. glaber (originally
described by Eiippell), and H. phillipsii, of which our knowledge
is due to Mr. Thomas. The length of the entire creature in-
cluding the tail is not more than 134 mm., both species being
approximately of the same dimensions. Mr. Lort Phillips, the
discoverer of the species which bears his name, writes " that this
little creature, called ' Farumfer ' by the Somali, throws up in
places groups of miniature craters, which exactly resemble
volcanoes in active eruption. When the little beasts were at

1 " Notes on the Rodent genus Heterocephalus," Proc. Zool. Soc. 1885, p. 84§.
VOL. X 2 I

482 BAMBOO RATS CHAP.

work, I used frequently to watch them, and found that the loose
earth from their excavations was brought to the bottom of the
crater, and sent with great force into the air in a succession of
rapid jerks, and that they themselves never ventured forth from
the shelter of the burrows." 1

Fam. 4. Spalacidae. — " The Spalacidae," observes Dr. Blan-
ford, " are sometimes called rodent moles, and resemble a mole in
general aspect, having cylindrical bodies, short limbs, small eyes
and ears, large claws, and a short or rudimentary tail." The
existence of a spiral valve in the caecum may perhaps characterise
this family ; but it has at present only been found in the two
genera Spalax and Rhizomys.

Spalax has inconspicuous eyes and external ears. The tail is
totally absent. The lower incisors are more developed than in
other Rodents ; they project in a bony sheath beyond the posterior
end of the ramus of the lower jaw. The scapula is long and
narrow. The large intestine is half the length of the small in-
testine. The animal seems to have only two pairs of teats, one
pectoral the other inguinal.

Spalax typhlus of Egypt, which is probably not different from
the European form, makes extensive burrows, some of the branches
being even 30 to 40 yards in length. In a "domical chamber,"
situated along the course of one of these burrows, Dr. Anderson
found no less than 68 bulbs stored up. Its eyes are mere black
specks among the muscles, but they appear, however, to have a
proper organisation. There are altogether eight species of the
genus, which is entirely Palaearctic in its range.

The genus Rhizomys, including a number of species known
as Bamboo Eats, is purely Oriental in range. Eh. sumatrensis
reaches a length of 19 inches; the better-known species, Rh.
ladius, is at most only 9 inches in length — in both cases the
measurements are exclusive of the tail, which is a quarter to
one-third of the length of the body, and is not scaly but nearly
naked, with a few scattered hairs. The molars are three, and
the incisors usually orange in colour ; but sometimes the upper
incisors are white as in Rh. ladius. There are thirteen dorsal
vertebrae. In Rh. pruinosus the large intestine is considerably
longer than the small intestine ; the lengths of the two sections
of the gut are 42 and 30 inches respectively. In another

1 Proc. Zool. Soc. 1885, p. 611.

xv HABITS OF RHIZOMYS 483

species the large intestine is slightly shorter than the small
intestine. In Rh. Radius the two parts of the gut are almost
exactly equal in length. There are three pairs of inguinal and
two pairs of pectoral teats. The name Rhizomys appears to have
been given to the animals of this genus for the reason that they
feed largely on roots. They burrow, and, like many other
burrowing animals, feed in the evening. As is the case with
other forms, Rhizomys is said to burrow with the assistance of
its teeth as well as of its claws.

FIG. 237. — Bamboo Rat. Rhizomys badius.

Tachyoryctes is an African genus closely allied to the last. There
are three Ethiopian species. It is mainly to be distinguished by
the different pattern upon the grinding surface of the molars.

Fam. 5. Geomyidae. — This family of burrowing Eodents is
limited to North and Central America. The animals have cheek
pouches, and small eyes and ears, in accordance with their mode
of life. The claws of the fere-limbs are very strongly developed.

The genus Geomys contains some eight species, which are
Central and North American, not extending, however, far north.
The incisors of the upper jaw are grooved with two grooves.
There are three pairs of teats — one axillary, and the two remain-
ing inguinal.

Thomomys, without grooves on the incisors, reaches to Canada
in the north, and does not extend as far south as the last genus.

484 THE JERBOA CHAP.

Allied to this family, and indeed united with it by Tullberg,
but kept separate by Thomas, is the

Fam. 6. Heteromyidae. — The members of this family are also
American, but are not confined to the northern-central regions
of that continent, for the genus Heteromys extends into South
America.

The genus Dipodomys, with twelve species, is of a Jerboa-like
form, as the following measurements of an example of D. merriami
will show. The length of the head and body was 85 mm.; of
the tail 127 mm.; the hind-foot is 32 mm. It has but four
toes. The hind-limb is longer than the front-limbs.

In Perodipus the same form is exhibited. There are, however,
five toes, and the sole of the foot is hairy. The axis vertebra
and the two following vertebrae are fused together.

Perognathus is a third genus. It has the same general slender
form, but the tail is not so long, being but little longer than the
body. The hind-limbs, too, are shorter. The teats of this and
of Perodipus are as in Geomys. The two remaining genera of the
family are Heteromys and Microdipodops.

Fam. 7. Dipodidae. — This family consists of small, plain-
living, and leaping or arboreal creatures, commonly known as

FIG. 238. — Jerboa. Dipus hirtipes. x J. Eastern Europe.

Jerboas. The main anatomical characters of the family are the
following: — There is a large infra-orbital foramen. The molars
are always reduced, the premolar being either absent in the
lower jaw alone or in both jaws. This family presents an
obvious likeness to Dipodomys (hence the name of the latter) and
to some other members of the American family Heteromyidae

STRUCTURE OF JERBOA

485

There is even the same ankylosis of the neck vertebrae. We
find, moreover, the same association of long-legged and shorter-
legged forms that characterises the Heteromyidae.

The typical genus Dipus is a smallish quadruped with long
naked ears and a long tail. The ten
species are all Palaearctic in range. The
fore-limbs are short and five fingered, and
the short pollex has no claw ; the hind-
limbs are excessively long and only three-
toed. . The bony structure of these limbs
is remarkable. The three metatarsals are
elongated almost like those of a bird, and
are ankylosed together. The digits have
long phalanges which alone reach the
ground as the animal hops. It is a curious
fact, and one not so easily identifiable with
the way of life, that the neck vertebrae of
this genus are ankylosed together with the
exception of the atlas, which is free ; the
arrangement is precisely like that of the
Sperm Whale. The last vertebra is, how-
ever, sometimes free.

only leap but they burrow, and their strong
incisors are said to be used in burrowing
through stony ground. They are eaten by
the Arabs, and are, or have been, called
Daman Israel, i.e. Lamb of Israel. In
D. liirtipes the body and tail measure respectively 4^- and 7
inches. The hind-feet have a tuft of long hairs below. Mr.
W. L. Sclater's newly-founded genus JZuchoreutes1 is somewhat
more primitive in its characters than is Dipus. The general
form is the same, with long ears and a long tail. But there
are five toes to the hind -limb, the two lateral ones though
nailed being much shorter than the middle three. It has a
" long pig-like snout," and the tail is cylindrical as in most other
Jerboas, with a tuft of longer hairs at the end. The incisor
teeth, grooved in Dipus, are here smooth, as in Alactaga. The
species was probably obtained " in the sandy plains round the
city of Yarkand."

1 Proc. Zool. Soc. 1890, p. 610.

Thp Tprboas not FIG. 239.— Bones of right pes
of Jerboa, Dipus aegyptius.
x f . a, Astragalus ; c,
calcaneum ; c2, middle
cuneiform ; c3, outer
cuneiform ; cb, cuboid ;
n, navicular ; I-IV, first-
to fourth toes. (From
Flower's Osteology.)

486 CAPE JUMPING HARE CHAP.

Alactaga is much like Euchoreutes ; it has five toes, a
cylindrical tufted tail, the hairs at the end distichous, smooth
incisors, and a premolar present in the upper jaw. It also
differs from Euchoreutes by the much smaller auditory bulla as well
as in the fact that the infra-orbital foramen has no separate
passage for the nerve, which passage is to be distinguished in
both Dipus and Euchoreutes. The best-known species is the
Siberian Jumping Rabbit, A. jaculus. Beneath the ends of the
three main toes of the feet are remarkable fan-shaped pads. In
A. decumana the body and tail measure 7 and 1 0 inches re-
spectively, the ears 2 inches. Platycercomys, a fourth genus of
the family, is much less known and is to be differentiated from the
last three genera by the fact that it has no premolars at all, the
grinding tooth formula being thus -f. The tail too is flattened
and " lancet shaped." It extends from Siberia to Kubia, and
thus just enters the Ethiopian region.

The above are the more typical Jerboas. There remain
several forms which are not at all Jerboa-like in their way of
life, but are nevertheless, on anatomical grounds, placed with
them. Zapus, an American genus, with the exception of one
Palaearctic species, is transitional in that -its hind-legs are
rather long, but there is not so much difference between them as
in the typical Dipodidae. Sminthus is at the opposite extreme
to Dipus. Its feet are short and of equal length ; it climbs in
trees, and may perhaps be looked upon as nearest of all Dipodidae
to the ancestral form of the group.

Fam. 8. Pedetidae. — The genus Pedetes contains but one
species, P. ca/er, the Cape Jumping Hare. The animal suggests
a large Jerboa in appearance on account of its jumping habits,
the long hind-limbs, and the long tail. The length of a fair-
sized example is some 1 7 inches, with a tail of the same length.
The eyes and ears are large. The hands are five-fingered and
the feet only four-toed, the hallux being of course the absent
digit. In the skeleton it is interesting to note that the second
and third cervical vertebrae are so close together that there can
be no free movement ; interesting because in Dipus the cervicals
are actually ankylosed. The dorsal vertebrae are twelve. The
small intestine is long, measuring 7 feet 4 inches, while the
caecum is short, being only 8 inches long. The large intestine
is 3 feet 10 inches long. The gall-bladder appears to be

xv SOUTH AMERICAN RODENTS 487

absent/ an exceptional state of affairs in Eodents. A singular
fact in the anatomy of this animal is the existence of a septum
dividing the lower part of the trachea. This is sometimes met
with in birds. As might be supposed from its large eyes, the
Spring Haas, as the animal is sometimes called, is nocturnal. Its
long hind-limbs permit it to leap enormous distances. It is a
burrowing Eodent.

SECTION 3. HYSTPJCOMORPHA.

Fam. 1. Octodontidae. — The Eodents of this family are of
small to moderate size, the only, relatively speaking, giant in the
family being the "Water- Eat," Myocastor. The toes are with
but one exception not reduced ; the tail is long in the majority
of the genera. The teats are placed high up on the sides of the
body. The clavicle is fully ossified. All the genera are South
or Central American in range with the exception of Petromys?

Sub-Fam. 1. Octodontinae. — Octodon has four species, which
are all Chilian, Peruvian, and Bolivian in distribution. The
Degu, 0. degus, has a length of 160 mm., with a tail 105 mm.
long. The ears are 18 mm. long. At the roots of the claws
are longish and stiff hairs which appear to serve as "combs."
The tail has long but sparsely scattered hairs. There are twelve
pairs of ribs. The lengths of the various sections of the intestine
are as follows : small intestine, 680 mm.; caecum, 90 mm.; large
intestine, 390 mm. These animals live in large companies.
Closely allied is the genus Jfalrocoma (more correctly, as it
appears, to be written Abrocoma), with two species. H. lennetti
is 204 mm. long, with a tail of 103 mm. The ears are long,
22 mm. The fore-feet have no outward trace of the thumb.
Stiff hairs like those that characterise Octodon are found also in
this genus. The fur is very soft. The furring of the tail is
much thicker than in Octodon.

Spalacopus with but a single species, S. poeppigi, is a burrow-
ing animal, from which indeed, and on account of its resemblance
to Spalax, it has received its name. The ears in accordance with
the underground life . are short, only 5 mm. in length in an

1 Parsons, Proc. Zool. Soc. 1898, p. 858.

2 Very probably this form should be rather, as it is by Thomas, referred to the
neighbourhood of Peetinator, which would clear up the geographical anomaly.

488 CTENOMYS AND PETROMYS CHAP.

example of 120 mm. The tail too is reduced, being in the same
example only 42 mm. in length. As in the last two genera the
large intestine is about one half of the length of the small
intestine.

The " Tuco-tuco," genus Ctenomys, has also short ears and
tail. The claws of the fore-feet are longer than those of the
hind-feet.

A related form is Aconaemys (better known as Schizodon),
with similar external characters ; it inhabits high localities on
the Andes.

Petromys is the only genus of the sub-family which is not
American in habitat. It is an African form and there is but one
species. Its anatomy conforms to that of the genera already
considered. The main difference in structure is shown by the
teeth. Their surface is uneven, and differs from that of other
Hystricomorphs " in that the enamel to the inside of each upper
jaw - tooth and outside on each lower jaw - tooth forms two
tubercles, to which correspond grooves in the reverse position of
the applied teeth."

Sub - Fam. 2. Loncherinae. — The genus Echinomys with
thirteen species belongs to the Neotropical region. The members of
the genus are entitled " Spiny Eats " since they have spines mixed
with the fur. The tail is long and the ears are very well developed.
Both feet are five-toed. The tail is scaly as well as haired.
Trichomys (also called Nelomys) is very close to the above, and is
also from the same part of the world.

The genus Canncibateomys contains but one species, C.
amblyonyx, which was formerly included in the genus Dactylomys,
but has lately been separated by Dr. Jentink.1 The animal is
Brazilian and has a total length of 520 mm., of which 320 mm.
belong to the tail. It is a climbing rat, and in accordance with
that way of life has undergone some modifications. The fore-
feet are four-toed, the two middle toes being markedly longer
than the outer ones. The hind-feet are five- toed with the same
greater development of the two middle toes. The claws are
small and somewhat nail-like.

Dactylomys, also Brazilian, and with but one species, D.
dactylinus, differs from the last in the fact that the molars are
simpler in form ; they are divided into two lobes, each of which

1 Notes Leyd. Mus. 1891, p. 105.

xv THE AFRICAN GROUND-PIG 489

has but a single enamel fold, whereas in Cannabateomys these
teeth have several enamel folds. The tail, moreover, is but
slightly hairy.

Loncheres with eighteen species is another Neotropical genus
allied to the foregoing. Small spines are, as in many of these
genera, intermingled with the fur. This genus has as many as
seventeen dorsal vertebrae, which is an unusually large number.
L. guianae is known as the " Porcupine Eat." Allied genera, also
South American, and without spines in their fur, are Mesomys,
Cercomys, and Carterodon.

The South American Thrinacodus is also known by one
species,1 T. alMcauda, which has rather more than the distal half
of the long tail of a white colour. The fore-feet have four toes.
The ears are broad and short.

Sub-Fam. 3. Capromyinae. — A third sub-family of the Octo-
dontidae is formed by the genera Myocastor, Capromys, Plagio-
dontia, and Th'rynomys, which are all Neotropical forms with the
exception of the last, which is African.

Thrynomys (better known perhaps as Aulacodus) is a genus
of African Eodent, containing some four species. The best-
known of these is T. sivinderniamis, the Ground-Eat of West and
South Africa. Its structure has been investigated by Garrod,2
by Tullberg,3 and by myself.4 The fur is mingled with flattish
bristles ; the tail is moderately long, about half as long as the
body. The fore-feet are five-toed, but the two toes at each end
of the series are quite small. The hind-feet are only four-toed,
the hallux being absent. The claws of the hind-feet are stronger
than those of the fore-feet. The ears are not long. The limbs
are decidedly short, hence the name of " Ground-Pig " sometimes
applied to this animal. The molars are four in number in both
jaws. The incisors of the upper jaw are twice grooved. There
are thirteen dorsal vertebrae. The length of the small intestine
is 60^ inches, that of the large 49 ; the caecum is short, being
only 8 inches long. It is a remarkable fact that the acromion
is joined to the rest of the spine of the scapula by a joint.

Myocastor, a name which seems to have the rights of priority
over the more familiar Myopotamus, applies to a large South
American aquatic Eodent. The general aspect of the animal

1 Giinther, Proc. Zool. Soc. 1879, p. 144. 2 Proc. Zool. Soc. 1873, p. 786.

3 Loc. cit. (on p. 458), p. 123. * Proc. Zool. Soc. 1892, p. 520.

4QO 'SPECIES OF CAPROMYS -CHAP.

suggests a Water-Eat of large size (it has been exhibited in shows
as a phenomenal product of London sewers !) ; the tail is nearly
as long as the body. The ears are small. The limbs are short.
The tail is naked. The hind-feet are webbed, but not so much
so as in Hydromys. A small thumb is present. The animal has
thirteen pairs of ribs ; the molars are four in each jaw. The large
intestine is more than three times the length of the small, and
the caecum is, as in the last genus, relatively short.

Gapromys is a genus l which is remarkable on account of its
restricted distribution. It is found only in the islands of Cuba
and Jamaica. There are four species, of which C. melanurus is a
dark brown-coloured animal with a blacker tail, nearly as large
as a Rabbit. The native name of this Rodent is " hutia" It is
also remarkable for having a stomach more complicated than is
the rule among the mammals of this group. The organ is
divided by two constrictions into three compartments. In
C. pilorides the liver is occasionally divided up in an extra-
ordinary fashion into small lobules. Capromys has the large
number of sixteen dorsal vertebrae.

Fam. 2. Ctenodactylidae. — For these African genera it seems
admissible to form a distinct family, though Thomas, and Flower
and Lydekker, only allow to the genera Ctenodactylus, Pectinator,
and Massoutiera sub-family rank. On the other hand, Tullberg
removed these genera entirely from the Hystricornorph section
and placed them as a section of the sub-tribe Myomorphi of the
tribe Sciurognathi. It was chiefly the form of the mandible
which led to this placing, for in these Rodents, as in all Squirrel-
and Rat-like Rodents, and unlike what is found in the Hystrici-
form genera, the angular process of the mandible is not bent
sideways.

The genus Ctenodactylus derives its name from the peculiar
strong bristles which form a comb-like structure upon the hind-
feet and hide the claws ; these are stated to be for the purpose
of dressing the fur. The Gundi of North Africa, C. gundi, has a
length of 190 mm., with a short tail of 17 mm. The ears are
only moderate in size. The dental formula of the molars is f .
The incisors are white. The feet have four digits, and the hind-
lirnbs are the longer. The large intestine is distinctly longer
than the small intestine.

1 See Dobson, Proc. Zool. Soc. 1884, p. 233.

THE CARPINCHO

491

Pectinator spekii is the only representative of a genus not far
removed from Ctenodactylus ; it is a smallish Rodent, 6 inches in
length, exclusive of a rather bushy tail nearly 3 inches long.
It comes from Abyssinia. It has somewhat the appearance of a
Squirrel, which is heightened by the fact that when sitting the
tail is arched over the back ; when running the tail is carried out
straight. There are only four toes visible externally on both fore-
and hind-limbs, but pollex and hallux exist in the skeleton, with
a single phalanx each. There is only a single pair of mammae,
and in correspondence with this but two or three young are
produced at a time. The hind-feet have bristles very much like
those of Ctenodactylus. The molars, however, are ^. There are
twelve ribs, of which six reach the sternum. The latter is made
up of six pieces, and the manubrium in its breadth anteriorly
suggests that of the Yizcachas. The clavicles are present.1

Fam. 3. Caviidae. — This family, which includes the Cavies and

FIG. 240. — Carpincho. Hydrochoerus capybara. x fa.

the Capybara, is entirely South American and West Indian in dis-
tribution. It embraces animals of fair to large size, the Capybara

1 Peters, Trans. Zool. Soc. vii. 1871, p. 397.

492 ANATOMY OF HYDROCHOERUS CHAP.

(or Carpi ncho) being the greatest of existing Rodents. The ears are
well developed. The toes are commonly reduced, and the members
of this family possess only a rudimentary tail. The hair though
rough is not spiny. Other characters had best be deferred until
the several genera are treated of. We shall begin with the giant
of the family, the genus Hydrochoerus. This genus contains but
a single species, H. capybara, of South America. It reaches a
length of some 4 or 5 feet. The ears are not large ; the tail is
completely absent. The fore-feet are four-toed, the hind-feet
three-toed; the digits are webbed, though not to a very great
degree, and the nails have the appearance of hoofs. There are
fourteen dorsal vertebrae ; the clavicle is absent. In the skull
the paroccipital processes are of great length. The infra-orbital
foramen is large. The most remarkable fact about the teeth is
the great size of the posterior molar of the upper jaw ; it has
fourteen folds of enamel, more than all the anterior teeth possess
collectively. The incisors are white and grooved in front. The
measurements of the alimentary tract as given by Tullberg are : small
intestine, 4350 mm.; caecum, 450 mm.; large intestine, 1500 mm.
The Capybara or Carpincho is largely aquatic in its habits.

FIG. 241.— Patagonian Cavy. Dolichotis patachonica. x^\.

Their " favourite locality," writes Mr. Aplin,1 " is a broad laguna

"Field Notes on the Mammals of Uruguay," Proe. Zool. Soc. 1894, p. 297.

xv GUIANA PIG AND RESTLESS CAVY 49 3

in the river, furnished with open water, and also beds of
' camelotes,' — -a sloping open grassy bank on one side, where the
Carpinchos can lie in the daytime in the cooler weather,
sleeping and basking in the sunshine ; on the other a low
shelving bank, clothed with ' Sarandi ' scrub growing out into
the black reeking mud and shallow water beyond." They always
take to the water when alarmed, at a rate and with a gait which
reminded Mr. Aplin of a Pig. When in the water they swim
slowly with the upper part of the head, including nose, eyes, and
ears, above the surface. But they can dive for a considerable
time and distance, and baffle their enemies by seeking the shelter
of a mass of water-plants, and lying there with their noses only
just above the surface.

The genus Dolichotis l has long ears, and generally resembles
a rather long-legged Hare in appearance. The front>-feet are four-
toed, the hind three-toed. The Patagonian Cavy, as this animal
is called, has twelve dorsal vertebrae, and rudimentary clavicles.2
The paroccipital processes are long ; the incisors are white, and
are not grooved in front. The sternum has six pieces, and
seven ribs reach it.

Cavia, including the species C. porcellus, the Guinea-pig (which
name is a corruption apparently of Guiana pig), has the same
number of toes on its hind- and fore-feet as has the Capybara.
The name applied to the wild stock whence our Guinea-pig is
derived is the Eestless Cavy. The fur is greyish ; of the domestic
animals the colour is too well known to need description.

Fam. 4. Dasyproctidae. — The genus Coelogenys includes but two
species. C. paca, known as the "Spotted Cavy" or "Paca," has a
brown body, with white spots like those of a Dasyure ; it is one of
the largest of Rodents, and has a quite short tail. The hand and
foot are both provided with five digits ; but the thumb is small,
and in the foot the three middle toes considerably exceed the
others in length. The hind-foot is practically three-toed.- The
fibula is not nearly so reduced as in Dolichotis. The skull of
the animal is remarkable for the extraordinary development in
breadth of the jugal arch, which is sculptured externally. There
is a large cavity formed below, at the maxillary end of this huge

1 Beddard, Proc. Zool. Soc. 1891, p. 236.

2 These are stated by Tullberg to be absent. I have found them, but they are
very small bones, not more than half an inch long.

494 PACA AND AGOUTI

arch, by the curving inwards of the bone, which lodges a cavity
continuous with the mouth. The palate has anteriorly a ridge
on either side, and is thus divided from the sides of the face in a way
which is not found l in the allies of Coelogenys. Clavicles are
present. There are thirteen dorsal vertebrae. The incisors are
coloured red in front. The animal is South American, and in
that continent is limited to the Brazilian sub-region. This, the
best-known species of Paca, is called the Gualilla by the natives of
Ecuador ; in the same district another form is met with which the
natives term Sachacui (signifying Forest Cavy). It is very often
the case that a different native name expresses a real specific
difference; and to the latter form M. T. Stolzmann has given
the name of C. taczanowskii? This form, unlike the common
Paca, which is fond of forests and low-lying ground in the neigh-
bourhood of water, is alpine in habitat, living upon mountains
of 6000 to 10,000 feet. It burrows in much the same way as its
congener, and is greatly sought after as food, its meat possessing
an " exquisite taste." It is pursued by dogs, by whose aid one of
the two entrances to the burrow is guarded, and the creature is
smoked out and killed with a stick.

The genus Dasyprocta, containing those Rodents known as
Agoutis, is divisible into several species, apparently about twelve,
all of which are, like the Pacas, confined to the Neotropical region.
They have, however, a much wider range within that region, and
occur as far north as in Central America and in some of the
West Indian Islands. They are of rather smaller size than the
Paca, and are without spots. The colour is of a golden brown in
some forms, but usually has a freckled, grizzled, greenish kind of
appearance. The tail is stumpy, the hind-limbs are distinctly
longer than those of the Paca, and the two lateral toes have dis-
appeared from the feet— a concomitant as it seems of the Agouti's
greater powers of running. The three metatarsals are closely
pressed together, and the foot is as it were on the way towards
the highly-modified foot of the Jerboa. The fore-feet are, how-
ever, five-toed. The clavicle is rudimentary,3 whereas it is well
developed in the Paca. The skull has not the peculiar modifica-
tions of that of the last-mentioned type. The sternum has seven

1 There is a faint development of these ridges, but behind the palatine foramina
in Dasyprocta aguti.

2 Proc. Zool. Soc. 1885, p. 161. s Or absent ?

xv THE GENUS DINOMYS 495

pieces, and eight ribs reach it. A curious difference between this
genus and the last is in the relative proportions of the regions of
the intestine. The figures given by Tullberg for the two animals
are — for Coelogenys, small' intestine, 4800 mm. ; caecum, 230
mm.; large intestine, 21,000 mm.; — for Dasyprocta aguti the same
author gives: small intestine, 4200 mm.; caecum, 200 mm.; large
intestine, 1000 mm. The Agouti, says Mr. Eodway,1 is as wily as

FIG. 242. — Agouti. Dasyprocta aguti. x ^.

the Fox. " If chased he will run along the shallows of a creek to
hide his scent from the dogs, or swim over and back again several
times for the same purpose. He never runs straight when pursued,
but doubles, often hiding until a dog has passed, and then making
off in a different direction. Like the fox he has been hunted for
a very long period, and, like Eeynard, has grown wiser with every
generation."

Fam. 5. Dinomyidae. — The genus Dinomys of Dr. Peters 2 is a
very little known and remarkable form from South America
allied to the Capybara, the Chinchilla, and other South American
Kodents. It is only known by a single example found wandering
about a courtyard in a town of Peru. It is externally like, and
of about the same size as the Paca, but has a hairy tail. The
animal is four-toed and plantigrade ; the ears are short, and the
nostrils are S-shaped. It is usually regarded as belonging

1 In the Guiana Forest, London, 1894. 2 MB. Ak. Berlin, 1873, p. 551.

496 CUVIER'S CHINCHILLA CHAP.

to a separate family which will include but the one species,
D. branickii.

Fam. 6. Chinchiliidae. — This family, likewise South American,
contains three genera,1 all of which agree in having long limbs,
especially the hind-limbs, and a bushy and well-developed tail.
The hair is exceedingly soft, hence the commercial value of
" chinchilla."

The genus Chinchilla, containing but a single species, C. lani-
ger, is a small and squirrel-like creature, living at considerable
heights in the Andes. The eyes, as it is a nocturnal creature,
are naturally large ; and so also are the ears. The fore-feet have
five toes, the hind-feet only four ; they are furnished with feeble
nails. The innermost toe of the hind-foot has a flat and nail-like
claw. There are thirteen dorsal vertebrae, and the long tail has
more than twenty. The clavicle is well developed, as in the
other genera of this family. The large intestine of this animal
is extraordinarily long; the proportions of the different regions
of the gut are shown by the following measurements : small
intestine, 820 mm.; caecum, 125 mm.; large intestine, 1340
mm. Such a disproportion between the large intestine and the
small, to the advantage of the former, is a very strange fact in
the anatomy of this Eodent.

The genus Lagidium (also called Lagotis), which includes
" Cuvier's Chinchilla," is also a mountain dweller. There are
several species of this genus, which differs from Chinchilla,
by the complete abortion of the thumb and of the great toe.
The intestinal proportions are those of Chinchilla. The ears and
tail are long. L. cuvieri measures 1-|- feet in length.

Lagostomus, again, has but one species, L. tricliodactylus. The
animal has a tail about half the length of the body. The digits
are reduced as compared with Chinchilla, there being but four on
the fore- and three on the hind-feet. There are only twelve dorsal
vertebrae, and seven ribs reach the sternum. In the skull a dis-
tinguishing mark from the last two genera is the separation of
the infra-orbital foramen into two by a thin lamella of bone. The
large intestine is between one-half and one -third the length
of the small intestine, and thus differs much from that of
Chinchilla.

1 An account of the three genera is to be found in Trans. Zool. Soc. i. 1833,
p. 35, by Mr. E. T. Bennett.

xv SOCIAL HABITS OF THE VIZCACHA 497

The Vizcacha lives in societies of twenty to thirty members,1
in a " village " (" Vizcachera "), a dozen or so of burrows, which
intercommunicate. They lie at home during the day and come
out in the evening. Their burrows, like those of the Prairie
Marmot, harbour other creatures, which apparently live on ami-
cable terms with the Vizcachas ; such are the burrowing owl, a
small swallow, and a Geositta. The Fox also affects these burrows,
but then he ejects the rightful owner of the particular burrow

FIG. 243. — Vizcacha. Lagostomv.s trichodactylus. x^.

which he selects. When the young Foxes are born the vixen
hunts the Vizcachas for food. The Vizcacha has a most varied
voice, producing " guttural, sighing, shrill, and deep tones," and
Mr. Hudson doubts if there is " any other four-footed beast so
loquacious or with a dialect so extensive." These animals are
very friendly, and pay visits from village to village ; they will
attempt to rescue their friends if attacked by a Weasel or a
Peccary, and to disinter those covered up in their burrows
by man.

Fam. 7. Cercolabidae. — A number of the characters which
differentiate this family from the Hystricidae or Ground Por-
cupines of the Old World are given under the description of the
latter. The principal external characters are the prehensile tail,
the admixture of spines with hairs, and the nature of the sole of
the foot. In these points the New-World Cercolabidae differ
from the Old-World Hystricidae. It is interesting to notice that

1 Hudson, "On the Habits of the Vizcacha," Proc. Zool. Soc. 1872, p. 822.
VOL. X 2 K

498

TREE PORCUPINES

in both families we have long -tailed and short -tailed forms.
Cercolabes corresponds to Atherura or Trichys, and Erethizon
to Hystrix.

The genus Erethizon, the " Urson " of Canada, has a short,
stumpy tail. Its spines are almost hidden by enveloping hair.
The fore-feet have four, the hind-feet five toes. The short tail of
this creature is remarkable when we reflect upon its climbing
habits. It appears, hpwever, to be a weapon with which it
strikes sideways at the enemy.

Of the Neotropical genus Cercolabes (sometimes called Sphin-
gurus, Synetheres, or Coenclou) there are some eight or nine
species, all found in Central and South America. The animal is

FIG. 244. — Brazilian Tree Porcupine. Sphiiigurus prehensilis. xj.

arboreal, and has in correspondence with that habit a prehensile
tail. The spines are not so stout as in the Ground Porcupines,
and are often coloured yellowish or reddish. In correlation with
its tree-frequenting habits the bones of Cercolabes show certain
differences from those of the Ground Porcupines. The scapula is
broader and rounder in front than is that of Hystrix; the
phalanges of the thumb (which is rudimentary) are fused
together as in the Canadian Erethizon ; but those of the very
small hallux are also fused, whereas in Erethizon, as in Hystrix,
they are separate. In one species, C. insidiosus, Sir W. Flower
states that there are as many as seventeen dorsal vertebrae
and thirty-six caudals. The tail is thus very long. In C.
villosus there are fifteen dorsals and twenty-seven caudals ; eight
ribs reach the sternum, which is composed of seven pieces, the

xv HYSTRICIDAE OR GROUND PORCUPINES 499

sixth being very small. The clavicles are well developed. A
curious fact about C. villosus is that the acetabular cavity is
perforate (on both sides), or at least only closed by membrane.
In many forms of Eodents the bone is very thin in this region.
This fact perhaps lessens the significance of the perforation of the
acetabulum of Echidna (see p. 109).

Of the allied genus Chaetomys, also Neotropical, there is but a
single species, which inhabits Brazil. It has a nearly completely
closed orbit, a feature which differentiates it from the last animal,
and one which also shows it to be a more modified form. The
spiny covering is less pronounced than in its allies.

Fam. 8. Hystricidae. — This family is characterised by the fact
that all its members possess spines ; but the tail, if at all long,
is not prehensile, and the soles of the feet are smooth and not
covered with rough tubercles, as in the Tree Porcupines of the
next family, Erethizontidae. The clavicle is less developed than
in the arboreal forms. In the organs of digestion there are
points of a family difference between the two groups of spiny
Rodents. The tongue has serrated scales arranged in transverse
rows, which are directed backwards. A gall-bladder, though not
always present, is sometimes found ; it apparently never exists
in the arboreal Porcupines and in Erethizon. The lungs show
a great tendency to subdivision, which appears to be especially
marked in the genus Atherura. The caecum seems also to be
shorter in the Ground Porcupines. In Hystrix cristata the
small intestine measures 1 5 feet 7 inches ; the caecum, 8 inches ;
the large intestine, 4 feet 4 inches : — in Atherura africana the
caecum measures 7^- inches ; the large intestine, 1 foot 10
inches. The corresponding measurements of Sy nether es villosus
were : small intestine, 7 feet 3 inches ; caecum, 1 foot 4 inches ;
large intestine, 2 feet 7 inches. In Erethizon the caecum is
2 feet 4 inches in length. These differences are too large
and too constant in a number of presumedly allied forms to be
overlooked.

Mr. Parsons has directed attention l also to a number of
muscular differences, such indeed as might be expected to occur
between animals of such different habits.

The genus Hystrix embraces the better-known Porcupines. It
is a genus of wide range, extending from the East Indies to Africa,
1 Proc. Zool. Soc. 1894, pp. 251, 680.

500

SPINES OF PORCUPINE

and even occurring in Europe. There are several species, of
which the common Hystrix cristata is the best known, and is the
one which is to be found in Europe.

The spines of the common form and of the others are solid in
the middle of the body, but on the tail they are expanded into
hollow quills, which make much rattling. They are as a rule
black and white, the middle of the spine being banded with
black. A great crest of coarse long hairs on the head is
responsible for the scientific name of the well-known form.

FlQ. 245. — Common Porcupine. Hystrix cristata. x^.

Sometimes in this genus, as in the Tree Porcupines of Brazil, the
spines are orange or yellow ; but it is said that the colour is soon
lost in this country. As a matter of fact it is the easiest thing
in the world to wash out with ordinary tap-water much of the
yellow colour of the spines of the South American Sphingurvs.
The same may be the case with the pigment of the Old-World
Porcupines. There are fourteen to fifteen dorsal vertebrae and
four or five lumbars. The tail varies in length, but is shorter
than the long tail of the arboreal New- World forms. It seems
impossible when mentioning the Porcupine to escape from some
observations about its alleged habit of shooting its quills. For
some reason or other Buffon has got the credit of inventing, or at

xv BRUSH-TAILED PORCUPINE $01

least promulgating, this legend, which has even grown so in the
telling that the quills are said to be capable of penetrating planks
of wood. "What Buffon said apropos of this matter is, " The
marvellous commonly is pleasingly believed, and increases in
proportion to the number of hands it passes through." It is of
course the rattling of the spines and the occasional falling out of
loose ones which has started the legend. They are, however,
excellent weapons of offence, and the animal charges somewhat
backwards to make the best use of them against the foe. The
spines, however, are by no means an absolute protection, since, as
Mr. Eidley informs us,1 Tigers will kill and eat these animals just
as the Thylacine is apparently indifferent to the spiny armature
of Echidna.

Of the Brush-tail Porcupine, Atherura,2 there are at any rate
two species, the West African A. africana and the Malayan A.
fasciculata. It is interesting that the gap in the present distribu-
tion is partially filled by the discovery of fossil teeth near Madras.
The genus does not differ widely in external appearance from
Hystrix ; it has, however, a rather longer tail ; there are fewer
large spines, and there is a tuft of them at the end of the tail,
whence is derived the name of the genus. The frontal bones
project a little distance between the nasals, a feature which does
not seem to appear in the true Porcupines. There are fourteen
dorsal vertebrae and five lumbars. The twenty-four caudal verte-
brae of this Porcupine shows how much longer is its tail than
that of Hystrix ; for in the latter twelve is about the number.

A third genus of Old-World Porcupine is the singular Trichys?
Of this there is but one species, T. lipura. It is a curious fact
that out of three examples, all from Borneo, two were quite
without a tail. But this appears to be merely a mutilation,
though it is singular that the natives state it to be without a
tail. One cannot help thinking of the way in which lizards
sometimes shed their tails when pecked at. The tail of this
genus is more than half the length of the body and head. Trichys
has sixteen dorsal and six lumbar vertebrae. There is a tuft of
quills at the end of the tail, which are thin and compressed,

1 Nat. Science, vi. 1895, p. 94.

2 See Parsons. Proc. Zool. Soc. 1894, p. 675.

3 Gunther, Proc. Zool. Soc. 1876, p. 739, and 1889, p. 75 ; and Cederblom, Zool.
Jahrb. Syst. AUh. xi. 1897-98, p. 497.

5O2 HARES AND RABBITS CHAP, xv

though truncate at the free extremity and hollow ; they represent
in a more rudimentary way the much stronger tuft at the end of
the tail of other Porcupines. It is a curious fact that this and
other Porcupines possess a mechanism for warning their foes
precisely comparable to that of the rattlesnake. There are
sixteen dorsal vertebrae.

SUB-OKDER 2. DUPLICIDENTATA.

The chief feature of this group is the existence of two pairs
of incisor teeth in the upper jaw, of which the inner are very
small and lie behind the outer. In the skull the infra-orbital
foramen is small ; the incisive foramina are very large. The tail
is short or absent.

Fam. 1. Leporidae. — This family is distinguished from the
Lagomyidae by the long ears, by the tail, which is present, though
short, and by the longer limbs. There are six teeth belonging to
the molar series in the upper jaw, and five of the same in the
lower. The clavicle is imperfect.

The longest known genus of this family, Lepus, was, until the
quite recent discovery of Romerolagus, the only genus. It is of
universal range, excepting Australasia and Madagascar, and con-
sists of about sixty species. These are the Hares and Babbits, to
the former being assigned the longer-limbed forms.

As every text-book of zoology contains a more or less elaborate
account of the structure of the Common Eabbit, and as there is
but little structural difference between the members of the genus,
a short account of the generic peculiarities of Lepus will suffice
here. The fore-feet are five-toed, the hind-limbs four-toed. The
hairy integument enters the mouth cavity, and the inside of the
cheeks have a hairy covering. The soles of the feet are, more-
over, hairy. The maxillary bones are curiously sculptured.

The Common Eabbit, L. cuniculus, differs from the Common
Hare in the comparatively shorter ears and legs. The ears have
not, to so marked a degree, the black tips of those of the Hare.
The animal, moreover, produces naked young, and lives in burrows
of its own excavation. A difference in the structure of the
caecum, which distinguishes the Eabbit from the Hare, has been

inl.pei

AS

Q^ttan^pr°

ffltf.maaz

b.sp/v
"b.oc

FlG. 246. — Lepus cuniculus. Skull. A, Lateral view ; B, ventral view, ang.pro,
Angular process of mandible ; as, alisphenoid (external pterygoid process) ; aud.me,
external auditory meatus ; b.oc, basioccipital ; b.sj)h, basisphenoid ; cond, condyle ;
cor, coronoid process ; fr, frontal ; int.pa, inter-parietal ; ju, jugal ; Icr, lachrymal ;
max, maxilla ; nas, nasal ; opt.fo, optic foramen ; o.sph, orbitosphcnoid ; j)a, parietal ;
pal, palatine ; pal.max, palatine plate of maxilla ; pal.p.max, palatine process of
premaxilla ; par.oc, paroccipital process; peri, periotic ; p.max, premaxilla; pt,
pterygoid ; p.t.sq, post-tympanic process of squamosal ; s.oc, supraoccipital ; sq,
squamosal ; ty.btil tympanic bulla ; vo, vomer ; zyg.max, zygomatic process of
maxilla, (From Parker and Haswell's Zoology.)

504 THE RABBIT OF POPOCATEPETL CHAP.

pointed out by Professor W. N. Parker.1 These differences have
led some to approve of its separation from the Hares into a genus
Oryctolagus. This animal is believed to be an introduced species,
and to have been brought by man into these islands. Its original
home is the Spanish Peninsula, the south of France, Algiers> and
some of the Mediterranean islands. Mr. Lydekker thinks that
the only other species of Lepus which can be considered to be a
" Rabbit " is the Asiatic L. hispidus.

Of Hares there are two species in this country. The Common
Hare, L. europaeus (the name L. timidus seems to be really
applicable to another species to be referred to presently), extends
all over Europe excepting the extreme north of Eussia and
Scandinavia. It is not known in Ireland, and, curiously enough,
attempts to acclimatise this animal in that island have failed — a
state of affairs which contrasts with the fatal ease with which
the Rabbit has been introduced into Australia. Ireland has,
however, the Variable Hare, L. timidus (also called L. varidbilis),
a species which is common in other parts of Europe, and which
extends as far east as Japan. This species differs from its ally
by the fact that it often turns white in winter with the ex-
ception of the black tips to the ears. In Ireland this change
does not always occur ; but Mr. Barrett-Hamilton has commented
upon the fact that Hares of this species do change on Irish
mountains. It appears that in this animal the change from the
winter to the summer dress is accomplished by the actual casting
off of the white hairs and their replacement by a fresh growth
of " blue " hairs. A similar change occurs in the American
L. americanus.

Dr. Forsyth Major has noted the fact that the various species
of Hares can be distinguished by the condition of the furrows
upon the upper incisors. Thus two African species, L. crawshayi
and L. whytei, are to be separated by the fact that in the former
the incisors are quite flat, whereas in L. whytei the groove is
more prominent and there is a second shallow furrow.

The genus Romerolayus 2 is quite a recent discovery.- It
occurs on the slopes of Popocatepetl in Mexico; it has the
general aspect of the last genus, and is spoken of as a " Rabbit."
It inhabits runs in the long grass which clothes the sides of the

1 Proc. Zool. Soc. 1881, p. 624.
2 Proc. Biol. Soc. Washington, x. 1896, p. 169.

xv THE PIKA 505

mountain. Externally it is something like the Pikas, since it
has no tail visible. The ears, too, are short, and the hind-legs
comparatively short. The skull is very like that of the Rabbit ;
but in other osteological details it is aberrant. Thus the clavicle
is quite complete, and only six ribs articulate with the sternum,
instead of the seven that we find in the Rabbit.

Fam. 2. Lagomyidae. — The animals of this family are smaller
than the Hares and Rabbits ; they have short Vole-like ears and
no external tail. The limbs also appear to be shorter. As there
is but a single genus, the characters of the family may be de-
scribed in connexion with those of the genus, which is known as
Lagomys (apparently more correctly Ochotona). Of this genus
there are about sixteen species, which are mainly Asiatic ; one
species extends its range into Eastern Europe, and three are
North American.

The skull has not the supra-orbital grooves of the Rabbits,
and has a well-marked backward process of the zygomatic arch.
There are eighteen dorsal vertebrae. The molars and premolars
are five.

The vernacular names of " Pika " and " Piping Hares " have
been applied to the members of this genus, the latter on account of
their peculiar call. They live among rocks in companies and they
burrow. They are usually found at considerable altitudes : thus
L. roylei, the " Himalayan Mouse Hare," is found at elevations
as high as 16,000 feet; while L. ladacensis gets even higher,
19,000 feet having been recorded. With the habits of a Marmot,
so far as concerns living in burrows and at great altitudes, the
animals of this genus, with their squat form and short ears, are
not unlike those animals. In the past this genus occurred more
generally over Europe. Species from Miocene beds have been
met with in England, France, Germany, and Italy.

Fossil Rodents. — Quite a large number of existing genera of
Rodents are known from even the earlier strata of the Tertiary
period. The Squirrels (and even the genus Sciurus itself) occur in
the Upper Eocene. So, too, do the genera Myoxus, and (in South
America) Lagostomus. Spermophilus, Acomys, Hystrix, Lagomys,
Lepus, Hesperomys are known from Miocene rocks. Rhizomys,
Castor, Cricetus, Mus, Microtus, and some others appear to have
originated so far as we know in the Pliocene, while a still
larger series of existing genera are Pleistocene. It is interesting

506 EXTINCT RODENTS

to note that some of the extinct genera were much larger than
recent forms. At present, Hydroclioerus is the biggest Eodent ;
but the genus Megamys from the Pampas formation of Argentina
was " nearly as large as an ox." The wider range of genera in
the past is illustrated by Hystrix, which, now an Old-World
form, is represented by remains in the Miocene and Pliocene of
America.

It is a significant fact that of living genera Sciurus is the
oldest ; for it has been pointed out that in a number of features
the Squirrels are among the most primitive of Kodents. The
zygomatic arch is slender, and has thus not acquired the specialisa-
tion that is to be found in that part of the skull in other Kodents ;
moreover, the " jugal bone is not supported by any process from
the maxilla exactly as in the primitive Ungulata." The feet,
too, are unspecialised, though that is the case with many other
genera. It may also be pointed out that the teeth bear not a
little likeness to those of Ornithorhynclius in their multituber-
culate character. -

Some few fossil forms have already been dealt with in the
preceding pages.

The two genera Castoroides and Amblyrhiza, from the Pleisto-
cene of North America and the West Indies, are usually regarded
as forming a family. The skull of the former genus indicates an
animal of the size of a Bear. It is compared to that of Castor,
but it has a wide infra-orbital foramen. The teeth are four in
each jaw, and are formed of three to five lamellae ; the incisors of
this animal are powerful but short. Amblyrhiza, on the other
hand, has long incisors which are longitudinally grooved anteriorly.
It has a free fibula. This latter as well as other characters have
led Tullberg to remove it from association with Castoroides.

Order X. TILLODONTIA.

This group of Eocene mammals is to be defined by a number
of characters, of which the more important are the following : —
The incisors are enlarged, grow from persistent pulps, and are
coated with enamel upon the outer surface only ; they are those
of the second pair only, the first and third having disappeared
or become small. The canines are reduced in the later forms.

THE TILLODONTIA 507

These animals have been regarded as ancestral Kodents, to
which the tooth characters just mentioned clearly show like-
nesses. The earliest known form is Esthonyx. This genus shows

FIG. 247. — Tillotherium fodiens. Left lateral view of skull. (From Flower,
after Marsh.)

such primitive characters, compared with its later representatives,
as the existence of all three pairs of incisors in the upper jaw,
but only two in the lower jaw. The enlarged incisors of both
jaws do not seem to have grown from persistent pulps.

Ancliippodus, a later form, still preserves the upper pair of first
incisors in a vestigial form ; the strong second incisors grew from
persistent pulps. The most recent genus, Tillotherium, shows the
characteristics of the group at their height. The strong Rodent-
like, chisel -shaped incisors, which are reinforced by a small
additional pair in the upper jaws only, are persistent. The
grinding teeth are of the tritubercular pattern ; there are three
of each kind in the upper jaw, but in the lower jaw only two
premolars on each side. This is at any rate the case with some,
while others have three. The canine, though present in both jaws,
is insignificant. As in many ancient types, there is an entepi-
condylar foramen in the humerus. The feet were five-toed, and
bore sharp, laterally-compressed claws. The skull has been com-
pared in general aspect to that of a Bear.

CHAPTER XVI

INSECTIVORA CHIKOPTERA

Order XI. INSECTIVORA.

THE Insectivora1 are an order of mammals to which it is (to
quote Professor Huxley) "exceedingly difficult to give a definition."
They are, however, none of them large animals, and most of them
are nocturnal in habit — two circumstances which may have had
something to do with their survival from past ages, as may have
also their modification to so many and diverse modes of life ;
for everything points to the antiquity of the group. They are,
for instance, more or less plantigrade. The snout is generally
long, and is often prolonged into a short proboscis.2 There is
a tendency for the teeth to be of a generalised type, and
their number is often the typical mammalian forty -four.
Moreover, trituberculate teeth, which are certainly an ancient
form of tooth, are common ; and indeed the Insectivora of the
southern regions of the globe, e.g. Centetidae, Solenodontidae, and
Chrysochloridae, have the most prevalent trituberculism, a fact
which is of importance in considering the age of the animal life
of these regions of the world. The limbs are, as a rule, provided
with five digits. The hemispheres of the brain are usually
smooth, and do not extend over the cerebellum. The palate
is often fenestrate as in the Marsupials, and as in that group
the lower jaw is sometimes inflected. But the latter character
also occurs in the Sea-lions and elsewhere. Clavicles are present,
as a rule, but not in Potamogale.

1 See especially Dobson, A Monograph of the Insectivora, London, 1886-90.

2 Even in the Otter-like Potamogale the upper jaw, though broad and flat,
projects considerably beyond the lower.

CHAP, xvi THE HEDGEHOG FAMILY 509

There is, furthermore, a distinct tendency towards a dis-
appearance of functional milk teeth, which is best seen in
Sdrex, where there are only seven milk teeth, none of which ever
cut the gum. This suppression of the milk dentition is like that
of the Marsupials, Edentates, and Whales, all of which appear to
be — the first certainly are — ancient forms of mammalian life.

There is also a fairly well-defined, though shallow, cloaca in
many genera. Finally, the testes are purely abdominal in some,
and in none is there a full descent into a scrotum, as in the more
highly-developed Eutheria.

SUB-ORDER 1. INSECTIVOKA VEEA.

Fam. 1. Erinaceidae. — This family contains the genera
Erinaceus, Hylomys, and Gymnura.

Hylomys, considered by Dobson to fall within Gymnura, is
kept separate by Leche.1 H. suillus is a Malayan animal, small in
size, about 5 inches long, with a short tail. Like Gymnura it is
spineless. The ears are decidedly large and nude. There is one
pair of inguinal and one pair of thoracic teats. The colour
above is a rusty brown with yellowish-white under parts. The
palms and soles are quite naked. In its general form it recalls
Tupaia very much more than its own immediate relatives.
There is no doubt, however, of its systematic position when the
skeleton and teeth are examined. A variety has been described
from altitudes of 3000 to 8000 feet on Mount Kina Balu in Borneo.
It has the complete dentition of forty-four teeth. There are
fourteen pairs of ribs. As in Gymnura the tibia and fibula are
united below. The genus is considered by Leche to be the oldest
existing type of Erinaceidae.

Gymnura^ is also a Malayan form with the complete dentition
of the last, but with fifteen pairs of ribs and a longer tail, con-
sisting of twenty-three vertebrae as against fourteen. There is,
as with Hylomys, but one species, G. rafflesii. This animal has
a peculiar odour, resembling decomposed cooked vegetables.

1 "Bemerkungen iiber die Genealogie der Erinaceen." In Festschrift f.
Liljeborg, 1896. See also Anderson, Trans. Zool. Soc. viii. 1874, p. 453.

2 Dobson, "Notes on the Anatomy of the Erinaceidae," Proc. Zool. Soc. 1881,
p. 389.

510 SPINELESS HEDGEHOGS CHAP.

The under surface of the tail is rough, and it is thought by
Dr. Blanford that it may be of use to the animal in climbing.
Its compressed terminal third and the fringe of stiff bristles on
the under surface of this indicate, according to Dr. Dobson,
powers of swimming, or at any rate a not very remote ancestry
of swimming creatures. It is purely insectivorous in diet.

Erinaceus, including the Hedgehogs, is a widely distributed
genus — Palaearctic, Oriental, and Ethiopian in range. There are
about twenty species. The familiar spines distinguish the Hedge-
hogs from their allies, as also the fact that they possess but thirty-
six teeth, the formula being I £ C -f Pm 4 M -§•. There are fifteen

O ^ _|_ £ O

or fourteen ribs, and the tail is very short, consisting of only
twelve vertebrae. As in Gymnura there is no caecum. The upper
canine has usually, as in other Erinaceidae, two roots, but not in
E. europaeus, which is one of the most modified of Hedgehogs.

The Hedgehog is a more omnivorous creature than Gymnura.
It eats not only insects and slugs, but also chickens and young
game birds, and lastly vipers. Four, or in some cases as many
as five or six, young are produced at a birth ; they are blind,
with soft and flexible white spines. In hot and dry weather
Hedgehogs disappear ; they come forth in rainy weather. The
English Hedgehog, as is well known, hibernates. The Indian
species do not. The Hedgehog is occasionally spineless, which
condition may be regarded as an atavistic reversion.1

The Hedgehog has acquired the reputation of carrying off apples
transfixed upon its spines. Blumenbach thus quaintly describes
this and other habits of the animal, whose English name he gives
as " hedgidog " : " II se nourrit des productions des deux regnes
organises, miaule comme un chat, et peut avaler une quantite
enorme de mouches cantharides. II est certain qu'il pique les
fruits avec les epines de son dos, et les porte ainsi dans son
terrier."2

The Miocene Palaeoerinaceus is so little different from
Erinaceus that it is really hardly genetically separable.
Erinaceus is therefore clearly one of the oldest living genera
of mammals.

Necrogymnura of the same epoch and the same beds (Quercy
Phosphorites) is doubtless an ancestral form. The palate is

1 See Natural Science, xiii. 1898, p. 156.
- Manuel d'Hist. Nat. French trans, by Artaud, 1803.

•

xvi MIMICRY IN TUPAIA 5 I I

perforated as in Erinaceus (it is not so in Gymnura and Hylomys),
but on the whole it comes nearest to Hylomys.

Fam. 2. Tupaiidae. — This family contains the genera Tupaia
and Ptilocercus. Tupaia is Oriental in range, extending as far
east as Borneo. There are a dozen or so of species, which are
generally arboreal and have the outward aspect of Squirrels. It
has been suggested that this is a case of mimicry, the animal
gaining some advantage by its likeness to the Rodent. The
name Tupaia, it should be added, means Squirrel, and the long-
nosed Squirrel, Sciurus laticaudatus, is so extraordinarily like it
that " one has to look at the teeth " to distinguish them.
Moreover, this Squirrel, like some Tupaias, lives largely on the
ground among fallen logs. Tupaia resembles a Lemur in the
complete orbit. The dental formula is 1^ C^ Pmf M^= 38.
The sublingua, too, is stated by Garrod to be like that of
Chiromys. There is a minute caecum in T. belangeri, none in
T. tana.

Ptilocercus1 has a pen -like posterior portion to the tail, a
modification which is found in other groups of animals. The
tail of certain Phalangers, for instance, shows this same modi-
fication. The rest of the tail is scaly. The animal, as was
pointed out by Dr. Gray,2 looks very much like a Phalanger.
The orbit is entire as in Tupaia. The fingers and toes are five.
The one species, called after Sir Hugh Low, G.C.M.G., P. lowi, is
a Bornean animal.

Fam. 3. Gentetidae. — This family is entirely confined to the
Island of Madagascar. It includes some seven genera. The best-
known genus is Centetes. C. ecaudatus, the Tanrec, Tenrec, or
Tendrac, is an animal a foot or so in length, without a tail, and with
forty-four teeth.3 The immature animal is so different from the
parent as to appear quite a different form. It has three narrow
rows of spines along the back, which do not wholly disappear
until the permanent dentition has been acquired. Even then
the hairs are of a rather spiny character, particularly those upon
the back of the head, which are erected when the animal is

1 "Notes on the Visceral Anatomy of the Tupaia of Burmah," Proc. Zool. Soc.
1879, p. 301.

2 Proc. Zool. Soc. 1848, p. 23.

3 I quote Woodward, Proc. Zool. Soc. 1896, for this dentition. The fourth
molar of the lower jaw is not always present. It comes late, and only old animals
possess it.

512

CENTS TES AND ITS ALLIES

CHAP.

annoyed. The Tanrec feeds mainly upon earthworms. It is
"probably the most prolific of all animals," since as many as
twenty-one young are said to have been brought forth at a birth.
Some Opossums, however, have twenty-five teats.

Hemicentetes1 is a genus with two species. These have spines
mixed with the fur of the back. There is no caecum in this or

jo. maze-

FIG. 248. — Skull of Tenrec. Centetes ecaudatus. fr, Frontal ; max, maxilla ; pa,
parietal ; p.max, premaxilla ; sq, squamosal. (After Dobsou.)

in other Centetidae. The teeth are forty in number, there being
only three molars.

Ericulus setosus is a small Insectivore, resembling externally
a small Hedgehog. It is covered with close-set spines which,
unlike what is found in Erinaceus, extend over the short tail.
The total number of teeth is thirty -six, the formula being
If C}Pmf Mf.

Uchinops2 is another spiny genus which is a stage in advance
of Ericulus, for still another molar has been lost, reducing the
total number of teeth to thirty-two. The dental formula is thus
I -| C ^ Pm ^ M -|. The zygomata are reduced to mere threads.

Microgale, a genus recently instituted by Mr. Thomas, is a
small furry Insectivore with a long tail, which is more than
double the length of the head and body. There are no less than
forty-seven vertebrae in the tail, which is relatively longer than
that of any other mammal.

Limnogale, discovered by Forsyth Major, is an aquatic genus,
also furry and not spiny, which has departed from the Cente-
tid type and taken to an aquatic life. The single species,

1 Mivart in Proc. Zool. Soc. 1871, p. 58.

2 Thomas, Proc. Zool. Soc. 1892, p. 500.

xvi THE INSECTIVOROUS OTTER 5 I 3

L. mergulus, is about the size of Mus rattus ; it has webbed
toes and a powerful laterally - compressed tail. Clavicles are
present, which is not the case with Potamogale.

Oryzoryctes is a Mole -like Centetid. It has fossorial fore-
limbs, but a fairly long tail. This genus is furry like the last
two. It is said to burrow in the rice -fields and to do much
harm. The teeth are forty in number, three incisors and three
molars in each half of each jaw.

Fam. 4. Potamogalidae. — This family contains two genera,
Potamogale and Geogale.

Potamogale velox is a West African animal, which though an
Insectivore has the habits of an Otter. It is " somewhat larger
than a stoat." The upper surface of the body is dark brown,
the belly brownish yellow. It has a flat head and a long tail
like the Stoat, but the tail is laterally compressed and very
thick. The eyes are very small ; the nostril has valves. The
toes are not webbed ; but the second and third toes are united for
the whole length of their first phalanges. Along the outer side
of the foot is a thin extension of the integument. In swimming
the feet are drawn up along the body, hence webbing would be
of no use ; but the thin flattening prevents the edge of the foot
from acting as a hindrance to the motion of the animal.
M. du Chaillu describes it as catching fish, which it pursues
with extreme rapidity in the clear mountain streams it
frequents ; but Dr. Dobson, remarking that no stomachs have
been examined, thinks that water insects are more probably its
prey. It is not known whether the animal possesses a caecum.
The tooth formula l is 1 1 C ^ Pm f M f . The animal is excep-
tional among the Insectivora in having no clavicles.2 There are
sixteen ribs ; there is no zygomatic arch, and the pterygoids
converge posteriorly.

Geogale, with one species, G. aurita, is a small representative of
this family from Madagascar. It has only thirty-four teeth. When
better known it may be necessary, thinks Mr. Lydekker, to make
this animal the type of a separate family. The tibia and fibula
are distinct, not confluent with one another as in Potamogale.

Fam. 5. Solenodontidae. — This family contains but a single
genus.

1 Allman states the canines to be absent. I follow Flower and Lydekker. /<O/o° ° *

2 See Allman in Trans. Zool. Soc. vi. 1869, p. 1. /^"^ '^••^~<Cf

VOL'X 2LUJLIBRARY

5 1 4 SOLE NO DON

Solenodon. This genus, including two species, one from Cuba,
the other from Hayti, was at one time referred to the Centetidae.
It offers, however, numerous points of difference from the
members of that family with some general points of agreement.
Possibly its isolation in the two West Indian islands mentioned is
comparable to the isolation of the Centetidae in Madagascar;
they are both survivors of an ancient group of Insectivores
extinct elsewhere. Solenodon has nearly the complete dentition.
It has lost only one premolar, and has therefore forty teeth in
all. The formula is thus I | C ^ Pm ^ M ^. It also differs from
the Centetidae in having only two inguinal mammae instead of
both inguinal and thoracic ; the penis of the male does not project
from a cloaca, but lies forward. On the other hand, the molars
have their cusps arranged in the V-fashion of the Centetidae, a
fact, however, which, in the opinion of some, merely points to an
ancient trituberculism not indicative of special affinity. It has,
moreover, no zygoma in the skull, and there is no caecum.
Dr. Dobson has furthermore tabulated a number of differences in
muscular anatomy between the two families. Solenodon has a
long naked tail. The snout, always developed in Insectivores, is
extraordinarily long in this genus. It is a furry, not a spiny
animal. S. cubanus is liable to fits of rage when irritated, a
feature which it has in common with Shrews and Moles ; it is
also stated to have the ostrich-like way of concealing its head in
a crevice, " apparently thinking itself then secure." But nothing
is known of the genus in a wild state.

Fam. 6. Chrysochloridae. — This family contains only the
genus Chrysochloris, comprising some five species, all natives of
Africa south of the equator. The scientific name of the genus, and
also the vernacular name Cape Golden Mole, are derived from the
beautiful iridescent hairs which are intermingled with softer and
non-iridescent fur. Chrysochloris has V-shaped cusped teeth like
those that are possessed by the Centetidae and Solenodontidae.
In the skull as in the Macroscelidae, etc., but not in the Centetidae,
there are complete zygomata. They are Moles in habit, and the
eyes are covered with skin ; the ears, moreover, have no conches.
The teeth are forty or thirty-six in number, the reduction being
caused by the losing of a molar in those forms which possess the
smaller number.1 It is interesting to notice that the adapta-

1 The generic name of Chalcochloris was proposed by Dr. Mivart for these.

THE GOLDEN MOLE

515

tion to a digging life is brought about in quite a different way
from that of the true Moles (Talpa}. In the latter the fore-
limbs are changed in position by the elongation of the manubrium
sterni, carrying with it the clavicles, which are extraordinarily
shortened (Fig. 251). In Chrysochloris, on the other hand, the
same need (i.e. that the limbs project as little as possible from the
sides of the body, while the length of the limbs is retained, and the
leverage of the muscles unaffected) is provided for by a hollowing

FIG. 249. — Golden Mole. Chrysochloris trevelyani. A, Lower surface of fore-foot,
x J. (After Giinther. )

out of the walls of the thorax, the ribs and the sternum being
here convex inwards. The sternum and the clavicles are not
modified. The tibia and fibula are ankylosed below. In the
manus, moreover, there are but four digits, of which the two
middle ones are greatly enlarged. In the Moles there are five
fingers, and all are enlarged ; there is, too, a great radial sesamoid
bone, which is as good as a sixth finger (which, indeed, it is con-
sidered to be, in common with similar structures in other animals,
by some anatomists). The foot has only four toes.

Fam. 7. Macroscelidae.1 — This family contains three genera,
all of them African in range, and mainly Ethiopian.

Macroscelides, the Elephant Shrews, are jumping creatures of
Shrew-like appearance, combined with a Marsupial look. Both
radius and ulna, and tibia and fibula, are ankylosed. There

1 See Peters, Reise nach Mosambique, 1852, for external characters and anatomy.

5i6

RHYNCHOCYON AND PETRODROMUS

CHAP.

are five fingers and toes. There is a caecum as in but few
Insectivores. The tooth formula, as revised by Thomas,1 is
I »- C \ Pin £• M -2-£-g, the total number being thus forty or
forty-two. There are several species of this genus.

Fia. 250. — Rhynchocyon chrysopygus. x £. (After Giinther.)

Rhynchocyon and Petrodromus differ from Macroscelides in
not having such long hind-legs. The dental formula of the
first is I-MP C ]- Pmf Mf = 34 or 36, of the latter If C i-
Pm 4- M £ = 40. In Petrodromus the toes are reduced to four ;

4 ^

in Rhynchocyon there are only four digits in the manus as well
as in the pes. This animal, as its name implies, has a longish
proboscis, which can be bent, and is really very like a miniature
Elephant's trunk, and also like that of the Desman (Myogale}.
It has thirteen pairs of ribs, and a well-developed caecum. Dr.
Giinther has pointed out that in Petrodromus tetradactylus the
hairs of the lower part of the tail are stiff elastic bristles 5 mm.
long, with a swelling at the free tip. The use of this singular
modification is not at all apparent. Pseudorhynchocyon, of
European Oligocene, is believed to be related to this family.

Fam. 8. Talpidae. — This family is confined to the Palaearctic
and Nearctic regions, or practically so, being fairly equally dis-

" Mammals collected by Dr. Emm Pasha," in Proc. Zool. Soc. 1890, p. 446.

XVI

SKELETON OF MOLE

517

tributed as regards genera ; a Mole just gets over the boundary
into the Oriental region. The genus Tcdpa is entirely Old World
in range, and includes several species, of which the Common Mole,
T. europaea, is the best known. There are forty teeth, one of
the molars of the full mammalian dentition not being represented.
In the milk dentition there is an additional preniolar, not repre-

FIG. 251. — Sternum and sternal ribs of the FIG. 252. — Bones of fore -ami and

Common Mole (Talpa europaea), with the
clavicles (d) and humeri (Jf) ; M. Manu-
brium sterni. Nat. size. (From Flower's
Osteology.)

mantis of Mole (Talpa europaea).
x 2. C, Cuneiform ; ce, centrale ;
I, lunar ; m, magnum ; p, pisi-
form ; R, radius ; rs, radial sesa-
moid (falciform) ; s, scaphoid ; td,
trapezoid ; tin, trapezium ; U,
ulna ; it, unciform ; / - 1', the
digits. (From Flower's Osttdogy.)

sented by a successor in the permanent dentition. The formula
is thus If C \ Pm % M §. There are no external ears, and the
eyes are rudimentary ; the soft silky fur is familiar to everybody.
The sternum has a strong crest, associated with a powerful
development of the pectoral muscles, so necessary to a burrowing
animal. The animal, it is hardly necessary to state, lives under-
ground in burrows excavated by itself, which have not, it has
been stated, the elaborate and, it appears, fanciful shape assigned
to them by many writers. At times Moles appear above ground.

5 I 8 AMERICAN MOLES

Their principal food consists of earthworms, and it may not be
out of place to quote Topsell's quaint account of their pursuit of
the annelids : " When the wormes are followed by molds (for by
digging and heaving they foreknow their owne perdition) they
fly to the superficies and very toppe of the earth, the silly beast
knowing that the molde, their adversary, dare not followe them
into the light, so that their wit in flying their enemy is greater
than in turning againe when they are troade upon." It has
lately been said l that Moles store up earthworms for consump-
tion during the winter, biting off their heads to prevent their
crawling away.

Scalops, an American genus, is a Mole-like creature of largely
aquatic habits, as its webbed hind-feet show ; it has a short, naked
tail. Apparently, like the Shrews, it has no lower canines.

Condylura, another American genus, is called the Star-nosed
Mole on account of a curious radiating structure at the end of
the snout.

Myogale, the Desman, is still more aquatic in habit, and
connects the Moles with the Shrews, though, as in many of the
former, it has lower canines. It has webbed hind-feet and a long-
tail. One species occurs in the Pyrenees, the other in Eussia. A
few other genera (Urotrichus, Uropsilus, Scaptonyx, Dymecodon,
Scapasius, Perascalops) belong to the same family.

Fam. 9. Soricidae. — The true Shrews have a much wider
range than other families of the present order. In the Palaearctic
region are found Sorex, Crossopus, Crocidura, Nectogale, Chimar-
rogale. The first is also Nearctic, and reaches Central America.
In the Ethiopian region is the single peculiar genus Myosorex, but
Crocidura occurs there also. Blarina and Notiosorex are " Sonoran "
in range ; Soriculus Oriental. Crocidura, Anurosorex, and Chimar-
rogale also enter this region. Sorex has teeth tipped with reddish
colour, its dental formula being, according to Mr. Woodward's
recent researches, I -^jr^ C^ Pmf Mf = 32 or 34.

As compared with other Insectivores, therefore, the most
remarkable fact found throughout the family is the absence of the
lower canines. In addition to this the genus may be known —
the family indeed — by the large size of the first pair of incisors.
In the above formula it is possible, thinks Mr. Woodward, that
there may be errors; he is not certain whether the supposed

1 Ritsenia Bos, Biol. Centralbl. xviii. 1898, p. 63.

xvi TOPSELL ON THE SHREW 5 I 9

upper canine may not be a fourth incisor, and whether the
first premolar may not be really the canine. Another peculiar
feature about the dentition of Sorex is the suppression of the teeth of
the milk dentition, which are functionless, and probably uncalcified.
The genus Sorex is terrestrial. The tail is long and covered with
hairs. There are two species in this country, S. vulgaris and 8.
minutus. The former is the Shrew of legend and superstition ;
and it is no doubt the species that has lent its name to the more
untameable members of the softer sex, though it is the males
which are especially pugnacious. As to legend, everybody has
heard of the shrew ash whose leaves, after a Shrew has been
inserted living into a hole cleft in the tree, are a specific for
diseases of cattle, caused by the Shrew itself creeping over
them.

The Eev. Edward Topsell, author of The Historie of Four-
footed Beastes, who defends his veracity by asserting that he does
not write " for the rude and vulgar sort, who being utterly
ignorant of the operation of learning, do presently condemne al
strange things," says of the Shrew that " it is a ravening beast,
feigning itself gentle and tame, but, being touched, it biteth deep
and poysoneth deadly. It bearetli a cruel minde, desiring to hurt
anything, 'neither is there any creature that it loveth, or it loveth
him, because it is feared of all." It is probable that all this
rustic feeling is due to the powerful effluvium which the Shrew
undoubtedly emits.

S. minutus has the distinction of being the smallest British
mammal ; it is scarcer than the last. This form is found upon
the Alps, as is also the peculiarly Alpine species S. alpinus, which
inhabits the Alps, Pyrenees, Carpathians, and the Hartz.

Crossopus fodiens, the Water Shrew, has also brown-stained
teeth. - It is not uncommon in this country, and lives in burrows
excavated by the sides of the streams which it affects.

Besides these two genera, Soriculus, Blarina, and Notiosorex
have red-tipped teeth. In Crocidura, Myosorex, Diplomesodon,
Anurosorex, Chimarrogale, and Nectogale the teeth are white-
tipped. These are all the genera of the family allowed by the
late Dr. Dobson in a review of that family.1

Chimarrogale and Nectogale are aquatic genera. The former

1 "A Synopsis of the Genera of the Family Soricidae," Proc. Zool. Soc. 1890,
p. 49.

52O

GALEOPITHECUS

CHAP.

consists of a Himalayan and Bornean, and of a Japanese species,
which have not webbed feet, but have a tail with a fringe of
elongated hairs.

Nectogale elegans is one of the characteristic animals of the
Thibetan plateau. It has webbed feet. The teeth are as in
Chimarrogale I f C ^ Pm y M §.

The other genera are terrestrial in habit.

SUB-ORDER 2. DEEMOPTEEA.

The family Galeopithecidae contains but one genus, which
has been at times referred to the Lemurs, to the Bats, or has been

Fio. 253. — Galeopithecus volans. x ^. (After Vogt and Specht.)

made the type of a special order of mammals. It is better to
regard it as an aberrant Insectivore — so different indeed from
other forms that it requires a special sub-order for its reception.

xvi INCISORS OF GALEOP1THECUS 5 2 I

Galeopithecus * inhabits the Oriental region. It is a larger
animal than any other Insectivore, about the size of a Cat, and
has a patagium extending between the neck and the fore-limb,
between the fore-limb and the hind-limb, and between the hind-limb
and the tail. This patagium is abundantly supplied with muscu-
lature, but the fingers are not elongated as in the Bats for its
support. In the degree of its development, however, the pata-
gium of this creature is midway between that of Sciuropterus
on the one hand, and the Bats on the other. It presents many
remarkable features in its organisation. The brain is like that of
the Insectivora in the exposure of the corpora quadrigemina by
the slight extension backward of the cerebral hemispheres ; but
its upper surface is marked by two longitudinal furrows on each
side, a state of affairs (in combination) which is unparalleled
among the Mammalia. The teeth are peculiar by reason of the
singular " comb-like " structure of the lower incisors. This, how-
ever, is an exaggeration of what is to be found in Rhynckocyon
and Petrodromus, while the same style of tooth, though not so
highly developed, characterises certain Bats. The Tupaiidaeand
certain Lemurs show what Dr. Leche regards as the beginning of
the same thing. As in Tupaia also there is an indication of
the characteristically Lemurine sublingua. The stomach is more
specialised than in other Insectivores, the pyloric region being
extended as a narrowish tube. There is a caecum. A peculi-
arity of the intestinal tract is that the large intestine is longer
than the small.

Order XII. CHIROPTERA.

We may thus define the Bats : — :Flying mammals, with the
phalanges of the four digits of the hand following the pollex
greatly elongated, and supporting between themselves and the
hind -limbs and tail a thin integumental membrane, which
forms the wing. The radius is long and curved ; the ulna rudi-
mentary. The knee is directed backwards, owing to the rota-
tion of the limb outward by the wing membrane. From the
inner side of the ankle-joint arises a cartilaginous process, the
calcar, which supports the interfemoral part of the wing mem-

1 Leche, " Uber Galeopithecus, " K. Svensk. Ak. Handl. 1886.

•

522 CHARACTERS OF BATS CHAP.

brane. The mammae are thoracic ; the placenta discoidal and
deciduate. The. cerebral hemispheres, which are smooth, do not
extend over the cerebellum.

This large order of mammals was once placed with the Primates.
There is no doubt, however, that they form a perfectly distinct
order; no knowledge of fossil forms in any way bridges over
the gap which distinguishes them from the highest mammals.
The most salient feature in their organisation is clearly the
wings. These consist of membrane, an expansion of the in-
tegument, provided with nerves, blood-vessels, etc., which mainly
lie stretched between the digits 2 to 5. These digits themselves,
which are enormously elongated, act like the ribs of an umbrella,
and when the wing is folded they come into contact. Besides

FIG. 254. — Barbastelle. Synotus barbastellus. x \. (After Vogt and Specht.)

this part of the flying apparatus there is a tract of membrane
lying in front of the arm, which corresponds to the wing mem-
brane of the bird, but which in the Bats takes quite a subordi-
nate place. In the bird, on the other hand, there is a metapa-
tagiuin, which is the main part of the wing of the Bat. It
seems just possible that in Archaeopteryx the metapatagium was
more Bat -like. Furthermore, a steering membrane, like that
which fringes the tail in some Pterosaurians, lies interfemorally
in Bats, and includes the whole or a part of the tail. The pollex
takes no share in the wing, but projects, strongly armed with a
"claw, from the upper margin.

The bones of this order of mammals are slender and marrowy ;
they are thus light, and subserve the function of flight. A
most remarkable feature among the external characters of the Bat
tribe is the extraordinary and often highly complicated mem-
branes which surround the nostrils. These are at least often

SKELETON OF BATS

523

more strongly developed
in males than in females,
and may perhaps be
partly relegated to the
category of secondary
sexual characters. But
it seems that they have
also an important tactile
function, and enable the
creatures to fly without
touching bodies which
intrude themselves upon
their way. The ears,
too, are frequently very
large, and it may be
supposed that the sense
of hearing is correspond-
ingly acute. In the
common Long-eared Bat
of this country, the ears
are not greatly inferior
in length to the head
and body of the animal
combined. The ears are
of every variety of shape,
and offer characters
which are valuable in
the systematic arrange-
ment of the members of
the order.

In the skull of Bats
there is very rarely a
complete separation be-
tween the orbital and
temporal fossae ; the
lachrymal duct is out-
side the orbit. The
tympanics are annular,
and in a rudimentary
condition. The centra

524

FLYING FOXES

of the vertebrae tend to become ankylosed in old individuals ;
the caudals have no processes, but are like those quite at the end
of the series in long-tailed animals. The sternum is keeled for
the better attachment of the pectoral muscles, the chief muscles
of flight. The ribs, which are much flattened, are occasionally
ankylosed together by their margins. There is a well-developed
clavicle. In the carpus the scaphoid, lunar, and cuneiform are
all fused together. In the hind-limb the fibula is rarely fully
developed.

The Bats are divisible into two primary groups, which are
those of the Megachiroptera and the Microchiroptera.

SUB-ORDER 1. MEGACHIKOPTEKA.

The Pteropodidae are frugivorous Bats, usually of large size.
The chief distinguishing feature is the fact that the molars
are not tubercular, but marked with a longitudinal furrow,
which is, however, concealed in the genus Pteralopex by cusps.
The palate is continued back behind the molars. The index
linger has three phalanges, and is usually clawed. The ears

FIG. 256. — Skull of Plerojnts fuscus. x §. (After de Blainville.)

are oval, and the two edges are in contact at the base of the
ear. The tail, if present, has nothing to do with the inter-
femoral membrane. This group is entirely Old World in range.
The genus Pteropus embraces the creatures known as Flying Foxes.
They are the largest forms in the sub-order, sometimes having
an expanse of wing of 5 feet (this is the case with P. edulis}.
The muzzle is long, and the face therefore " foxy " in appearance.

DISTRIBUTION OF PTEROPUS

525

The inner margin of the nostrils projects, a preparation for the
tubular nostrils of Harpyia. The tail is absent. The pre-
molars are three and the molars two. The pyloric region of the
stomach is extended and twisted upon itself. Of this genus
there are nearly sixty species, extending from Madagascar to
Queensland. Thirty species inhabit the Australian, twenty the
Oriental region. Madagascar has seven, and one species just
enters the Palaearctic. The occurrence of this genus in India
and in Madagascar is one of those facts which favour the view
supported, on these and other grounds, by Dr. Dobson and Dr.
Blanford. that a connexion between India and Madagascar must
once have existed ; for these slow-flying creatures could hardly
be believed capable of traversing vast stretches of ocean by
their unaided efforts.1

Pteropus is represented in the Ethiopian region by the allied
genus Epomophorus. Of this there are perhaps a dozen species.

FIG. 257. — Flying Fox. Pteropus poliocephalus. x £.

The teeth are reduced to two premolars in the upper jaw,

1 See Dobson, Ann. Nat. Hist. (5) xiv. 1884, p. 153.

526 GENERA OF FRUIT BATS CHAP.

three remaining below; while there is but one molar in each
upper jaw, and two in each lower. Dr. Dobson has studied
the structure of the remarkable pharyngeal sa'cs which exist in
the neck of the male, and are capable of inflation.

Pteralopex of the Solomon Islands has shorter ears than have
many Pteropus, otherwise its external characters are the same.
As in Pteropus nicobaricus, this genus has the orbits shut off by a
bony ring, an extremely rare phenomenon in Bats. The canines
have two cusps. The characters of the grinding teeth have
already been mentioned. It is uncertain whether the only
species of this genus, P. atrata, is, or is not, a vegetable feeder.
Harpyia has shortish ears and extraordinarily prolonged and
tubular nostrils. There is a hint of the accessory cusp to the
canines mentioned above in Pteralopex. The incisors are reduced
to one on each upper jaw, and none below. Cynopterus has also
often bituberculate canines. It is an Oriental genus with several
species.

Nesonycteris, with one species from the Solomon Islands, N.
ivoodfordi. has the dental formula I ^ C ^ Pm |^ M -|. The index
finger has no claw ; the tail is absent. The premaxillae are
separated anteriorly.

Eonycteris, with a single cave-dwelling species from Burmah,
E. spelaea, has also no claw upon the index ; the tooth formula
is fuller by reason of the presence of an additional incisor below.
The tongue is very long and is armed with papillae. There is a
short but distinct tail.

Notopteris, from New Guinea and the Fiji Islands, is dis-
tinguished from the related genera by its long tail.

The remaining genera of Fruit Bats are Boneia, Harpyionyc-
teris, Cephalotes, Callinycteris, and Macroglossus, from the Oriental
region, and Scotonycteris, Liponyx, and Megaloglossus from the
Ethiopian region ; finally, there is the Australian Melonycteris.

SUB-ORDER 2. MICEOCHIEOPTEEA.

The members of this sub-order are mostly insectivorous though
occasionally " frugivorous or sanguivorous " Bats. The molars are
multicuspid with sharp cusps. The palate is not continued back
behind the last molar. The second finger has but one phalanx, or

HORSE-SHOE BATS 527

none ; occasionally there are two. It has no claw. The ear has
its two sides separate from their point of origin upon the head.
The group is of Old- World distribution.

Fam. 1. Rhinolophidae. — The Bats of this family possess the
leafy outgrowths around the nostrils. The ears are large, but
have no tragus. The index finger has no phalanx at all. The
premaxillary bones are quite rudimentary, and are suspended from
the nasal cartilages. In addition to the pectoral mammae they
have two teat-like processes situated abdominally. The tail is
long, and extends to the end of the interfemoral membrane.

The genus Rhinolophus has a large nose leaf, and an anti-
tragus to the ear. The first toe has two joints, the remaining toes
have three joints each. The dentition is I ^ C ^ Pm ^ M -|. There
are nearly thirty species of the genus, which are restricted to the
Old World. Two species occur in this country, viz. It. ferruin
equinum, the Great Horse-shoe Bat, and the Lesser Horse-shoe Bat,
JR. hipposiderus. The name is of course derived from the shape
of the nose leaf.

The genus Hipposiderus and some allied forms are placed
away from Rhinolophus and its immediate allies in a sub-family
Hipposiderinae. The type genus Hipposiderus, or, as it ought
apparently to be called, Phyllorhina, is Old World in range, like
all the other members of the family.

The nose leaf is complicated, and there are only two phalanges
in all the toes ; there is no antitragus to the ear. A curious
feature in the osteology of the genus, and indeed of the sub-family,
is the fact that the ileo-pectineal process is connected with the
ilium by a bony bridge ; this arrangement is unique among
mammals.

The genus Anthops, only known from the Solomon Islands,
and represented there by but a single species (A. ornatus}, has
an extraordinarily complicated nose leaf. The tail, like that of
the Oriental Goelops, likewise represented by a single species
(C. frithii}, is rudimentary.

Triaenops, Ethiopian and Malagasy, has, like the Australian
Rhinonycteris, a well-developed tail. Triaenops has also a highly-
complicated nose leaf.

Fam. 2. Nycteridae. — This family is to be distinguished from
the Rhinolophidae by the fact that the ear has a small tragus,
and by the small and cartilaginous premaxillae. In addition to

528 THE SEROTINE AND PIPISTRELLE CHAP.

these two characters it may be added that the nose leaf is well
developed, but is not so complicated as in the last family. The
type genus Nycteris is Ethiopian and Oriental, nine species being
African, and only one, N. javanica, being, as the specific name
denotes, from the East. Megaderma is to be distinguished by the
loss of the upper incisors. There is no tail, and the ears are
particularly large. They are carnivorous Bats, and M. lyra, called
the " Indian Vampire Bat," chiefly affects frogs as an article of diet.

Fam. 3. Vespertilionidae. — This family has not the nose
leaf of other families. The apertures of the nostrils are simple,
round, or crescentic apertures. The ear has a tragus, and the tail
is not produced to any great degree behind the interfemoral
membrane. There are two phalanges to the index digit.

This family in numbers of species is vastly in excess of any
other family of Bats. The most recent estimate, that of P. L.
and W. L. Sclater, allows 190. But the generic types are by no
means so numerous as in the Phyllostomatidae. This is a signi-
ficant fact when we reflect upon the geographical range of the two
families. The Vespertilionidae range over the whole earth, while
the Phyllostomatidae are practically limited to the South American
continent, only just getting into the Nearctic region. They in-
habit, therefore, a more restricted area, and, in consequence of
competition, have specialised more freely than the widely -spread
and therefore not crowded Vespertilionidae.

The genus Vesperugo is by far the largest genus of this family,
embracing no less than seventy species. The tail is shorter than
the head and body together ; the ears are separate, and moderate
or short in size ; the tragus is generally short and obtuse. The
dentition is I 2, C 1, Pm 2 or 1, M 3. It is a remarkable fact
that this genus, unlike most Bats, produces two young at a time.
The genus is universal in range, and one species, the Serotine Bat,
known in this country, even ranges from the New World to the
Old ; but with so small a creature the possibility of accidental
transportation by man must not be left out of sight. The British
species are — V. serotinus, the Serotine already mentioned; V.
discolor, a single example only of which has occurred, and may
have been introduced; V. noctula, the habits of which were
described by Gilbert White ; V. leisleri ; and the Pipistrelle, V.
pipistrellus, which is the best-known member of the genus in this
country.

xvi THE LONG-EARED AND OTHER BATS 529

The genus Vespertilio contains some forty-five species, and is
world-wide in range. It has one more premolar in the upper
jaw than has Vesperugo. There are no less than six British
species, of which V. murinus is the largest species of Bat recorded
from this country, but is not quite certainly indigenous.

Plecotus has very long ears. The dentition is I f C ^ Pm f M §.
The tragus is very large. There are but two or possibly three
species, of which one is North American, and the other is the
Long-eared Bat, P. auritus, of this country, but ranging as far as
India. The shrill voice, inaudible to some ears, of this Bat has
been heard of by everybody.

Synotus includes the British Barbastelle, S. barbastellus, as
well as an Eastern form. It differs from the last genus principally
by the loss of a lower premolar. The ears, too, are not so large.
Otonycteris, Nyctophilus, and Antrozous are allied genera; the
last is Californian, the others Old- World forms.

Kerivoula (or Cerivoula) has a long, pointed, narrow tragus.
The tail is as long as or longer than the head and body. The
dentition is as in Vespertilio ; but the upper incisors are parallel
instead of divergent as in that genus. The brilliantly-coloured
K. picta is, on account of this very fact, the best-known species.
The name Kerivoula, a corruption of the Cinghalese " Kehel vulha,"
signifies plantain bat. This Bat has been described as looking,
when disturbed in the daytime, more like a huge butterfly than a
Bat, which is naturally associated with sombre hues. Other
species occur in the Oriental, Australian, and Ethiopian regions.

Miniopterus has a premolar less in the upper jaw; it has a
long tail as in the last genus. One species, M. scheibersi, has
almost the widest range of any Bat, it being found from South
Europe to Africa, Asia, Madagascar, and Australia.

Natalus is an allied form from Tropical America and the West
Indies. It is chiefly to be separated from Kerivoula by the short
tragus to the ear.

Thyroptera is also South American. It is distinguished by
the curious sucker-bike discs upon the thumb and foot. These
" resemble in miniature the sucking cups of cuttle-fishes." The
Madagascar genus, Myxopoda, with but one species, has also an
adhesive but horse-shoe-shaped pad upon the thumb and foot.

Scotophilus has shortish ears with a tapering tragus. The tail
is shorter than the head and body, and is nearly contained within

VOL. x 2 M

53O BATS WITHOUT A NOSE LEAF CHAP.

f

the interfemoral membrane. The dentition is I £ C ^- Pm ^ M f ,
with another upper incisor in the young. It is African, Asiatic,
and Australian.

This genus appears to be connected with Vesperugo by Mr.
1 )obson's proposed genus, or sub-genus as it is generally held to be,
Scotozous.1 The genus Nycticejus, founded for the inclusion of
Scotozous dormeri, an Indian species, should, according to Dr.
Blanford, replace on grounds of priority the name Scotophilus.
But as this name (Nycticejus} is one introduced by Eafinesque,
whose work was so uncertain and untrustworthy, it seems prefer-
able to retain the better-known name of Scotophilus, introduced
by William Elford Leach.

The genus Chalinolobus 2 has short, broad ears with an
expanded tragus. A distinct fleshy lobule projects from the
lower lip on either side of the mouth. The tail is as long as the
head and the body. The dental formula is 1 1 C ^ Pm -| or -J M |.
The genus occurs in Africa, Australia, and New Zealand ; but the
African species, with diminished premolars and pale coloration,
have been distinguished as Glauconycteris.

Fam. 4. Emballonuridae. — The Bats belonging to this family
have no nose leaf. The tragus is present, but often very small.
The ears in this family are often united. There are two phalanges
in the middle finger. The tail is partly free, either perforating
the interfemoral membrane and appearing upon its upper surface,
or prolonged beyond its end. The face is obliquely truncated in
front, the nostrils appearing beyond the lower lip.

Eniballonura is Australian, Oriental, and Mascarene in range.

O

The ears arise separately, and there is a fairly developed and
narrow tragus. The tail perforates the interfemoral membrane.
The dental formula is I f C { Pm f M -|.

Rhinopoma has the ears united ; the incisors are reduced by
one on each side of each jaw, and the premolars are similarly
reduced, but only in the upper jaw.

Noctilio is an American genus of two or three species, which has
one pair of markedly large upper incisors, which completely con-
ceal the outer pair. On these grounds this Bat was removed from
its allies and placed by Linnaeus among the Rodents, an instance
of the disadvantage of the artificial scheme of classification. The
species named N. leporinus has been shown to feed upon fish.

1 Dobson, Proc. Zool. Soc. 1875, p. 370. 2 IMd. p. 381.

VAMPYRUS 531

Furia, Amorphochttus, Rhynchonycteris, Saccoptei'yx, Cormura,
and Diclidurus are other Neotropical genera of the same family.

The genus T'aphozous1 has a tail which perforates the inter-
femoral membrane, appearing on its upper surface ; it is capable
of being withdrawn. The premaxillaries are cartilaginous. The
dentition is I -^ C \ Pm % M f . The upper incisors often disappear.
Many species of the genus have a gular sac, opening anteriorly
between the jaws. This is better developed in the males. The
genus ranges from Africa through Asia to New Guinea and
Australia. There are some twelve species.

The genus Molossus 2 has short legs and well- developed fibulae.
The tail is thick and fleshy, and is prolonged far beyond the margin
of the interfemoral membrane. The ears are united together
above the nose ; the tragus is minute. The dentition is I ^ or ^ C ^
Pm * or § M |. This genus, which is confined to the tropical and
subtropical portions of America, has long and narrow wings. The
Bats can thus fly rapidly, twist about with ease, and capture
strongly-flying insects. There are a large number of species.

Nyctinomus is an allied genus, and also has many species.
These range through both hemispheres. The chief differences from
Molossus are that the premaxillary bones are separate in front or
united by cartilage, and that the incisors may be three in the
lower jaw.

Fam. 5. Phyllostomatidae. — The Bats of this family are ex-
tremely numerous and almost entirely confined to South America.
None of them occur outside the New World. There are some
thirty-five genera. The members of the family are to be distin-
guished by the presence of the nose leaf, by the well-developed
premaxillae, and by the possession of three phalanges by the
middle finger. They are large, and the tragus of the ear is well
developed.

Vampyrus of South America contains the large species V. spec-
trum, which, mainly on account as it seems of its " forbidding
aspect," was supposed to be a bloodsucker. This genus has two
incisors on each side of the upper jaw.

The genus Glossophaga represents another type of structure in
this family. The tongue is long and extensile, and is much attenu-
ated towards the tip, where it is covered with strong and recurved
papillae. This structure was at one time thought to indicate a.

1 Uobson, Proc. Zool. Soc. 1875, p. 546. " IMd- 1876, p. 701.

532 STOMACH OF VAMPIRE CHAP, xvi

bloodsucking habit ; but its use appears to be merely that of
scooping out the soft insides of fruits, upon which the Bat mainly
lives. The incisors are only one on each side of the upper
jaw. The really bloodsucking Bats of this family belong to the
genera Desmodus and Diphylla. The former is the Vampire, the
species being known as Desmodus rufus. These Bats have no
tail ; there is no true molar tooth ; the canines are large, and the
single pair of upper incisors quite caniniform, and very sharp and
strong. These are the main teeth for aggression. In accordance
with its diet of blood, the Vampire has a peculiarly modified
intestine. The gullet is provided with a bore so small that
nothing but fluid food could pass down it ; the stomach is
intestiniform in shape.

CHAPTER XVII

PRIMATES

Order XIII. PRIMATES.

THE highest of mammals, the Primates,1 may be thus differ-
entiated from other groups : — Completely hairy, generally arboreal
mammals, with five digits on fore- and hind-limbs, provided with
flat nails (except in the case of certain Lemurs and the Marmosets),
the phalanges that bear these being flattened at the extremity and
expanded rather than diminished in size. The fore-feet are grasp-
ing hands as a rule, and the hind-feet walking as well as (generally)
grasping organs, and the mode of progression is plantigrade. The
teats, except in Chiromys, are thoracic, and even axillary in
position. The skull is characterised by the fact that the orbital
and the temporal vacuities are, at least partly, separated by bone.
The clavicles are always present. The carpus has separate lunar
and scaphoid bones, and the centrale is often present. There is
rarely an entepicondylar foramen in the humerus, except in some
archaic Lemurs. The femur has no third trochanter. The stomach
is usually simple, being sacculated only in Semnopithecinae. The
caecum is always present, and often large.

This great group could be easily divided into two separate
orders, the Apes and the Lemurs, if it were not for certain fossil
types. As will be seen from the description of Nesopithecus and
of Tarsius, the actual hard and fast lines between all Apes and all
Lemurs are very few. On the other hand, it is a little difficult to
draw a hard and fast line between the Primates as a whole — or
at least between the Lemurine section — and the Creodonta, a

1 For a general account of the Primates, see Forbes in Allen's Naturalists'
Library, London, 1894.

534 CHARACTERS OF PRIMATES CHAP.

group to which so many others appear to converge. It is dis-
puted, for example, whether the Chriacidae among extinct Lemurs
are rightly placed, or whether they should be referred to the
Creodonta. The number of primitive characters seen among the
Primates, even in Man himself, is remarkable. Of these the more
important are the five digits of both limbs and the plantigrade
walk, the presence of clavicles and of a centrale, and the absence
of a third trochanter. All these features distinguish the early
Eutheria.

SUB-ORDER 1. LEMUKOIDEA.1

The animals known as Lemurs, from their nocturnal and ghostly
habits, are on a lower level of organisation than the other division
of the Primates. Even the external form enables the members
of the present sub-order to be readily distinguished from the
higher Anthropoidea. The head is more like that of a Fox, with
a sharp muzzle ; it lacks the human expression of the face of even
the lower among the Apes. The long tail is never prehensile, and
there is never any trace of cheek pouches or of integumental
callosities, which are frequently so characteristic of the Apes.
The Lemurs agree with the remainder of the Primates in having
pectoral mammae (sometimes abdominal ones are present in
addition, and in Hapalemur — in the male at least — there is a
mamma upon each shoulder), in having opposable thumbs and
toes, and in the flattened digits. The tail varies from complete
absence (in the Loris) to a great length and bushiness in
the Aye -aye. The pectoral limbs are always shorter than
the hind -limbs; the reverse is occasionally the case in the
Anthropoidea. A curious contrast between the two divisions
of the Primates concerns the digits of the hands and feet. In
the Anthropoidea it is the hallux or pollex which is subject to
great variation. In the Lemurs, on the contrary, the thumb
and great toe are always well developed, but the second or
the third digit constantly shows some abnormality; thus the
singular elongation of the third digit of the hand in CMromys
and the absence of the index in the Potto.2 In all Lemurs the

1 See Dr. Mivart's papers in Proc. Zool. Soc. 1864, -65, -66, -67, and -73 foi
osteology and teeth.

2 Murie and Mivart, Trans. Zool. Soc. vii. 1869, p. 1.

xvii CHARACTERS OF LEMURS 535

second toe is furnished with a sharp nail, unlike the flattened
nails of the other fingers and toes, and in Tarshis the third also
is thus provided. As to osteology, the shape of the head, already
referred to, indicates some of the differences in the skull which
mark off the Lemurs from the Anthropoidea. The brain case is
small relatively to the face : the orbital and temporal fossae are
in communication, though the frontal and jugal bones are united
behind the orbit. The two halves of the lower jaw are not in-
variably ossified to form one piece, as is the case with most Apes.
The lachrymal foramen lies upon the face in front of the orbit.
The teeth are characteristic, not so much in their number (the
dental formula is usually 12, C I, Pm 3, M 3 = 36) as in the
disposition of the incisors. The incisors of the lower jaw and the
canines project forwards in a way only found in a few American
Monkeys ; as in the Apes there are four incisors in each jaw, but,
with the exception of the highly aberrant Chiromys, there is a
space in the upper jaw between the incisors of the two sides.
The canines of the lower jaw, moreover, are often incisiform.
There is a well- developed sublingua beneath the tongue (see p. 61).
The stomach is perfectly simple ; and the caecum, always present
and varying in length, never has a vermiform appendix. The
gall-bladder is always present. The brain differs from that of
the Anthropoidea in that the cerebellum is, as in the lower
Mammalia, exposed. The convolutions upon the cerebral hemi-
spheres are not greatly developed, a circumstance, however, which
(see p. 77) may have more relation to the size of the animals
than to their mental development. Though the brain in its
general outlines is not like that of the other Primates, there are
certain resemblances ; the most striking of these is perhaps the
presence, though in rather a rudimentary condition, of the " Simian
fissure."

The Lemurine brain has been chiefly studied .by Flower,1
by Milne-Edwards,2 and by myself.3 There are also a number
of scattered papers dealing with particular types, such as the
memoirs of Owen4 and Oudemans,5 upon the brain (and the
general anatomy) of Chiromys. Without going into great

1 Trans. Zool. Soc. v. 1863, p. 103.
2 Hist. Nat. de Madagascar, Mamm. 1875.

3 Proc. Zool. Soc. 1895, p. 142.

4 Trans. Zool. Soc. v. 1863, p. 33.

5 Verli. Ak. Amsterdam, xxvii. 1890, Art. '2.

536 ARM GLAND OF HAPALEMUR CHAP.

detail it may be stated generally that the anatomy of the brain of
this group confirms the classification which is adopted in this
work.

A curious feature in the anatomy of the Lemurs, which they
share with animals so remote from them in the system as the
Edentata, is the breaking up of some of the arteries of the limbs
to form retia mirabilia ; nothing of the kind is known among
the other Primates.

Perhaps the most remarkable difference between the Lemurs
and the Anthropoidea, which are really in many respects more
closely allied than might be inferred from the above summary of
differences, is in the structure of the placenta. The Lemurs agree
with the Ungulates in having a non-deciduate placenta.

A curious feature confined to the sub-family Lemurinae was
first discovered by myself in Hapalemur griseus.1 On the
forearm (see Fig. 258) is an area of hardened skin, which is
raised into spine-like processes. Fully developed, this organ is
characteristic of the male, the area being marked off in the
female, but without the spiny outgrowths. On removing the
skin a gland about the size and shape of an almond is brought
into view. In other Lemurs there is no modified skin, but a
small tuft of particularly long hairs, which are also present in
Hapalemur, and a small gland beneath the skin. The gland of
Hapalemur may be comparable with a tract of hardened skin in
Lemur catta, which projects to a large extent and has been spoken
of as a " climbing organ."

An almost exactly similar tuft of spine-like outgrowths exists
upon the lower end of the ankle of Galago garnetti. The spines
are black and bent, just as they are in Hapalemur. There
appears also to be a gland. This structure is not universal in
the genus Galago any more than is the patch of spines in the
genus Hapalemur.

In addition to this gland and to the patch of spines which
cover it, the same Lemur as well as Chirogahus and certain
species of Lemur possess to the inner side of it a bundle of long
and stiff bristles associated with unusually large sebaceous
glands ; these structures are, of course, not homologous with the
gland of the arm of Hapalemur, as they coexist in the same

"On some Points in the Structure of Hapalemur griseus " Proc. Zool. Soc.
1884, p. 301.

CARPAL VIBRISSAE

537

species. They are, moreover, not peculiar to the Lemurs, but
exist in the Squirrel, in the Domestic Cat, in the Leopard, in
Bassaricyon? the Otter, various Marsupials, and doubtless in

FIG. 258.— A, left arm of Hapalemur griseus $ . a, Teat ; b, spines on arm gland ; c,
tactile bristle. B, left foot of Xycticebus tardigradus. 1 to 5, Pads upon sole of
foot. (After Sutton, and Mivart and Murie.2)

many mammals which require a tactile organ, for these hairs are
associated with a large branch of the radial nerve.

The Lemurs have at the present time a most remarkable dis-
tribution. There are altogether about fifty species, referable to
seventeen genera. Thirty-six species are confined to Madagascar

1 Beddard, Proc. Zool. Soc. 1900, p. 661.
2 On the Arm Glands of the Lemurs, Proc. Zool. Soc. 1887, p. 369.

538 THE INDRI

and to some small neighbouring islands. The rest occur in the
Ethiopian and in the Oriental region. The rest of the world is
at present totally without Lemurs, though, as will be seen in the
sequel, the order was more widely spread over the globe in past
times.

Fam. 1. Lemuridae. — This family can be usefully subdivided
into four sub-families.

Sub-Fam. 1. Indrisinae. — This sub-family is limited to Mada-
gascar, and has been exhaustively treated of by M. Grandidier
and Professor Milne-Edwards in the Histoire de Madagascar.
These Lemurs contrast with others by the large size of the hind-
as compared with the fore-limbs. The ears are short. The tail
varies in length. The thumb is but slightly opposable, and the
toes are webbed. Correlated with the first two of these
characters, these Lemurs when upon the ground progress by
means of the hind-limbs, holding their arms above their heads.
The number of teeth is reduced, the total being thirty. The
formula l is I f C -J- Pm f M •§-. The colon or large intestine, as
figured by Milne-Edwards, has a remarkable watch-spring-like coil,
highly suggestive of the Euminants and of certain Kodents. This,
however, is only in Propithecus and Avahis. The caecum in
this sub-family is specially large. The brain is characterised by
the comparatively slight development of the angular fissure in
Propithecus and Avahis ; it is in them anterior in position. In
Indris it is more S-shaped and larger as in Lemur. The
parieto-occipital fissure is fairly well developed, so too is the
antero-ternporal.

The genus Indris has more pronounced external ears than
have the two other genera of the sub-family. The tail is
rudimentary. The incisors of the upper jaw are sub-equal and
set close together, those of the lower jaw have marked longitudinal
ridges upon the outer surface, which suggests Galeopithecus (see
p. 520). The molars are quadricuspidate. There is but a single
species, I. brevicaudata, which is of a black colour, diversified with
white upon the rump and the limbs. The term " Indri " 2 means,
as does "Aye-aye," "look." One of the native names for the

1 So at least the formula has been given ; but it is very possible that the
supposed second incisor is really, judging from the other Lemurs, a canine.

The Malagasy, however, must be vague in definition, or their interpreters
not well grounded in the rudiments of the language ; for Sonnerat states that
Indri signifies " homme des bois."

THE SIFAKAS 5 39

animal, " Amboanala," signifies " dog of the forest/' and is derived
not only from the woeful howls of the creature, but from the fact
that in certain parts of the island it is used as a dog to chase
birds.

These howls are largely effected by means of a laryngeal pouch,
which is described as different from that of Apes ; the mechanism
must also differ from that of Megaladapis, inasmuch as the lower
jaw is not deep as in that extinct Lemur. The Indri is the
largest of Lemurs, measuring about two feet in length. It is
arboreal and social, travelling in large companies. As is the
case with the Propithecus, the natives of Madagascar hold the
Indri in awe and veneration. It is curious that the name
Lemur or ghost is peculiarly applicable to the Indri or Babakote
in another sense from that which led to its adoption by Linnaeus.
The natives, in fact, believe that men after death become Indris.
Naturally, therefore, these Lemurs have reaped the advantage of
this superstition in almost perfect immunity from destruction.
Their "long-drawn-out, melancholy cries" are probably at the
root of much of the ghostly terrors which they inspire.

The genus Avahis 1 has but a single species, A. laniger, which
is the smallest of this sub-family. It is a foot long without the
tail. The Avahi has a long tail (15 inches in length) like
Propithecus. The outer incisors are larger than the inner, thus
differentiating the genus from Propithecus. The molars of the
upper jaw are quadricuspidate, of the lower jaw five cusped. This
genus has only eleven pairs of ribs instead of the twelve of
Indris and Propithecus. The Avahis, unlike the Sifakas and
Indrinas, lead a solitary life, or go about in pairs. They are,
moreover, completely nocturnal.

The genus Propithecus is characterised by the fur being rather
silky than woolly, which latter is the kind of fur found in the
two other genera of the sub-family. They are also rather
larger animals, the body reaching a length of nearly 2 feet.
The tail is long as in Arahis ; the inner incisors are larger than
the outer. The " Sifakas," as these Lemurs are termed, have a
reputation for gentleness of character, but, as is the case with
other animals, the males fight for the possession of the femal
the breeding season. They are mainly vegetarian in habit, and
travel in large companies. There are at least three species, and

* J Svn. Micrurhijnchus.

540

THE TRUE LEMURS

several varieties are allowed. The colours of these Lemurs are
bright, and distributed so as to form contrasting bands ; thus
P. coquereli, a variety of P. verreauxi, has a black face and a
body mainly white, with splashes of a rich maroon upon the
limbs and upon the chest.

These Lemurs are diurnal, and are especially active in the
early morning and evening, sleeping, or at any rate remaining
quiet, during the heat of the day. Their fitness for an arboreal
life is shown by the existence of a parachute-like fold of skin
between the arms and the body, which suggests a commencement
of the more complete parachute of Flying Foxes, etc. These
Lemurs are said to be reverenced and therefore shielded from
injury by the natives of Madagascar.

Sub-Fam. 2. Lemurinae. — The "True Lemurs"are all inhabit-
ants of Madagascar and of the Comoro Islands. They have not such
long hind-limbs as have the members of the last sub-family, nor
are the toes webbed. The tooth formula differs from that of the
Indrisinae in that there is one more premolar on each side of
the upper jaw, and often one more incisor in the lower jaw,
making thus a total of thirty-six teeth. Sometimes, however,
the incisors of the upper jaw are totally wanting.

The Hattock, genus Mixocebus, is a scarce creature, only known
from a single species, M. caniceps. As it is rare, nothing is
known of its habits. It has one pair of upper incisors. The
creature is one foot and half an inch long, exclusive of the tail,
which is an inch longer than the body.

Genus Lepilemur.- — -The Lemurs belonging to this genus,
entirely confined to Madagascar, as are all the Lemurinae, have
received the perfectly unnecessary and pseudo-vernacular name of
" Sportive Lemurs " ; an equally inappropriate and" not at all
ingenious name of "Gentle Lemurs" being bestowed Upon the
allied genus Hapalemur. In Lepilemur there are seven species,
which are to be distinguished from Mixocebus in having the tail
shorter than the body. There are no incisors in the upper jaw.
The last molar is tricuspidate in the upper jaw ; that of the lower
jaw has five cusps. They are nocturnal creatures, and but little
is known of their habits. Previously to Dr. Forsyth Major's visit
to Madagascar only two species of the genus were known ; he has
added five others. The length of the body is 14 inches, and that of
the tail 10 inches, in L. mustelinus, which is the Jargest species.

xvn HA PA LEMUR 5 4 1

The genus Hapaletnur l has a shorter muzzle than Lemur, and
shorter ears. There are two pairs of mammae instead of only
one ; these are upon the breast and abdomen. In the male there
is a pair upon the shoulder. The incisors are small, sub-equal,
and placed one behind the other ; the last one is at the inside of
the canines. The molars of the upper jaw and the last premolar
have only three well-marked cusps ; in the lower jaw they have
four. The caecum is blunter and is not so long as in Lemur ; it
differs from that of other Lemurinae in having only two support-
ing mesenteries, which are both furnished with blood-vessels.' As
in Lepilemur and the Indrisinae the carpus has no os centrale.

The genus, which is confined to the island of Madagascar, has
two species, of which one, H. sim,us, is the larger and has a broader
muzzle, and does not possess the peculiar arm gland (Fig. 258)
already described in H. griseus. The former species is stated by Mr.
Shaw to be chiefly a grass-eater, and to dislike berries and fruits,
which are usually so popular with Lemurs. It is, however, believed
by some that there is but one species of Hapalemur. H. griseus is
1 5 inches long, and has a tail of the same length. Its native name
is " Bokombouli." It is nocturnal, and is especially addicted to
bamboos, upon the shoots of which it feeds and among which it
lives. It is often exhibited in the Zoological Society's Gardens ;
but the specimens seem to be always males. This Lemur is of a
dark iron-grey colour with a tinge of yellow, more marked in
individuals which have received the separate specific name of
H. 'olivaceus.

The genus Lemur is distinguished by the long tail, half as long
as the body at the least, by the elongated face, and by the Fox-
like muzzle ; the teeth are present to the full number of the family,
viz. thirty-six ; the incisors are small and equal in size, and are
separated from each other and from the canines by spaces. The
molars of the upper jaw have five cusps, but there are only four
in the lower jaw.

This genus is entirely confined to Madagascar and the Comoro
Islands, and consists of several species, the exact number of which
is doubtful. Wallace in his Geographical Distribution allows
fifteen ; Dr. Forbes only eight, with a plentiful allowance of
varieties. One of the best-known species is Lemur catta, the

1 Beddard, Proc. Zool. Soc. 1884, p. 391, and 1891, p. 449 ; and Jentink, Notes
Leyd. Mus. 1885, p. 33.

542

SPECIES OF THE GENUS LEMUR

King-tailed Lemur, or the " Madagascar Cat " of sailors. Lemur
macaco shows a remarkable sexual dimorphism, the male being
black, and the female— formerly described as a distinct species,
L. leucomystax — being reddish brown with white whiskers and
ear tufts. This led to a confusion with a totally distinct species,
L. rufipes, of which the male (regarded as distinct and called
L. nigerrimus} is entirely black. This latter identification is,
however, considered by Dr. Forsyth Major1 to be not quite
certain at present.

The young Lemur is at least sometimes carried by the mother
across her belly ; its tail passes round her back and then round
its own neck.

The Lemurs of this genus agree with those of some other
o-enera in the loudness of their voice, which is constantly

r*

FIG. 259. — Ruffed Lemur. Lemur varius. x I.

exercised. Some move about by day and others by night. They
are insectivorous and carnivorous as well as vegetarian ; and Mr.
Lydekker suggests 2 that their abundance and hardiness is to be
traced to this fondness for a mixed diet. Lemur catta seems to
be the only member of the genus that is not arboreal. It lives
among rocks where but few trees, and those much stunted, occur.
Many species of Lemur are always to be seen in the Zoological
Society's Gardens. Fourteen " species " have at one time or
another been exhibited.

Sub-Fam. 3. Galagininae. — This sub-family is found on the
continent of Africa as well as in Madagascar ; but the genera are

1 Proc. Zool. Soc. 1899, p. 554.
- Royal Xatural History, London, 1894, p. 211.

xvn GALAGO AND CH1ROGALE 543

different in the two districts. In Madagascar we have Opolemur,
Microcebus, and Chirogale ; on the continent, Galago. The
members of this sub-family have markedly large ears, which are
but little furry; the tail is long. A very marked skeletal
character distinguishes this sub-family from other Lemuridae,
and allies them to Tarsius, that is the lengthening of the
calcaneum and naviculare in the ankle. The dental formula is
as in Lemur. The supporting bands of the caecum are in this
sub-family as in the genus Lemur. There are but two folds, of
which one is median and non-vascular; the lateral fold bears a
blood-vessel, and is joined by the median frenum. The brain is
but little known. The only figure of the brain of Galago is one
by myself. There are four mammae, two on the breast and two
upon the abdomen.

The genus Galago comprises at any rate six distinct species.
They are all African, and range right across the continent from
Abyssinia as far south as Natal, and to Senegambia in the west.
The incisors of the upper jaw are small and equal ; there is a
gap between the canine and the first premolar. The molars and
the last premolar have four cusps ; the last molar of the lower
jaw has an additional fifth cusp as in Macacus, etc. The Galagos
are chiefly nocturnal, and are more or less omnivorous. Owing
to their long hind-legs these animals when they leave the trees
advance upon the ground by hops like a Kangaroo. Galago
senegalensis makes a nest in the fork of two branches, where it
sleeps during the day. The Great Galago (G. crassicaudatus) is
named by the Portuguese " Eat of the Cocoa-nut Palm." Sir
John Kirk, after whom a variety of this species is called, relates
that it is incapable of resisting the fascinations of palm wine,
upon which it will readily intoxicate itself, and as a consequence
brave probable captivity. I have referred above (p. 536) to the
patch of spines upon the tarsus of G. garnetti.

The genus Chirogale is entirely confined to Madagascar. It is
to be distinguished from Galago by the fact that the inner incisors
are larger than the outer. There are five species of the genus
known : four previously to Dr. Forsyth Major's recent visit to
Madagascar, and a fifth brought back by him.1 In connexion
with this genus the naturalist just mentioned has observed
that all the Lemurs of Madagascar, including the aberrant

1 See JYovitates Zoologicac, vol. i. 1894, p. 2.

544

MINUTE LEMURS

Chiromys, differ from the African forms by the fact that the
tympanic ring " is completely enclosed by the bulla ossea, but
without osseous connexion with the same." This character he

FIG. 260. — Smith's Dwarf Lemur. Microcebus smithii. x f.

thinks so important as to justify the inclusion of all the
Mascarene forms in one group as opposed to another group
consisting of the continental Lemurs. In this event Chirogale

FIG. 261. — Mouse Lemur. Chirogale coquereli. x ^.

will have to be separated from its close association with Galago.
For the present, however, it is left in the more generally accepted
position.

Microcebus contains the most minute among the Lemurs. M.
smithii has a body only 5 inches long, the tail being another
6 inches. It occurs in Madagascar, and includes five species.

Opolemur, the Fat-tailed Lemur, was so called on account of a

THE SLOW LORIS 545

deposit of fat formed chiefly at the root of the tail, and intended
to tide over the time of the creature's hibernation. But, as a
matter of fact, this peculiarity also exists in Cliirogcde. Of
Opolemur but two species are known, and of one of these, named
after Mr. Thomas of the British Museum, only three examples are
in existence in museums, that is to say in one museum — our own
at South Kensington. Many of these dwarf Lemurs are exceed-
ingly rare. In this genus and in the last two the palate has a pair
of posterior fenestrae, of which there are also traces in other Lemurs,
but which are particularly large in Microcebus. This is, of course,
a well-known character of the Marsupials, and also, which is more
important in the present connexion, of certain Insectivores.

Sub-Fam. 4. Lorisinae. — This sub-family is the only one with
a wide distribution, and it contains, with the exception of Tarsius,
the only Asiatic members of the group. Correlated with its
wide distribution there is more divergence in anatomical characters
than is the case with the other sub-families of the Lemuridae.
In external features all the three genera of this sub-family agree
in their small size, their short or entirely deficient tail, large
staring eyes, and the rudimentary character, or absence, of the
index finger, which is never provided with a nail ; in all of them
the thumb diverges widely from the other fingers, and the great
toe is so divergent as to be directed backwards. In the brain
there is one character common to all three genera, and that is
the small length of the angular fissure. The caecum, which is
long, is supported by three folds, of which the median is anangious,
and is sometimes attached to the longer of the two lateral folds,
which are vascular. The members of this sub-family have more
dorsal vertebrae than are found in other Lemurs; the range is
from fourteen in Loris, to sixteen in Nycticebus.

The genus Nycticebus contains only a single species, N. tardi-
gradus, though four other names have been given to supposed
varieties. Moreover, the genus itself has been named Stenops, as
also the next genus Loris. The body of this animal is stouter than
that of the next to be described. Professor Mivart has pointed
out that, though Asiatic like the Loris, it presents more resem-
blances to the African Potto. The index finger is small ; the
inner of the two incisors is smaller than the outer, but both of
one side are close together. They may be reduced to one on each
side of the upper jaw.

VOL. x 2 N

546

LEGENDS CONNECTED WITH THE LORIS

The animal has a wide distribution in the East, occurring in
Assam and Burmah, the Malay Peninsula, Siam, and Cochin-Chiiia,
Sumatra, Java, Borneo, and the Philippines. Its vernacular
names signify " Bashful Cat " and " Bashful Monkey " in allusion
to its nocturnal and shy habits. It lives among trees, which it
does not voluntarily leave. Its movements are deliberate, as its
popular name, Slow Loris, implies ; but it makes up for this by
a vigorous tenacity of grasp. The animals " make a curious

FIG. 262. — Slow Loris. Nycticebus tardigradus. x J.

chattering when angry, and when pleased at night they utter a
short though tuneful whistle of one unvaried note, which is
thought by Chinese sailors to presage wind." Much superstition
has collected round this harmless though rather weird-looking
creature. Its influence over human beings is as active when it
is dead as when it is alive. " Thus," writes Mr. Stanley Flower,1
" a Malay may commit a crime he did not premeditate, and then
find that an enemy had buried a particular part of a loris under
his threshold, which had, unknown to him, compelled him to act
to his own disadvantage." The life of the Loris, adds Mr. Flower,

1 Proc. Zool. Soc. 1900, p. 321.

xvii THE POTTO AND ANGWANTIBO 547

" is not a happy one, for it is continually seeing ghosts; and that
is why it hides its face in its hands ! "

The genus Perodicticus contains two quite recognisable species,
known respectively as the Angwantibo and Bosnian's Potto. The
former has been regarded as referable to a distinct genus, Arcto-
cebus. A curious internal character of the Potto which is visible,
or at least can be felt, externally, is the long neural processes of
the cervical vertebrae, which project beyond the level of the skin.
The index finger is rudimentary and so is the tail, being only
just visible (about an inch in length) in the Potto. The colour
of both genera is a reddish grey, redder in the Potto. The
incisors are equal and minute. Both species are confined in
their range to West Africa, and are arboreal like the other
members of the sub-family. The Potto seems to share the
leisurely mode of progression of its Asiatic relatives, if Bosnian,
its original describer, is to be trusted. He says : " By the negroes
called Potto, but known to us by the name of Sluggard, doubtless
from its lazy, sluggish nature ; a whole day being little enough
for it to advance ten steps forward." The same writer did not
at all appreciate his addition to zoological knowledge, for he
remarked that the Potto " hath nothing very particular but his
odious ugliness." The Angwantibo is rare and but little known.
Our knowledge of its anatomy is derived from a paper by
Huxley.1 It is an animal measuring about 10^- inches in total
length to the end of the tail, which is only a quarter of an inch
long. The hands and feet are smaller than those of Perodicticus.
The index finger is rudimentary and has but two phalanges, and
it has no trace of a nail. In this it agrees with the Potto, but
" the spinous processes of the cervical vertebrae do not project in
the manner described by van der Hoeveu in the Potto, though
they can be readily felt through the skin." The dental formula
of this genus as of the last is I f C ^ Pm f M if. The last lower
molar has a fifth cusp, which is wanting in the Potto. The last
upper molar is tricuspid. It is bicuspid in the Potto. It seems
impossible to avoid agreeing with Professor Huxley that the
Angwantibo is entitled to generic separation.

The genus ZoHs.also contains but a single species, L. gracilis,
and is, as its name denotes, an animal of more slender build than
the Slow Loris. Its eyes are very large, and the limbs excessively
1 "On the Angwantibo," Proc. Zool. SQC. 1864, p. 314.

548

THE AYE-AYE

CHAP.

slender. ^The index finger is much as in Nycticebus. The colour,
too, is not widely different, being of a yellowish grey, but it lacks
the dorsal stripe which distinguishes its relative. The incisor
teeth are equal and very small. The last upper molar has four
cusps instead of the three of Nycticebus. This Lemur is confined
to Southern India and Ceylon, and has much the same habits as
the last. But it is rather more active, and can capture small
birds when sleeping upon the trees ; its diet, however, is mixed,
and is vegetarian as well as animal.

A mysterious Lemur, which we conveniently place as a kind
of appendix to the present family on account of its locality, has
been shortly described by Nachtrieb from the Philippines. The
tail is rudimentary ; there are two upper incisors, but as many as
six lower. It is doubtful what the beast really is.

Fam. 2. Chiromyidae. — This family contains but a single
genus and species, the Aye-aye, Cliiromys madagascariensis,

FIG. 263. — Aye-aye. Chiromys madagascariensis. x ^,.

whose characters therefore are for the present those of the
family as well as of the genus and species. The external
features of this extraordinary animal will be gathered from
an inspection of Fig. 263, from which it will be seen that
the earlier name of Sciurus given to the creature was not by
any means a misnomer. The Squirrel-like appearance is due,
of course, chiefly to the strong and long incisor teeth. As
to the external characters, which are of systematic importance,

xvn ANATOMY OF THE AYE-AYE 549

attention may be called to the long and bushy tail, to the greater
length of the hind-limbs, to the abdominal teats (one pair) in the
female, and above all to the singular third digit of the hand,
which is thin and elongated. The thumb is, as in other Lemurs,
opposable, and has a flat nail ; the remaining digits have claws,
as have also the toes with the exception of the great toe, which
has a flat nail like the thumb.

The anatomy of this animal has occupied the attention of a
considerable number of observers, dating from Sir E. Owen, who
was the first to give a connected account of its entire organisa-
tion. The most recent paper of importance is by Dr. Oudemans.1
The teeth are very unlike those of other Lemurs. The most
remarkable divergence is in the incisors, which are present to
the number of but a single pair in each jaw, and are shaped
like those of the Eodentia, and in the same way as in that
group grow from persistent pulps. There are likewise, as in
the Eodents, no 'canines. There are two premolars in the
upper jaw (none in the lower) and altogether twelve molars, so
that there is a total of eighteen teeth. The intestine has a
moderately long caecum. The brain has been most fully described
by Oudemans, who had fresh material to work with, the brain
described by Owen having been extracted from a spirit-preserved
carcase. The angular fissure is well developed, as in Lemur and
the Indri ; but it does not join the infero-frontal. The antero-
temporal fissure is also well developed.

" The name of Aye-aye," wrote Sonnerat, the discoverer of the
animal, " which I have retained for it, is a cry of surprise of the
inhabitants of Madagascar." It is, however, usually said that
the animal itself makes a sound which may be written in the
same way (or with an initial H). It is an arboreal and
nocturnal animal, which accounts for its excessive rarity at one
time. In one of his many eloquent essays upon natural history
the late Mr. P. H. Gosse adduced the Aye-aye as an example of
a creature on the verge of extinction. It is, however, now more
frequently met with, though the superstition of the natives
renders its capture a matter of some difficulty. There is a
specimen at the moment of writing in the Zoological Society's
Gardens. There has been some discussion as to the use of the
slender middle finger : it is stated that it can thrust it into the
1 Verh, Ak. Amsterdam, xxvii. 1890.

55O AFFINITIES OF TARSIUS CHAP.

borings of the larva of a certain beetle of which the Lemur is
particularly fond, and can extract the insect, or at any rate
discover its position, when it may be extracted by the powerful
chisel-shaped teeth. The partiality of the Aye-aye for animal
food of any kind including insects has been both reaffirmed and
denied ; and Mr. Bartlett has seen the creature use its slender
finger for combing out its hair, and for other purposes of the
" toilet." Dr. Oudemans has figured in his paper an apple which
has been largely eaten by the Chiromys ; the fleshy pulp has
been entirely excavated, leaving only the core and the skin, which
are untouched. The Eev. Mr. Baron is one of the latest writers
upon the ways of life of Chiromys}- He states that it inhabits
the most dense parts of the forests. It has the habit of prowling
about in pairs, and the female produces but a single young one
at a birth. A nest, which is about 2 feet across, is made of twigs
in lofty branches. This is occupied during the day, and entered
by a hole in the side. With regard to the superstitious venera-
tion in which the animal is held, it is said that if a person sleeps
in the forest the Aye-aye will bring him a pillow. " If a pillow
for the head, the person will become rich ; if for the feet, he will
shortly succumb to the creature's fatal power, or at least will
become bewitched." But a counter-charm may be obtained. It
is said that the reverence for this beast leads the natives to bury
carefully a specimen found dead.

Fam. 3. Tarsiidae. — This family also consists of but a single
genus, Tarsius, to which it is the general opinion that but a
single species belongs ; there are, however, at least four different
specific names on record. The general aspect of the animal is
not unlike that of a Galago, with which it also agrees in the
elongation of the ankle ; but the elongation is more pronounced
in the present genus. The ears are large, and the eyes are
extraordinarily developed. The fingers and toes terminate in
large expanded discs, and are furnished with flattened nails
except on the second and third toes, whicli have claws. The tail
is longer than the body and is tufted at the end. The skull is
more like that of the Anthropoidea than is the skull of any
other Lemur. The resemblance is by reason of the almost com-
plete separation of the orbit and the temporal fossa by bone ;

1 Proc. Zool. Soc. 1882, p. 639 ; see also Rev. G. A. Shaw, Proc. Zool. Soc. 1883,
p. 44, 2nd Art.

HABITS AND DISTRIBUTION OF TARSIUS

551

there is, however, a gap left to mark the Lemurine characters of
the animal. The placenta, too, has been compared to that of the
Apes. The dental formula is that of the genus Lemur, save for
the absence of an incisor on each side of the lower jaw ; the
number of teeth is therefore thirty-four. The incisors of the
lower jaw are upright, and not procumbent as in other Lemurs.
The caecum is of moderate length. The brain is almost smooth,
but there is a Sylvian fissure and an antero-temporal, which latter
does not reach the lower margin of
the brain, but divides the middle part
of the temporal lobe. The name
Tarsier, as may be inferred, was
originally given to this creature by
Buffon on account of the abnormal
ankle, and it was compared by him
with the Jerboa, like which animal
the Tarsier leaps when it descends to
the ground. The genus is Malayan,
but its range extends to the Philip-
pines and to Celebes and Borneo.
The Tarsiers are nocturnal and parti-
cularly arboreal ; they live in pairs,
in holes in tree stems, and are mainly
insectivorous in their food. One,
rarely two young are produced at a FIG. 264.— Right pes of Tarrius
birth. Contrary to what is found in
many Lemurs, the Tarsier is a silent
creature, and at most emits a " sharp,
shrill call." Dr. Charles Hose, who
has studied this creature, has noticed
that the mother often carries her young one about in her mouth
like a Cat. Like so many Lemurs this animal is held in super-
stitious dread, which no doubt is the result of its most weird
appearance.1

Fossil Lemurs. — The Lemuroids are a very ancient race;
they extend back to the very earliest strata of the Eocene, the
Torrejon and Puerco beds, which, as already said, are thought to
be more referable to the Cretaceous than to the 'Tertiary epoch.

1 For a survey of the position of Tarsius, see Earle, Amer. Naturalist, xxxi.
1897, p. 569 ; and Nat. Science, x. 1897, p. 309.

cuneiform ; <?, middle cunei-
form ; c3. external cuneiform ;
cb, cuboid: w, uavicular : 7-1',
the digits. (From Flower's
Osteology.)

552 ORIGIN OF LEMURS

One of these early forms is referred to the genus Mixodectes, a
genus which has been placed, though with a query, in the order
Kodentia. It appears, however, to be a Lemuroid, and is of
American range. The incisor teeth have been held to argue
that it lies on the direct track of Chiromys ; but other features,
more especially the form of the astragalus, have been used to
argue the justice of the inclusion of thrs type within the order
Rodentia. Allied, as it is supposed, to this form is Indrodon,
also of the lowest Eocene deposits of the United States. Indrodon
mcdaris is known from fragments of nearly all parts of the
skeleton. They indicate the existence of a creature of about one-
half the size of Lemur varius. It had slender limbs and a
long and powerful tail. The humerus, as in so many archaic
beasts, has an entepicondylar foramen. The femur has three
trochanters, and the fibula articulates with the astragalus. It is
not always easy to distinguish these primitive mammals from
each other, so that the minutest of characters have to be called
in to our assistance. One of the contemporaneous groups with
which these early Lemurs might be confused is that of the
Condylarthra ; it is important, therefore, to note that in Indrodon
the calcaneo-cuboidal articulation is nearly flat, and not bent as
it is in the former group. The teeth are of the tritubercular
pattern. The incisors are not known, but the molars and pre-
molars are each three. To the same family, which has been
termed Anaptomorphidae, is referred the genus Anaptomorphus,
which has been specially compared to Tarsius. This small
animal has a Lemurine face with huge orbits. It has a pre-
molar less than Indrodon. It has been ascertained that A.
homunculus had an external lachrymal foramen.1

Another family, that of the Chriacidae, appear to hover on
the border line of Lemurs and Creodorits, having been referred to
both by various palaeontologists. Professor Scott suggests their
Lemurine or at least Primate relationships, while Cope urged
their Creodont affinities. A difficulty raised by Scott was, that
m Chriacus the premolars of the lower jaw were spaced. But it
appears that this is not fatal to their inclusion in the Primates,
since Tomithtrium, an "undoubted Primate," shows the same
feature. If G'kriatus is a Lemur it is an earlier type than those

1 See Schlosser, BeitragePal. Osterr. Hung. 1888 ; also Osborn and Earle, Bull.
Arner. Mus. Xat. ffist. vii. 1895, ]>. 16.

xvn NESOPITHECUS, AN INTERMEDIATE FORM 553

which have been considered ; for it has the typical Eutherian
dentition of four premolars and three molars. These teeth,
especially the superior molars, are particularly compared to the
corresponding teeth of Lemur and Galago. Of this and the
allied genus, Protochriacus, several species are known.

Adapis, a representative of another family, is one of the best
known of ancient Lemuroids. It has the typical mammalian
dentition of forty-four teeth in a close series without diastemata.
The orbits are completely separated from the temporal cavity,
the eyes looking forwards. The canines are large and caniniform.
The skull is deeply ridged behind with the usual sagittal crest.
This genus is European, and corresponds to the already mentioned
American Eocene Tomitherium, perhaps belonging to the same
family.

Nesopithecus is an extinct genus from Madagascar, lately de-
scribed by Dr. Forsyth Major.1 There are two species, N. roberti
and N. australis. The dental formula is I 2, C 1, Pm 3,.M 3, for
the upper jaw, the lower jaw having but a single pair of incisors.
The lachrymal foramen is just inside, or on the edge, of the orbit,
so that one distinctive Lemurine character is lost. The genus is
also Ape-like in the form of the canines and incisors, these
having been especially compared by Dr. Forsyth Major with
those of the Cercopithecidae. The molars, too, agree with those
of the same family. There is, however, one important feature in
which Nesopithecus resembles not only the Lemurs as opposed to
the Apes, but the Malagasy Lemurs. As already mentioned (p. 544),
Dr. Major has shown that in the Malagasy Lemurs, even including
the aberrant Chiromys, and in the Tertiary and European Adapis,
the bulla tympani is not produced by an ossified extension of the
annulus tympanicus, but from the adjacent periotic bone, the
annulus remaining separate and lying within the fully-formed
bulla. This feature shows conclusively that Adapis is a Lemur,
and that Nesopithecus, originally supposed to be a Monkey, cannot
be removed from the Lemuroidea, many though its likenesses to
the higher Primates undoubtedly are. However, this feature,
combined with the fact that the orbital and temporal cavities are
in communication, shows the Lemuroid position of Nesopithecus,
though it is quite conceivable that it is on the way to become
an Ape.

1 Proc. Zool. Soc. 1899, p. 987.

554 LARGE EXTINCT LEMUR CHAP.

A family, Megaladapididae, lias been quite lately founded
by Dr. Forsyth Major1 to include the remains of a gigantic
extinct Lemur from Madagascar, which when alive, so far as we
can judge from the skull, must have been three or four times the
size of the Common Gat. The name Megcdadapis madagascariensis
was given to the fossil, on account of certain resemblances to the
also extinct Adapis. It differs from other Lemurs in a number
of characters which jointly warrant its inclusion in a distinct
family. The small size of the orbits suggest a diurnal life ; the
deep mandibles, which, unlike what is found in other Lemurs,
are completely blended at the suture, point to the existence of a
howling apparatus, as in Mycetes. The low brain -case is a
character which is found in so many extinct Mammalia belonging
to many different orders that it weighs neither ono way nor
the other in considering the systematic position of the animal.
The shape of the molars, which are three in each half of each jaw
as in other Lemurs, is, according to the discoverer, like that of
the genus Lepilemur. The incisors and the canines are not
known. Of a still larger form, M. insignis, the molar teeth are
known*

SUB-ORDER 2. ANTHEOPOIDEA.

The Apes differ from the Lemurs in that the teats are
always restricted to the thoracic region ; the orbit, though sur-
rounded by bone as in the Lemurs (and in Tupaia, a very Lemur-
like Insectivore), does not open freely behind into the temporal
fossa as in Lemurs (except Tarsius). The lachrymal opening is
inside the orbit instead of outside ; the cerebral hemispheres
are more highly developed, and conceal, or nearly conceal, the
cerebellum ; the upper incisors are in close contact ; a few other
points are mentioned under the description of the characters of
the Lemurs. There are altogether about 212 species of Monkeys
and Apes. They are tropical and subtropical in range, and, with
but few exceptions, are impatient of cold.

The Monkeys are primarily divisible into two great divisions,
which have been termed, on account of the characters of the nose,

1 Phil. Trans, clxxxv. B, 1894, p. 15.

2 It seems to be possible that this great Lemur was extant so lately as 1658,
when a creature possibly answering to it was described by de Flacourt.

xvn CHARACTERS OF ANTHROPOIDEA 555

the Catarrhi nes and Platyrrhiries. In the former the nostrils look
downward and are close together ; in the latter they are separated
by a broad cartilaginous septum, and the apertures are directed
outwards. But numerous other points of difference separate these
two groups of the Monkey tribe. The Catarrhines often have
those remarkable ischial callosities, patches of hard skin brightly
coloured ; the tail may be totally wanting as a distinct organ, as
is the case, for instance, with the Anthropoid Apes ; there are
often cheek pouches, so that, as Mr. Lydekker has remarked, if a
Monkey be observed to stow nuts away in its cheeks for future
reference, we may be certain that its home is in the Old World,
for the Catarrhines are exclusively denizens of the Old World,
while the Platyrrhines are as exclusively New World in range.
Again, those of the Catarrhines which do possess a 'long tail,
such as the members of the genus Cercocebus, never show the
least sign of prehensility in that tail. The teeth of Jbhe Catar-
rhines are invariably thirty-two in number, the formula being
If C^PmfMf = :',:>.

In the Old -World Apes there is a bony external auditory
meatus, which is wanting (as a bony structure) in the' Platyr-
rhines. The late Mr. W. A. Forbes pointed out that in most of
the New- World forms the parietals and the malars come into
contact ; in the Monkeys of the Old World they are hindered
from coming into contact by the frontals and the alisphenoids.
The Platyrrhines may have the same number of teeth ; this is
the case with the Marmosets, but in them there are three pre-
molars and two molars ; in the remaining New-World Monkeys
there are thirty-six teeth, but of these three are premolars and
three molars.

Not only are these two groups of the Primates absolutely
distinct at the present day, but they have been, so far as we
know, for a very long time, since no fossil remains of Monkeys
at all intermediate have been so far discovered. This has led to
the suggestion that the Monkeys are what is termed diphyletic,
i.e. that they have originated from two separate stocks of
ancestors. It is hard, however, to understand on this view the
very great similarities which underlie the divergences that havt-
just been mentioned. But, on the other hand, it is equally hard
to understand how it is that, having been separated from each
other for so long a period, they have not diverged further in

556 MARMOSETS

structure than they have. The Platyrrhines seem to stand at
the base of the series. This is another example of the existence of
archaic creatures in South America.

GROUP I. PLATYRRHINA.

Fam. 1. Hapalidae. — We may begin the account of the
Platyrrhine Monkeys with the Hapalidae or Marmosets ; for this
family is structurally lower than the rest. They have thirty-two
teeth, arranged as in the following formula : I^| C ^ Pm | M|^ = 32.
The molars have three main tubercles, and not four as in the
higher forms. The digits are for the most part clawed, not
nailed, as in the higher types ; the great toe alone bears a flat
nail. The tail, too, is ringed, a condition which is characteristic
of many of the lower groups of mammals, but not of the higher
Apes. The cerebral hemispheres are smooth, but this is a matter
rather connected with their small size than with low zoological
position. The tails of the Marmosets, unlike those of -so many
other American Monkeys, are not prehensile though they are
long.

The genus Hapale is broadly distinguished from the other
genus, Midas, by the fact that the lower incisors slant forwards
as in the Lemurs. They are small, soft -furred, long- tailed
Monkeys, familiar to every one. There are some seven species,
which are entirely restricted in range to Brazil, Bolivia, and
Colombia, one species only, H. pygmaea, extending northward into
Mexico.

Of Tamarins, genus Midas, there are rather more species —
about fourteen. They are South and Central American in dis-
tribution. Since both these genera are arboreal in habit, it is
extraordinary that they have not the prehensile tails of their
American allies. As, however, the late Mr. Bates observed an
individual of the species M. nigricollis fall head-foremost from a
height of at least 50 feet, alight on its feet, and run off as if
nothing in particular had occurred, it is evident that no extra
prehensile powers are absolutely necessary. Some of the Tamarins
have a long marie ; this is well seen in M. rosalia, or rather in
M. leoninus, which, if not identical with it, is at least very closely
allied to it. The name is obviously derived from the character

xvn OTHER SOUTH AMERICAN MONKEYS 557

referred to, and the Monkey, originally described by the traveller
von Humboldt, is said to have " the appearance of a diminutive
lion." M. bicolor is an example of the species with no mane,
but with a patch of white round the mouth, looking like " a ball
of snow-white cotton " held in the teeth.

Fam. 2. Cebidae. — The remaining American Monkeys are
comprised in the family Cebidae. This is to be distinguished
from the last by the fact that there is an additional molar, thus
making thirty-six teeth in all. The tail, sometimes very short,
is more generally long and highly prehensile, being nude at the
extremity, which • part is therefore especially prehensile ; this
state of affairs is often to be seen in animals with prehensile
tails. The Cebidae, though for the most part larger than the
Marmosets, never approach in size the Old-\Vorld Apes.

Typical of the family is the genus Oebus, including the
'• Capuchin " Monkeys, and consisting of nearly twenty species ;
the tail, though prehensile, is covered with hair to the tip, a fact
which is indicative of a less perfect prehensility than is exhibited
in some Monkeys with a naked under surface to the tip of the
tail. The thumb is well developed. The genus ranges from
Costa Rica to Paraguay. The commonest Monkey which accom-
panies the street organs of this country is a Cebus. It is a
popular delusion that these and other monkeys are purely
vegetable-feeding animals. Cebus is in fact particularly fond of
caterpillars, as are also the Marmosets.

Allied to Cebus is Lagothrix, the Woolly Monkey, of which
L. humboldti is the best-known species, there being indeed but
one other. It is a larger and heavier animal than any species of
Cebus; and the Hare -like woolliness of the fur suggested its
scientific name to its original describer, von Humboldt. It has
a perfectly prehensile tail, naked at the tip. The thumb and
great toe are well developed. These are purely fruit -eating
Monkeys, and are known as " Barrigudos " by the Portuguese of
the Amazon country on account of their prominent belly, due
apparently to the immense amount of fruit consumed. They are,
or were, much eaten by natives.

Brachyteles is a little-known genus, connecting the last with
the next genus. The under fur is woolly ; the thumb is small
or absent. The tail is naked below.

The Spider monkeys, Ateles or Coaitas, have been described as

558

SPIDER MONKEYS

CHAP.

the most typically arboreal of American monkeys. The use of
the prehensile tail can frequently be studied in living examples
in the Zoological Society's Gardens. With this " fifth hand " the
Monkey feels for a place to grasp, and securely twists its tail
round, moving it with the greatest ease from point to point.
When the tail is being thus used it is carried erect over the head.
The fact that this genus possesses no functional thumb is thought

FIG. 265.— Spider Monkey. A teles aler. x^.
%

to be associated with the extreme perfection of its adaptability to
an exclusively arboreal life. The hand without a thumb can
act as an equally efficient hook for suspending the body ; and
what is useless in nature tends to disappear. These Monkeys
have a wide range, extending from Mexico in the north to
Uruguay in the south. There are ten species. The flesh of
many Monkeys is eaten not only by natives but by Europeans ;
but the Spider Monkeys are said to furnish the most sapid food
of all.

The Howling Monkeys, genus Mycetes, have also received the
appropriate generic names of Alouatta and Stentor. The former
of these two names, indeed, is that which should properly be
applied to the genus. But Mycetes is perhaps better known.
The " howling " is produced by saccular diverticula of the larynx,
larger than those of other American Monkeys, such as Aides,
where, however, they are also developed. The hyoid bones, too,

xvii SQUIRREL MONKEYS 559

are enormously enlarged and cavernous, while the jaw — in order
to accommodate and protect these various structures — is unusually
large and deep. The Howlers are furnished with a fully pre-
hensile tail. The thumb is present. They are described as
being the most hideous in aspect of the American Monkeys, and
of the lowest intelligence, with which latter characteristic is
associated a less convoluted brain than in Ateles, for example.
The noise produced by these Monkeys is audible for miles, and
is said not to be due to emulation, i.e. not to be comparable to
singing or talking, but to serve to intimidate their enemies.
The story told of these and otber Monkeys with prehensile tails,
tliat they cross rivers by means of a bridge of intertwined Monkeys,
is apparently devoid of truth. There are six species, which are
Central and South American in range.

The Squirrel Monkeys, genus Chrysothrix, are small creatures
with a long head, the occiput projecting. Their tail, though long,
lias no naked area at the extremity and is non-prehensile. It is
a remarkable fact that the proportions of the cranium as compared
with the face are greater, not only than in other Monkeys, but
than in Man himself. The thumb is short, but not so short as
in the Spider Monkeys. The cerebral hemispheres are very
smooth ; but, as already remarked, this is a matter of size, and
not of low position in the series. It may appear at first sight
that this statement contradicts the one made concerning the
Howlers. But the latter are large Monkeys, and therefore ought,
so to speak, to have a more complex brain ; but they have not.
Like so many of the American Monkeys, the Squirrel Monkeys
are gregarious, and, in spite of their tails, arboreal. They are
largely insect -feeders, and also catch small birds and devour
eggs. There are four species, of which C. sciurea is the commonest,
and is constantly an inmate of the Zoological Society's Gardens.
Humboldt asserted of it that when vexed its eyes tilled with
tears ; but Darwin did not succeed in seeing this very human
expression of an emotion.

Ccdlithrix is a genus not far removed from the last, and, like
it, occurs both in, Central and in South America. It is chiefly
to be distinguished from Chrysothrix by the non-extension back-
wards of the head, and by the more furry character of the tail.
The lower jaw is rather deep, as in the Howlers ; but there is
not, or there has not been discovered, a howling apparatus like

560

DOROUCOULIS

CHAP.

that of Mycetes. Nevertheless Professor Weldon1 has found in
a female of C. yigot a patch of ossification on the thyroid cartilage
of the larynx which may be an indication of something more in
the male. There are eleven species.

NyctipithecAis, the Doroucouli Monkeys, is a genus of some-
what Lemurine appearance, caused by their large eyes. But they
reminded Bates of an Owl or a Tiger-cat ! They have a long, but
not prehensile tail. As in the Marmosets, the lower incisors
project forwards in a Lemurine fashion. The thumb is very
short. A peculiarity of this genus is the twenty- two dorso-
lumbar. vertebrae. As in Chrysothrix, but nut as in Callithrix,
the hemispheres of the brain are smooth. There are five species,
of which one occurs so far north as Nicaragua ; the rest are
Brazilian, extending down to the Argentine.

The Ouakari Monkeys, Brachyurus? are, as the name denotes,
short -tailed forms. Two species, B. rubicundus and £. calvus,

FIG. 266. — Red-faced Ouakari. Brachyv.rus rubicundus. x i.

have bright red faces ; B. melanocephalus has a black one.
There is a small thumb. The brain is fairly convoluted, and is
to be specially compared with that of Celus and Pithecia. The

"Notes on Callithrix gigot," Proc. Zool. Soc. 1884, p. 6.
2 Forbes, Proc. Zool. Soc. 1880, p. 639.

XVII

OUAKARI MONKEYS

56l

species B. rubicundus at any rate has an absolutely as well as
a relatively greater length of intestines and caecum than any
other American Monkey known.

Not the least remarkable fact about these Ouakari Monkeys is
their distribution in South America. We cannot do better than

FIG. 267. — White-nosed Saki. Pitheda albinasa.

(From Nature.)

quote the summary given by Messrs. P. L. and W. L. Sclater in
their Geography of Mammals, which is as follows: "Each of
them, as first shown by Bates and afterwards further explained
by Forbes, is limited to a comparatively small tract of forest on
the banks of the Amazon and its affluents. The Black-headed
Ouakari (B. melanocephalv.s) ... is met with only in a tract
VOL. x 2 0

562 THE SAKIS CHAP.

traversed by the Eio Negro ; the Bald-headed Ouakari appears
to be confined to the triangle formed by the union of the
Amazon with another affluent, the Japura ; and the Eed Ouakari
to the forests on the north bank of the Amazon opposite Olivenc,a,
and lying between the main stream and the Eiver Ic,a. Each of
them evidently takes the place of the others in its particular
district. Of this peculiar kind of distribution few instances are
known amongst mammals, but many somewhat similar cases have
been observed in birds, reptiles, and insects."

The genus Pithecia, the Sakis, consists of five species with
long bushy tails, which are non-prehensile. They are bearded
and have a thumb. Like the last genus, Pithecia does not
extend into Central America. The incisors project forwards, and
the lower jaw is deep, though the howling apparatus of Mycetes
is wanting. The thin, closely-set, and projecting incisors are very
suggestive of those of the Lemurs. Brachyurus is much like
Pithecia in this respect, and both differ markedly from such a
genus as Cebus, where the lower incisors are vertical. An
anatomical peculiarity of Pithecia is the breadth of the ribs.
P. satanas is perhaps the best-known species, but all five have
been exhibited at the Zoological Society's Gardens. As its
name suggests, P. satanas is entirely black ; it shows a curious
point of difference from P. cheiropotes in its way of drinking.
The latter species, as its name denotes, uses its hand to drink,
while P. satanas puts its mouth to the water. P. albinasa is
black with a red patch on the nose, within which again is a
small white patch.

GROUP II. CATARRHINA.

The Catarrhine Apes are divisible into three or perhaps only
two families, the Cercopithecidae and the Simiidae, to which
must be added the Hominidae. The Simiidae are sometimes
spoken of as the Anthropoid Apes.

Pam. 1. Cercopithecidae. — Of the Cercopithecidae there are
eight genera (perhaps nine) to be recognised, which may be dis-
tributed into two sub-families. The first of these two sub-families,
that of the Cercopithecinae, has the following characters : — There
are cheek pouches in which the animals store food temporarily.

XVII

THE GIBRALTAR APE

563

The stomach is simple and globular; this corresponds with a
mixed diet. The tail is long or short, or practically absent.

The most familiar genus is undoubtedly Macacus. This
includes all the common so-called Macaques, the Bonnet Monkey,

FIG. 268. — Tcheli Monkey. Macacus tcheliensis. x J. (From Xature.)

the Pig -tailed Monkey, etc. In this genus we find that the
males are larger than the females, and have stronger canine teeth.
Ischial callosities are well developed. The genus is purely
Asiatic, reaching as far east as« Japan, with the exception of the
Barbary Ape, M. inuus, also known as the Gibraltar Ape. There
are altogether some seventeen species.

Macacus inuus is doubtfully indigenous to Gibraltar. It is,
however, definitely established there at present, and is carefully
fostered. It is a large Ape with no external tail, in which

564 EXTRATROPICAL MONKEYS CHAP.

particular it is unique among the members of its genus. At one
time its extinction on the " Hock " was nearly accomplished, but
three individuals being known. In 1893 the Governor of
Gibraltar informed Mr. Sclater that he had himself counted as
many as thirty in one herd. Its depredations seem to have led
to the expression of a wish in some quarters that the numbers
should be thinned ; but feeling on the opposite side appears to be
stronger, so that whatever was the actual mode of its introduction
on to the " Kock " it will at any rate remain there unmolested
for the present.

M. tcheliensis is a species found in the Yung-ling Mountains
in North China. It is, with the possible exception of M. speciosus,
the most northerly form of Monkey. It is interesting on account
of the fact that like the Tiger of those regions it has put on an
extra coating of fur to enable it to combat with the bitter
winters. It is doubtful whether it is more than a variety of
the Rhesus Monkey (M. rhesus}.

M. nemestrinus, " the Pig-tailed Macaque," is trained by the
natives of the east to climb cocoa-nut palms and to carefully
select and throw down only the ripe fruit. Sir Stamford Baffles
apparently was the first to report upon this useful intelligence of
the animal, and Dr. Charles Hose of Borneo has confirmed him.

The Japanese Macaque (M. speciosus} is well known from the
work of Japanese artists. It is the only species of Monkey
found in Japan, and goes very far north.

A rather rare form is M. leoninus. It has a short tail, and
occurs in Burmah. M. silenus is distinguished by a ruff of long
light-coloured hair surrounding the face. It is sometimes called
the "Wanderoo ; but this is apparently quite inaccurate, since that
term is used by the Ceylonese for a Semnopithecus. For those
who wish a " pseudo-vernacular " name Dr. Blanford suggests
Pennant's name of " Lion-tailed Monkey."

The commonest species of the genus are M. cynomolgus,
M. sinicus, and M. rhesus.

The genus Cercocebus, including those Monkeys known as
Mangabeys, is confined to West Africa. They have always a
long tail, quite as long as the body. The upper eyelids are pure
white in colour. The ischial callosities are more pronounced
than in the Macaques. In the Mangabeys also the hairs are not
ringed with differently coloured bars, as is the case with both

xvii MANGABEYS AND GUENONS' 565

Macaques and Cercopithecus, giving to them the greenish hue
which characterises so many of the last two genera. There are
no laryngeal air sacs as in the Macaques. There are not more
than seven species.

The genus Cercopithecus (the Guenons) represents in Africa
the Oriental and Palaearctic Macaques ; the genus has a long
tail. The cheek pouches are larger than in the genus Macacus.
The ischial callosities are less extensive than in that genus. A
tooth character also distinguishes this genus from Macacus ; the
last molar of the lower jaw has, as a very general rule, only
four cusps instead of the five which are found in Macacus. The
supraciliary ridges in the skull are by no means so marked as in
the allied genera.

One species, the Talapoin, C. talapoin, has been separated into
a distinct genus, Miopithecus, on account of the fact that the
lower molars have only three tubercles instead of the usual four.
But if this be done, then Cercopithecus moloneyi, which has a
lower molar with five tubercles, should also be separated.

The genus Cercopithecus is limited to Africa, and its numerous

FIG. 269. — Diana Monkey. Cercopithecus diana. x J.

species have often a very limited range. They are frequently
rather brightly coloured, with blue and white patches on the face.
The Diana Monkey has a pointed white beard. Of the Vervet
Monkey (C. lalandii) a curious fact was noticed at the Zoological

5 66 BABOONS

Society's Gardens a year or two back : the young was observed to
take both teats of the mother into its mouth at once. Mr. Sclater1
in a recent list of the group allows forty-seven species, of which
thirty-three were examined by himself. Subsequently, however,
the list has been reduced to forty by the same authority. One
of the rarest species is C. stairsi, first described from a skin
stripped from a specimen which lived for a short time at the
Zoological Gardens.

The genus Cynocephalus (or Papio) includes the Baboons ; and
the scientific name indicates the Dog-like aspect of these animals,
due to the projecting snout. Cynocephalus is confined to
Africa and Arabia. Several of the species of the genus are well
known. The Mandrill, C. mormon (or maimori), has blue ridges
on the muzzle, the bridge of the nose being red. The animal
lives in herds, and is ferocious and omnivorous. The Chacma
Baboon, C. porcarius, is the largest of Baboons. It lives in Soutli
Africa in large herds. The Arabian Baboon, C. hamadryas, is
the Sacred Baboon of the Egyptians. The names of two other
species, C. thoth and C. anubis, serve also to remind us of the
ancient Egyptians. There are altogether eleven species of
Cynocephalus.

Gelada (or Theropithecus) is separated as a distinct genus.
Though regarded as a Baboon, Garrod has pointed out many
points of likeness to Cercopithecus.2 The two species are, like
the other Baboons, African.

Cynopithecus niger is a small black Baboon from Celebes. It
has swellings on the muzzle as in other Baboons, but differs from
them in being a more amiable creature as well as in its smaller
size. It has a rudimentary tail, smaller even than the small tail
of the typical Baboons. It has, like them, ischial callosities.

In the second sub -family, Semnopithecinae, the following
characters are distinctive : — All the Apes of this group are slender
in form, with a long tail. There are no cheek pouches. The
stomach is sacculated ; it is divided into three portions. This is
accompanied by an apparently more exclusively vegetarian diet
than characterises other Apes, which mingle with their diet of
fruit a large proportion of insects, eggs, etc.

"On a new African Monkey of the genus Cercopithecus, with a List of the
known Species," Proc. Zool. Soc. 1893, p. 243 ; see also p. 441.
2 Proc. Zool. Soc. 1879, p. 451.

THE GUEREZA MONKEYS

567

The first with which we shall deal is Colobus, containing
the Monkeys known as Guerezas. These creatures are entirely
confined to the African continent, and they are arboreal in habit.
It has been attempted to show that their affinities are more with
the Platyrrhines than with the group in which they are really to
be placed. In favour of regarding them as nearer akin to the
American monkeys are only two facts of importance : the first is
the practical absence of the thumb, which of course recalls the

FIG. 270.— Black Celebesian Ape. CyiwpUhecus niger. x £.

condition characteristic of Ateles ; in the second place, the nostrils
in their wideness somewhat resemble those of the Platyrrhines.
They are slender Monkeys with well-marked callosities. They
have a complex sacculated stomach, resembling the large intestine
of some other animals ; it is not divided into distinct chambers
like the stomach of a Euminant or of a Whale. Correlated
apparently with this large stomach is the small development of the
cheek pouches. This genus, of which there are about ten species,
is characterised by beautiful skins, which are largely collected.
The Arabs have a legend to the effect that one species, when
wounded, and seeing its capture and the removal of its skin

568

SACRED MONKEY OF THE HINDUS

CHAP.

inevitable, carefully tears the latter, that its captors may not
profit by it. The species of this genus are most abundant on
the west coast of Africa. It is interesting that one species,
0. kirki, is limited to the Island of Zanzibar, where, however, it
is nearly extinct.

The " Holy Apes," or Langurs, genus Semnopithecus, are allied
to the last, but they are Asiatic in range. The thumb is better
developed, but still shorter than in other Cercopithecidae ; the
callosities are small, and the cheek pouches are absent. There is
a single large laryngeal sac, and the stomach is complex.

This genus is, like the Tiger, often quoted as an example of a
race supposed to be characteristically tropical, existing habitually
in the coldest climate. A species of Semnopithecus has been
observed climbing snow-laden branches at a height of 11,000
feet in the Himalayas. There are some thirty species, which
extend as far east as Borneo.

The name Semnopithecus is derived from the fact that
the Hanuman is regarded as sacred by the Hindus. The best-

FIG. 271. — Entellus Monkey or Haiiuman. Semnopithecus entellus. x J.

known species of Semnopithecus is this Langur or Hanuman,
8. entellus. Being regarded as a sacred animal, and with the
advantage thus gained, it has become a fell nuisance in gardens
and to crops. Though the veneration with which the Hindoos
regard these animals will not allow them to slay them, they are
exceedingly thankful to a European who will enable them to

LONG-NOSED APES 569

commit a sin vicariously. This Ape has immense powers of
leaping — a space of 20 to 30 feet can be cleared by them if
one side, that from which the leap is taken, be considerably higher
than the other. They are useful to the Tiger hunter, as they
follow and hoot at this, their deadly enemy. 8. schistaceus is a
species which lives at great heights, not less than 5000 feet, in
the Himalayas.

The genus NasaMs is hardly separable from the genus
Semnopithecus. It is a Bornean animal, and is distinguished by
a comical long nose, which not only suggests, but goes beyond, the
aquiline nose of the human species. It is no doubt on this
account that the Borneans, unconsciously imitating our habit of
comparing " natives " in general to Monkeys, call it by a name
which signifies " white man." RMnopithecus has also a long, but
a more definitely upturned nose.

Fossil Monkeys. — Several of the existing genera of Old-World
Apes are also known to have existed in past times ; in some cases
their past distribution indicates a greater range. Thus Macacus
is now represented — and that doubtfully — in Europe by the
Barbary Ape alone. But from Montpellier have been unearthed
the remains of M. priscus, from Pliocene beds. The Asiatic
Semnopithecus is known to have lived during the Pliocene period ;
its remains are discovered in France and Italy, as well as in
Asia. In addition to these existing forms, a number of totally
extinct Old- World genera are known. The rich formation at
Pikermi near Athens has produced Mesopithecus pentelici ; this
Monkey has a skull which recalls that of Semnopithecus, while
the stout limbs are rather Macaque-like. As is the case with
many living Catarrhines, the males have stronger canines. The
animal had a long tail.

An analogous annectent character is shown by the Italian
fossil, Oreopithecus bambolii. This animal was referred by one
palaeontologist to the Man-like Apes, by another to the Cerco-
pithecidae. It suggests a common ancestral form, and is Middle
Miocene in horizon.

Just as there are no Platyrrhine Apes in the Old World so
there are no Catarrhines met with in a fossil condition in the New
World ; the two great divisions of the Apes were as distinct in
the past, so far as we know, as they are now — a strong argument
in favour of those who would derive them from two sources. The

57O CHARACTERS OF SIMIIDAE CHAP.

existing genera, Cebus, Mycetes, and Callitlirix, now living in South
America, are also known in a fossil state. The extinct genus
Homunculus is known from the Tertiary strata of Patagonia, and
an apparently allied form is Anthropops. These creatures, how-
ever, are at present far from exhaustively known.

Fam. 2. Simiidae. — The Anthropoid, or Man-like Apes,1
may be separated from the lower Apes as a group, Simiae,
or perhaps better, on account of the after all slender points of
difference, a family Simiidae, which has the following distinctive
characters.

Though arboreal creatures for the most part, these Apes, when
they come to the ground, progress in at least a semi-erect fashion.
Moreover, when they, as is usually the case, put their hands
upon the ground to aid in walking, they do not rest as do the
lower Apes upon the flat of the hand, but upon the back of the
fingers. None of the Anthropoids has a tail, or cheek pouches.
Ischial callosities are only seen in the Gibbons. There is
commonly a laryngeal pouch, which is of large size, and aids in
the production of the generally loud voice of these creatures.
The hair is rather more scanty than in the Cercopithecidae, which
is an approach to Man. The placenta differs in detail from that of
the lower Apes, and is exactly like that of Man. These Apes show
as further differences from the underlying Cercopithecidae, the
greater length of the arms as compared with the legs, and the
presence of a vermiform appendix to the caecum. In the latter
but not the former character they agree with Man, whom we
shall place in a separate family, Hominidae. The Anthropoid Apes
are entirely Old World and intratropical in range at the present
time.

The Gibbons, genus Hylobates, stand quite at the base of the
series of existing Anthropoid Apes. They are the smallest and
the most purely tree-frequenting of all the members of that
group. Connected with this habit is the structural peculiarity
that their arms are proportionately longer than in the other
Anthropoids. The affinity of the Gibbons to the Catarrhines is
proved by the presence of distinct but small ischial callosities.
The arms are so long that when walking upright the hands
reach the ground. The hallux is well developed. The ribs are
thirteen pairs. In the skull the chief noteworthy character as

1 See the books quoted on p. 576 (footnote).

XVII

SPECIES OF GIBBON

571

compared with other Anthropoids is the fact of the large size of
the canines, which are of equal or nearly equal size in the two
sexes. The molars on the other hand have been particularly

FIG. 272. — Hoolock. Hylobates lioolock. x £.

compared to those of Man. The brain is simpler than in the
higher forms. But it is not clear that this may not be a case
of diminished complexity of convolution going hand in hand
with smallness of size.

The Gibbons range through south-eastern Asia from Assam
and Burmah to Hainan. The number of species is a little
doubtful. It is clear that in the first place we may distinguish
the Siamang, H. syndactylus, which indeed some regard as a
separate genus. It is mainly to be defined by the syndactylous
character of the second and third toes ; they are united by skin
as far as the last joint. The Hainan species, H. hainanus, is
probably distinct, and the following names have been given to
various other species or races, viz. H. agilis, H. leuciscus, H. leuco-
genys, H. lar, If. hoolock. These animals can walk erect ; and
when they do so, the big toe is separated as in unsophisticated
or at least unbooted man. The voice is well known to be loud,

572

EXTERNAL CHARACTERS OF GORILLA

and it is a curious fact that the Siamang, which has a large
laryngeal pouch, is not excelled in this respect by species in
which this sac is not developed.

pa.

FIG. 273. — Cerebrum of the Gibbon (Hylobates). (Lateral aspect.) c.c', c.c", Anterior
and posterior central convolution ; /», interparietal fissure ; fr, frontal lobe ;
f.s, Sylvian fissure ; oc, occipital lobe ; pa, parietal lobe ; s.c, fissure of Rolando ;
tp, temporal lobe; *, fron to - orbital fissure. (From Wiedersheim's Structure of
Man.)

Of Gorillas, genus Gorilla, there is but one species, which must
apparently and rather unfortunately be called Gorilla gorilla.

The misfortune is double : in the first place the repetition of
the same word as both generic and specific appellation is tiresome
to the ears and barbarous in its suggestion ; in the second place
it is now well known that the " Gorilla " of Hanno, observed by that
Carthaginian voyager on an island off the African coast, was not
a Gorilla at all as the word is now understood, but probably a
Baboon. The external aspect of this great Anthropoid is familiar
from many reproductions. The male, as is usual, is larger than
the female, and his characters are more pronounced.

The face is naked and black, and the skin generally is deep
black, even at birth. The ear is comparatively small, and is
adpressed to the side of the head ; it is altogether more human
in form than that of the Chimpanzee, and this statement applies
also to the rudimentary condition of the muscles of the ear,
which are more rudimentary than in the Chimpanzee. The nose
has an obvious median ridge, and is thus pronounced as an external
feature ; the nostrils are very wide. The hands and feet are
short, thick, and broad ; the digits are webbed. In the foot the
heel is more apparent than in other Anthropoids. It is not,

GORILLA AND CHIMPANZEE

573

however, so marked as in Man, and the phrase "Ex pede
Herculem " has been aptly supplemented by " Ex calce hominem."
The hair upon the head forms a kind of crest, which can be
elevated when the animal is enraged. The neck is thick and
short, and the beast has massive shoulders and a broad chest.

If it were not for the fewness of the Anthropoid Apes, and
their nearness to Man, it is doubtful whether the Gorilla would
be ranked as a distinct genus,1 for in internal structure it is

FIG. 274. — Gorilla. Gorilla gorilla, ?. x |.

very near the Chimpanzee. The microscopic character of the
investigations into the anatomy of Man have somewhat dimmed
the proper sense of perspective, and have tended to throw into
greater prominence than seems necessary the divergences of
structure seen in the Gorilla. Dr. Keith 2 has recently summed
up and commented upon these divergences, and the following
account of this Anthropoid is mainly deduced from his memoir.

The cranial capacity of the Gorilla is greater than that
of the Chimpanzee. It is not possible, however, to decide from
this point of view whether a given skull is that of one or of the
other of these Apes. Some Chimpanzees are higher in capacity

1 It is not so ranked by everybody. a Proc. Zool. Soc. 1899, p. 296.

574 GORILLA AND MAN CHAP.

than some Gorillas. But the average is undoubtedly as stated.
It is to be noted that there is a correspondence between cranial
capacity and size of palate, the correspondence being converse, i.e.
the greater the brain the smaller the palate. This applies to
Man as compared with his Ape-like relatives, but does not apply
so accurately to the Gorilla, which has a more extensive palate
than the Chimpanzee ; its " brute development " is much greater
than that of the Chimpanzee. Not only is the palate larger, but
the molar teeth, slightly different in form, are also larger and
stronger. This is so plainly marked that " one may say almost
with certainty, that any upper molar tooth over 12 mm. in
length is that of a Gorilla, and under 1 2 mm. is that of a Chim-
panzee." In the skeleton generally it may be said that the
crests for muscular attachments upon the bones are greater in
the Gorilla. The nasal bones are more like those of lower Apes
in their length, and they have a sharp ridge more marked than
in the Chimpanzee, which, however, disappears in aged animals.
It is a curious fact that Gorillas often have a "cleft palate,"
owing to the failure of the palatal part of the palatine bones
to meet completely. The general conformation of the skull is
less brachycephalic in the Gorilla.

The limbs show a number of small differences, which are
associated with a more completely arboreal life in the Chimpanzee
as compared with the Gorilla. The latter is approaching the
human way of life. In spite, however, of these differences, no
hard and fast lines of divergence can be laid down between the
two African Anthropoids, for it appears from the many memoirs
that have been written upon both that "there is scarcely a
feature in any muscle or bone found in one animal which is not
also found in the other." The heel of the Gorilla has already
been referred to. This is, of course, associated with a plantigrade
and therefore non-arboreal mode of progression. Certain of the
muscles of the calf of the leg attached to the heel show a more
human arrangement in the Gorilla than in the Chimpanzee. It
is interesting to find that the muscles of the little toe are
diminishing in the Gorilla as in Man. This is most clearly due
to terrestrial progression and we may apply the same explanation
to Man and ignore tight boots ! The arm of the Gorilla is less
adapted to arboreal progression. Its proportions differ from those
of the arm of the Chimpanzee in that the fore-arm is shorter. In

xvii MENTAL CHARACTERS 575

both animals the thumb is not of much use, and this digit is
more retrograde in the Gorilla, not only in proportionate length
but in its muscular supply. The hip girdle tells the same tale.
It is broader in the Gorilla, and the glutaei muscles are more
prominent, all these features being connected with the more
erect gait.

The brain of both animals have been studied, but not in the
case of the Gorilla from a sufficiently large number of examples to
make any generalisations of great value. On the whole, the
Gorilla has the larger brain, but this must be discounted by the
fact that it also has the larger body. It is a remarkable fact that
the Gorilla's liver is much more like that of lower Apes than the
liver of other Anthropoids. It has, as has the Chimpanzee,
laryngeal sacs. The general conclusion concerning the relative
position of the two African Anthropoids 'seems to be that the
Gorilla is the more primitive ; and as thus it must approach more
nearly to the original parent than does the Chimpanzee, it may be
said that it also comes rather nearer to Man, since the Chimpanzee
has travelled away from the common stock on another line. The
detailed likenesses to Man, however, are not to be unduly dwelt
upon ; for they mainly come from a tendency to assume the
plantigrade mode of progression.

In mental characteristics there is the widest difference be-
tween the two Apes that we are considering. The Chimpanzee
is lively, and — at least when young — teachable and tameable.
The Gorilla, on the other hand, is gloomy and ferocious, and
quite untameable. When angry the Gorilla beats its breast, a
statement that was originally made, we believe, by M. du Chaillu,
but which has been disputed, though it appears to be perfectly
true. A young Gorilla, exhibited some time since in the Gardens
of the Zoological Society, could be observed to do so. The cry
of the Chimpanzee is different from the " howl " of the Gorilla.
An immense amount has been written upon the ways of this
animal in its own home, including much that is legendary. The
Gorilla has been said to lurk in the depths of the forest, and to
stretch down a prehensile foot to grasp and strangle an un-
fortunate black man passing below. It is said, too, to vanquish
the Elephant by hitting it hard upon the trunk with a stout
stick, and to crumple up the barrel of a rifle with its powerful
teeth.

576 THE PONGO OF BATTEL CHAP.

Apart from the doubtful " Pongo " and " Engeco " of Andrew
Battel, our first intelligence concerning the Gorilla is due to Dr.
Savage, after whom, indeed, the late Sir Eichard Owen called the

O ? '

animal Troglodytes savagei, a name which has to be abandoned
in favour of an earlier name.

The Gorilla is limited in its distribution to the forest tract of
the Gaboon. It goes about in families, with but one adult male,
who later has to dispute his position as leader of the band with
another male, whom he kills or drives away, or by whom he is
killed or driven away. The animal is said to make a nest in a
tree like the Orang ; but this statement has been questioned.

It feeds upon the berries of various plants, and upon other
vegetable substances ; there is apparently not so marked an
inclination for animal food as is exhibited by the Chimpanzee.
In search of their food they wTander through the forest, walking
partly upon the bent hand, and progressing with a shuffling gait.
It is noteworthy that the Gorilla has been said to walk upon the
palm of the hand and not upon the back, as is the case with the
Chimpanzee. It can readily assume the upright posture, and,
in this case, balances itself largely with its arms. Professor
Hartmann, however, states that the back of the hand is also used.
Unlike most or many wild beasts, the Gorilla exhibits no desire
to run away when he views a human enemy. Dr. Savage
remarks that " when the male is first seen, he gives a terrific yell,
that resounds far and wide through the forest, something like
kh-ah ! kh-ah ! prolonged and shrill." This is accompanied by
offensive tactics, which the natives do not willingly encounter.
When making an attack the Gorilla rises to his feet, and as a
full-grown animal reaches a height of some five feet, he is a most
formidable antagonist. The attack of one of these animals is said to
be made with the hand, with which he strikes his adversary to the
ground, and then uses the powerful canines. The beating of the
breast which heralds an attack is a statement made by M. du
Chaillu. It has been denied with a vigour and asperity quite
incommensurate with the importance of the matter.1

The Chimpanzees, genus Antliropopithecus (or Troglodytes), are

1 For accounts of the habits of the Gorilla, compiled from various sources, see
Hartmann's "Anthropoid Apes," International Scient. Ser. London, 1885; H.
O. Forbes, "Monkeys," in Allen's Naturalists' Series, London, 1894 ; and Huxley,
" Man's Place in Nature," vol. vii. of Collected Essays, London, 1894.

BRAIN OF CHIMPANZEE

577

to be distinguished from the Gorilla by the characters mentioned
in the account of the latter animal. Briefly summed up they

fr.

v

Fro. 275. — A, Cerebrum of a female Chimpanzee two years old. x J. (Dorsal aspect,
showing asymmetrical development.) c.c', c.c", Anterior and posterior central con-
volutions ; f.i, interparietal fissure ; f.l, the longitudinal fissure ; f.po, parieto-
occipital fissure ; fr, frontal lobes ; oc, occipital lobes ; s.c, sulcus central is. B,
Brain of a female Chimpanzee two years old. x J. (Lateral aspect.) cb, Cere-
bellum ; c.c', c.c", anterior and posterior central convolutions ; fr, frontal lobe; /.*,
fissnra Sylvii ; is, island of Reil ; md, medulla oblongata ; oc, occipital lobe ; pa,
parietal lobe ; s.c, sulcus centralis ; tp, temporal lobe. (From Wiedersheim's
Structure of Man.)

are mainly as follows : — The ears are large, and generally stand
out from the head ; but there are exceptions to be noted pre-
VOL. x 2 P

578

RANGE OF CHIMPANZEES

sently. The pigmentation of the body is not always so pro-
nounced as in the Gorilla. The nasal bones are shorter. The
skull as a whole is more brachycephalic, and the molar teeth are
smaller. The hands and feet are much longer, the animal being
more purely arboreal than the Gorilla. The female Chimpanzee is
slightly smaller than the male, but the great disparity observable
in the Gorilla does not characterise its ally. The animal, like
the Gorilla, has large air sacs.

Chimpanzees are entirely restricted to Africa, and though

Fid. 276. — Skull of Chimpanzee. Anthropopithecus troglodytes, x ^.
(After de Blainville.)

they appear to extend rather farther east than the Gorilla, the
forest-clad region of the equatorial belt is their home.

It has been mentioned in treating of the Gorilla that the
main feature of this animal, which affords a constant difference
from the Chimpanzee, is its gloomy and ferocious manner. The
Chimpanzee, on the other hand, is lively and playful, though
often maliciously so, and quite tameable, as many instances —
particularly the notorious " Sally " of the Zoological Gardens —
show. The earliest mention of animals that are probably Chim-
panzees is to be found in a work upon the Kingdom of Congo,
published in 1598. In a cut illustrating that work, and of

xvn SPECIES OF CHIMPANZEE 579

which a part is reproduced in Professor Huxley's essay referred
to below,1 the Apes, which correspond roughly in their appearance
to Chimpanzees, are represented as being captured by the device
of limed boots, which the Apes are putting on. This idea has
been subsequently imitated and acted upon. A little later, Andrew
Battel wrote of the Pongo and of another creature the Engeco.
This latter, whatever may be the case with the former, is in all
probability the Chimpanzee, since the word 'Nchego, now applied
to those creatures, seems to be the same word. From this seems
also to be derived the sailor's term " Jacko." Whether there are
or are not more than one species of Chimpanzee, is a matter
which has exercised and perplexed naturalists. That there are
plain differences of external features, at any rate between in-
dividuals, is perfectly clear. We are justified in recognising
three forms, but the question of their specific distinctness may
for the present be held in reserve. The commonest of these is
the variety known as A. troglodytes. This is frequent in mena-
geries, though the specimens on view are nearly always young
and small. The face and the hands are flesh-coloured, and the
ears are very large. The black hair gets a reddish tinge on the
flanks. The second variety is that which was termed by du
Chaillu Troglodytes Jcooloo-kamba. This animal appears to be
also the T. aiibryi of MM. Gratiolet and Alix,2 and to be identical
with two Apes known by the names of " Mafuca " and " Johanna." 3
The former of these was exhibited in Dresden, the latter at
Messrs. Barnum and Bailey's show. The two animals have been
carefully studied. They differ from the common Chimpanzee by
the dark colour of the face, and in the case of Mafuca the ear
was Gorilline in form. So too was the ear of A. aubryi, while
Johanna has a larger one. These features have led to the sug-
gestion that the Kooloo-kamba was the result of a mesalliance
between a Gorilla and a common Chimpanzee.

It has at any rate been stated that the two Anthropoids do
go about in company; but there seems to be little doubt that
there is no question here of a hybrid. Dr. Keith's careful
studies 4 upon Johanna have demonstrated the impossibility of

1 "Man's Place in Nature," vol. vii. of Collected Essays, London, 1894.

2 Hartmann's "Anthropoid Apes," in International Sci. Ser. London, 1885.

3 Nouv. Arch. Mus. Hist. Nat. Paris, ii. 1866.

4 Proc. Zool. Soc. 1899, p. 296.

580

DR. ROMANES AND SALLY

CHAP.

regarding this Ape as anything but a Chimpanzee. The animal
has the ways and manners of the Chimpanzee ; has a cry exactly
like that of A. troglodytes ; does not beat her breast like a Gorilla
when annoyed. Anatomical knowledge, however, of this specimen
is at present wanting.

Antliropopithecus calvus l seems to be at least as much entitled
to distinction as the last. It was originally described by du

A •-;

FIG. 277. — Young Orang-Utan. Simia satyrus. Zeitschrift fur Ethnologie (Anthropolog.
Gesdlschaft), Bd. viii. (From Wiedersheim's Structure of Man.)

Chaillu ; but Dr. Gray who examined the skins thought that the
baldness was accidental, and then after this wise caution pro-
ceeded to describe, under the name of A. vellerosus, perhaps the
"worst" species of Chimpanzee that has been added to the
unnecessarily long list of "species" of Chimpanzees. To this
variety belonged " Sally " 2 of the Zoo, whose intelligence has been
celebrated by the late Dr. Komanes. The form is characterised

1 See also Duckworth, Proc. Zool. Soc. 1898, p. 989.

2 For the structure of this Ape see Beddard, Trans. Zool. Soc. xiii. 1893, p. 177 ;
and for experiments on her intelligence, Romanes, Proc. Zool. Soc. 1889, p. 316.

SALLY AND JOHANNA

58l

by its intense blackness, the red reflection of other Chim-
panzees not being visible ; also by the bald head, whence of
course the name. The nostrils of this Ape, as of Johanna, were
somewhat expanded, and thus present a certain likeness to the
Gorilla. But there can be no suggestion that A. calvus is the
product of a union between the two African Anthropoids. As is

FIG. 278.— Young Oraug-Utan. Simia satyrus. Zeitschrift fiir Ethnologic (Anthropolog.
Gesellschaft), Bd. viii. (From Wiedersheim's Structure of Man.)

the case with Johanna, Sally was given and enjoyed animal food
on occasions. It is a curious fact that both Sally and Johanna
appear to have been colour-blind.

The Orang Utan, genus Simia, has but one definable species,
viz. 8. satyrus. The supposed species of Owen, S. morio, cannot
be satisfactorily defined. Plenty of other specific names have
also been given to what is in all probability but a single species
of large Anthropoid Ape inhabiting the islands of Borneo and
Sumatra.

582

SKELETON OF ORANG

CHAP.

The name Orang-Utan, now applied exclusively to the subject
of the present description, was formerly applied also to the
Chimpanzee, and to that animal, moreover, under the latinised
version of Homo sylvestris. The Orang is a large and heavy Ape
with a particularly protuberant belly and a melancholy expression.
The face of the old male is broadened by a kind of callous expan-
sion of naked skin at the sides. The colour of the animal is a

FIG. 279. — Skeleton of Orang. Simia satyrus. (After de Blainville.)

yellow brown, varying in the exact shade. The ears are parti-
cularly small and graceful in appearance, pressed closely to the
sides of the head. The head is very brachycephalic. The arms
are very long, and when the animal is in the erect posture they
reach as far as the ankle. The hallux is very short and usually
destitute of a nail. It is a curious fact that the head of the thigh
bone is unattached by a ligament to the socket of the pelvis in
which it articulates, a state of affairs which may give the limb
greater freedom in movement, but does not add to its strength ;

NEST OF ORANG

583

indeed, the Orang has been described as moving with laborious
caution.

This Ape inhabits flat and forest-clad ground, and lives mainly
in the trees. The male leads a solitary life except at the pairing
season, but the female goes about with her family. On the
ground the Orang walks with no great ease, and uses his arms as
crutches to swing the body along. Even on trees the rate of
progress is not rapid, and is accomplished with careful investiga-
tions as to the capabilities of the branches to bear his weight.

FIG. 280. — A, Skull of a young Orang-Utan. Simia satyrus. (One-third natural size.)
B, Skull of an adult Orang-Utan. (One-third natural size.) (From Wiedersheim's
Structure of Man.)

The " Man of the Woods " has been stated to build a hut in trees.
This is an exaggeration of the fact that it constructs a temporary
nest.

One of these nests has lately been described elaborately by
Dr. Moebius. It was found (by Dr. Selenka) on the fork of a
tree at a height of 11 metres from the ground. Every night, as
it appears, or every second night, the animal constructs a new nest
for himself, abandoning the old one. So numerous, therefore, are
these nests in localities frequented by Orangs, that a dozen can be
readily found in a day. The particular nest which Dr. Moebius
examined was 1*42 metres long, and at most '80 metre broad. It
was built of about twenty-five branches, broken off and laid for

584 THE FOSSIL MAN OF JAVA CHAP.

the most part parallel to each other. Above this framework a
number of loose leaves lay. There is no doubt, therefore, that
these nests are not by any means elaborate structures, and that
they only serve as sleeping-places, and not as nurseries for the up-
bringing of the young, as has been asserted.

The Orang seems to be usually of a fairly mild disposition ; it
will rarely attack a man unprovoked. Bub Dr. Wallace, who has
accumulated a large number of observations upon these animals,
describes a female Orang who " on a durian tree kept up for at
least ten minutes a continuous shower of branches, and of the
heavy-spined fruits as large as 32-pounders, which most effectually
kept us clear of the tree she was on. She could be seen breaking
them off and throwing them down with every appearance of rage,
uttering at intervals a loud pumping grunt, and evidently mean-
ing mischief." The name given by the Dyaks to the Orang is
Mias Pappan.1

Fossil Anthropoid Apes. — Undoubtedly the most interesting
of fossil Anthropoids is the now famous Pithecanthropus erectus.
Our knowledge of it is due in the first place to Dubois.2 But there
is hardly an anatomist or an anthropologist who has not had his say
upon this regrettably very incomplete remnant. The creature is only
known by a calvarium, two separate teeth, and a femur. And the
femur, moreover, is diseased. M. Dubois discovered these remains
in the island of Java in andesite tufa of Pliocene or at least
early Pleistocene age. The remains were found in company with
Stegodon, which is now extinct, and Hippopotamus, which is no
longer found in that part of the world. The name Pithecanthropus
was given to it by the discoverer in order to furnish with a
definite habitation and a name the theoretical Pithecanthropus of
HaeckeL Even the most particular of students of mammalian
nomenclature will hardly object to the utilisation of a name for a
second time which is with some clearness a nomen nudum !
The animal when erect must have stood 5 feet 6 inches high.
The contents of the cranium must have been 1000 cm., that is to
say 400 cm. more than the cranial capacity of any Anthropoid

1 For the external appearance of the Orang see Hermes, Zeitschr. f. Ethn. 1876,
a paper which has coloured plates.

2 Pithecanthropus erectus. Eine nienschendhnliche Uebergangsform aus Java,
Batavia, 1894. See also Ernst Haeckel, The Last Link (with notes by H. Gadow),
London, 1898 ; Manouvrier, Ainer. Journ. Sci. 1897, p. 213 (extracts) ; and Klaatsch,
Zoolog. Centralbl. vi. 1899, p. 217.

MEN AND APES 585

Ape, and quite as great as or even a trifle greater than the cranial
capacity of some female Australians and Veddahs. But as these
latter are not 5 feet in height, the Ape-like man had really a
less capacious cerebral cavity. The skull in its profile outline
stands roughly midway between that of a young Chimpanzee
(young in order to do away with the secondary modifications
caused by the crest) and the lowest human skull, that of
Neanderthal Man. This creature is truly, as Professor Haeckel
put it, " the long searched for ' missing link,' " in other words
represents " the commencement of humanity."

The remains of Apes, more distinctly Apes than Pithecanthropus,
are known from Miocene strata of France. Two genera, Plio-
pithecus and Dryopithecus, are known. The former appears to be
close to Hylobates. Dryopithecus is more Man-like than any
other, and seems to have been as large as a Chimpanzee. The
incisors are human in their relatively small size. But it has
been pointed out that the long and narrow symphysis of the
lower jaw is a point of likeness to the Cercopithecidae.

Fam. 3. Hominidae. — Apart from Pithecanthropus, which per-
haps is a member of this family, but whose remains permit us to
leave it among the Simiidae, at least for the present, the family
Hominidae contains but one genus, Homo, and probably but one
species, H. sapiens. The characters of the family may therefore
be merged in those of the genus.1

Though it is easy enough to distinguish a Man from an Ape, it
is by no means easy to find absolutely distinctive characters which
are other than " relative." As Professor Haeckel has pointed out,
there are really only four characters which differentiate Man :
these are the erect walk, and the consequent modification of the
fore- and hind-limbs to that position ; the existence of articulate
speech ; the faculty of reason. Whether one body of psycholo-
gists are right who argue that reason is a distinctive human
attribute, not to be confused with the apparent reasoning powers
of lower animals, or whether others are justified in separating
Man only in degree from the lower animals, it is clear that this
very diversity of opinion prevents us for the present from utilising
such characters as absolute differences. In any case the dis-
cussion of these matters is beyond the scope of the present book.

1 See especially Wiedersheim, The Structure of Han, transl. by Howes, London,

1895.

586

HAIRY COVERING

CHAP.

Anatomically there are a number of small points which distinguish
Man ; but they are mainly due to the erect gait. It is some-
times attempted to divide Man as a naked animal. But this is an
apparent difference only ; the hair is not so much developed upon
the body as in the Apes, save in occasional abnormalities, such as

FIG. 281.— Skull of Immanuel Kant. (After C. voii Kupffer.) The great size of the
cranium is a noteworthy feature. (From Wiedersheim's Structure of Man.}

the various hairy men arid women who can be seen in travelling
shows, and to a less extent the Japanese Ainos, but it is present
everywhere, as is shown by microscopical investigation of the skin.
The skull in Man " is a smooth and imposing, rounded or oval
bony case," which contrasts with the smaller and deeply ridged
skull of the Anthropoid Apes. The shape of the skull is largely
in accord with the large brain. The face does not project so
much as in the Anthropoid Apes, though this character must not

HAND AND FOOT

587

be insisted upon too strongly, as in some American Monkeys the
face is as little projecting. Still we are now comparing Man with
his undoubtedly nearest relatives the Simiidae. In the lower jaw
the anterior line at the symphysis is an approximately straight
one, that is at right angles to the long axis of the jaw, while Apes
have a more retreating chin. The " beautiful sigmoid curve
formed by the lumbar and dorsal vertebrae " is more pronounced

Man

Otang

FIG. 282. — Foot of Man, Gorilla, and Orang of the same absolute length, to show the
difference in proportions. The line «'«' indicates the boundary between the tarsus
and metatarsus ; b'b', that between the latter and the proximal phalanges : and c'c'
bounds the ends of the distal phalanges, as, Astragalus ; ca, calcaneum ; sc,
scaphoid. (After Huxley.)

in Man, but exists not only in the Anthropoids, but in other
Apes.1

The fore-limbs are relatively short, the extreme length of the
arm being such that the outstretched hand does not reach the
knee. The thumb is a large and useful digit in Man, much more
so than in the Anthropoids. On the other hand the hallux is not
opposable. This is, of course, correlated with the upright attitude,
as is also the greater relative thickness of that digit, upon which

1 Cunningham, "Cunningham Memoirs," No. II. Royal Irish Acad. 1886.

588

MUSCLES OF MAN

CHAP.

the greatest stress is laid in walking. As to muscles, the glutaeus
raaximus is more developed in Man — the Ape which most nearly
approaches him being the Gorilla, in which animal the life is
less thoroughly arboreal than in some others. The so-called
"scansorius" is only present in Man as an occasional occurrence.

tb

FIG. 283. — Skeleton of the left pes of a Chimpanzee. (Dorsal aspect.) as, Astragalus ;
cb, cuboid ; cl, calcaneum ; ec, ectocuneiform ; en, endocuneiform ; ms, mesocunei-
form ; nv, navicular ; I-V, digits. (From Wiedersheim's Structure of Man.)

The rudimentary character of the ear muscles for the movement of
the external ear in Man has often been insisted upon, as also
their occasional functional activity. But here and elsewhere, so
numerous are the abnormalities, that "the gap which usually
separates the muscular system of Man from that of the Anthro-
poids appears to be completely bridged over." These are words
of Professor Wiedersheim quoted from Testut, and express a final
summary of the matter of muscles in Man and the Apes.

RUDIMENTARY VOCAL SACS

589

In his teeth Man differs by the small exaggeration of the

c

B

pi. • pi- pi.

FIG. 284. — The hard palate, A, of a Caucasian; B, of a Negro ; C, of an adult Orang-
utan, showing the differences in shape of the bones. The palate of the Negro
represents a type transitional between that of the Caucasian and that of the Orang.
mx, Maxilla ; pi, palatine ; p.mx, premaxilla. (From Wiedersheim'a Structure of
Man.)

canines, which hardly, if at all, differ in the two sexes.

also a complete absence of a diastema.

The teeth are also on the whole

weaker than in the Anthropoids,

though Hylolates is very human in

this particular.

There is a tendency in Man
towards the disappearance of the
upper outer incisors, and more
markedly still of the wisdom teeth,
which appear very late, and are
often imperfect. In a large number
of cases the tooth does not appear
at all. In the larynx there is no
great development of the great
throat pouches of the Anthropoids.
The minute diverticula of that
organ, known to human anatomists
as the ventricles of Morgagni, alone
remain to testify to a former howl-
ing apparatus in the ancestors of Man.

There is

-I t/i.

i. 285. — Human Larynx in frontal
section, cr, Cricoid cartilage ; sn,
sinus of Morgagni ; t.c, first
tracheal cartilage ; tlu, thyroid
cartilage. (From Wiedersheim's
Structure of Man.)

INDEX

Every reference is to the page : words in italics are names of genera or species ; figures
in thick type refer to an illustration ; f. = and in following page or pages ;
n. = note.

Aard Vark, 187

Aard Wolf, 413

Abderites, 148

Aceratherium, 258 ; A. incisivum, 259 n. ;

A. platycephalum, molar teeth of, 51 ;

A. tridactylum, 259
Achaenod&n, 279
Acodon, 480

Acomys, 472 ; A. cahirinus, 473
Aconaemys, 488
Acrobates, 142
Adapts, 553

Addax, 315 ; A. nasomaculata, 315
Aeluroidea, 390

Aeluropus, 444 ; Ae. melanoleucus, 444
Aelurus, 431

Aepyceros, 311 ; Ae. melampus, 311
Aepyprymnus, 137 ; Ae. rufescens, 137
Aethurus, 463 ; A. glirinus, 463
Agouti, 495
Agriochoerus, 331 ; skeleton of, 332 ; A.

latifrons, 332
Ai, 171
Alactaga, 486; A. decutnana, 486; A.

jaculus, 486
Alcelaphus, 308
Alces machlis, 297, 298
Alimentary canal, 61
Allodon, 99
Allotheria, 105
Alouatta, 558
Alpaca, 286
Ambergris, 364
Amblotherium, 99
Amblydactyla, 206
Amblypoda, 205 f.
Amblyrhiza, 506

AmTnodorcas, 313 ; A. clarkei, 313
Amwphochilus, 531
Amphicyon, 423 ; A. giganteus, 424
Amphilestes, 99, 100

Amphimeryx, 330

Amphipromverra, 160

Amphitherium, 99 ; ^4. prevostii, 100

Amphitragulus, 301

Anaptomorphidae, 552

Anaptomoi-phus, 552 ; ^4. homunculus, 552

Anchippodus, 507

Anchitherium, 248 ; *•!. aureliense, 249 ;

^4. eqtiinum, 249
Ancodus, 329 ; *4. brachyrhynchus, 329 ;

^4. velaurms, 329
Aucy lopoda, 211 f.
Ancylot/ierium, 211
Angwantibo, 547
Anisodon, 211
Ankle, 43

Jiwtw, 317 ; J. depressicornis, 317
Anomaluridae, 462
.Iwoj/Mi^MrM*, 462 ; .4. _peft, 463
Anoplotheriidae, 332
Anoplotherium, 333
Ant-eater, 166

Anteckinomys, 153 ; .4. lanigera, 153
Antelopes, 307 f.
Anthops, 527 ; .4. ornatus, 527
Anthracotherium, 328
Anthropoid Apes, 570 f.
Anthropoidea, 554 f.
Anthropopitfiectis, 580; ^1. calvus, 580;

^1. troglodytes, 579
Anthropops, 570

Antidorcas, 312 ; .4. euchore, 312
AntUocapra americana, 306
Antilocapridae, 306
Antilope, 311 ; /I. cenncopro, 311
Antlers ; «ee Horns
^4n/roi»tw, 529
^iMMrosoroc, 518, 519
^4orf<w do/ei, 369
,4 onyx, 441
Apar, 177

592

INDEX

Apes, 554 f.

Aplwlops, 260

Archaelurus, 401

Archaeoceti, 384

Arctic region, mammals of, 81

Arctictis, 407 ; A. Unturong, 407

Arctocebus, 547

Arctogaea, 85

Arctogale, 408 ; A. leucotis, 408 ; A.

stigmatica, 408
Arctoidea, 424
Arctomys, 465 ; A. bobac, 466 ; A.

caudatus, 465 ; A. marmotta, 466 ; A.

monax, 466 ^

Arctonyx, 432 ; ^4. collaris, 432
Arctotherium, 445

Argyrocetus, 383 ; -4. patagonicus, 384
Armadillo, 177
Arteries, 66
Artiodactyla, 209
Artionyx, 211 ; five-toed, 271 ; extinct

forms of, 328
Arviacanthis, 473
Arvicola, 477
Ass, Wild, 242
Astrapotherium, 210, 216
^4<eZes, 557 ; ^4. ate/% 558
Atdodus, 254
Atheriogaea, 85
Atherura, 499 ; ^4. africana, 501 ; .4.

fascicidata, 501
Auchenia, 286
Aulacodus, 489
Aurochs, 318, 321
Avahis, 539 ; .4. laniger, 539
Aye-aye, 548 ; legends relating to, 550

Babirusa, 277 ; £. atfurus, 278

Baboon, 566

Badger, 432

Balaena, 358 ; U. australis, 359 ; jB. 6is-
cayensis, 360 ; .B. glacialis, 359 ; -B.
mysticetus, 359 ; bones of hind-limbs of,
353 ; cervical vertebrae of, 352 ; ster-
num of, 352

Balaenidae, 358

Balaenoptera, 355 ; B. australis, 356 ; B.
borealis, 356 ; sternum of, 33 ; B.
musadus, 356 ; sternum of, 352 ; B.
patachonica, 356 ; B. rostrata, 356 ;
B. sibbaldii, 356

Balaenopteridae, 355

Baleen, 354

Bandicoot, 156

Banteng, 317, 318

Bassaricyon, 428 ; B. alleni, 428 ; B.
gabbii, 428

Bassnriscus, 428 ; B. astutus, 429

Bat, 521 f. ; wings of, 522

Bathyergus, 481

Batomys, 473, 474 ; B. granli, 473

Bdeogale, 410

Bear, 442 ; Grizzly, 442 ; Himalayan,

442 ; Malayan, 443 ; Polar, 443 ; Sloth-,

443

Beatrix Antelope, 314
Beaver, 467

Behemoth, equivalent to Mammoth, '228
Beisa, 315
Beluga, 372
Berardius, 368 ; B. arnouxi, 368 ; B.

vegae, 369

Bettongia, 137 ; B. lesueuri, 138
Bezoar stones, 287, 325
Bibos frontalis, 317, 318
Binturong, 407
Bison, 318 ; B. americanus, 319 ; B.

europaeus, 318
Blaaubok, 314
Blarina, 518
Blarinomys, 480
Blessbok, 309

Boar, Wild, of Europe, 275, 276
Body cavity, 69
Bolodon, 99
Boneia, 526
Bontebok, 309
Bos, 317 ; B. alleni, 319 ; B. antiquus,

319 ; B. bonasus, 318 ; B. depressicor-
nis, 320 ; B. ferox, 319 ; B. frontalis,
318 ; B. gaums, 317 ; B. grunniens,

320 ; B. latifrons, 319 ; B. mindorensis^
320 ; B. primigenius, 321 ; B. sondai-
cus, 318 ; B. taurus, 320

Boselaphus, 316 ; B, tragocamelus, 316

Bovidae, 307

Brachymeryx, 331

Brachyodont, 48

Brachytarsomys, 480

Brachyuromys, 480

Brachyurus, 560 ; 5. calvus, 560 ; 5.

melanocephalus, 561 ; .B. rubicundus*

560

Brady podidae, 170
Bradypus, 170 ; 5. tridactylus, 171 ;

skull of, 171 ; skeleton of, 172
Brain, 75 ; of Dog, 76 ; of Rabbit, 77 ;.

of Echidna, 110, 117 ; of Wallaby, 118
BramatJierium, 306
Brontops, 264
Brontotherium, 264
Bruta, 161

Bubalis, 308 ; B. caama, 308
Buffalo, 321
Bunodont, 272
Bush Dog, 414

Cachalot, 363 ; ferocity of, 365 ; food of,.

365

Cadurcotherium, 264
Caenolestes, 146 ; C. obscurus, 146
Caenopus, 262
Caenotheriidae, 329
Caenotherium, 329

INDEX

593

Calamodon, 192 ; C. simplex, 192

Ccdlignatkus, 367

Callinycteris, 526

Callithrb:, 559 ; C. gigot, 560

Caloprymnus, 138 ; C. co.mpestris, 138

Camel, 285

Camelidae, 285 ; ancestry of, 289

Camelus, 285 ; C. bactrianus, 285, 286 ;

C. dromedarius, 285
Canidae, 413
Canine teeth, 49
Canis, 416 ; C. aegi/ptiacus, 419 ; C.

antarcticus, 418 ; C. anthus, 420 ; C.

aureus, 420 ; C. azarae, 418 ; C. caw-

crivorus, 414 ; C. chama, 418 ; (7.

chanco, 421 ; C*. dingo, 421, 422 ; C,

famtiiaris, 422 ; (7. ferus, 422 ; C.

hodqphyla.>:, 418 ; (7. jubatus, 414 ; C.

lagopus, 419 ; (7. lanvjer, 421 ; (7.

kiteralis, 420 ; C. ta<r«ns, 417 ; C.

lupus, 420 ; (7. magdlanicus, 418 ; C.

mesomelas, 420 ; C. mikii, 422 ; C'.

niger, 418; (7. pallipes, 418; C. par-

videns, 415 ; (7. primaevus, 423 ; (7.

rudis, 418 ; (7. urost ictus, 415 ; C.

mriegalus, 420 ; C. rete, 418 ; C'.

virginianus, 418 ; C. vulpes, 419 ; C'.

zerda, 417
Cannabateomys, 488 ; C. amblyonyx,

488

Caperea, 358
C"«/w«, 324 ; C1. aegagrus, 325 ; C. /«/-

coneri, 325 ; C, ifiec, 325 ; C. jemlaica,

324 ; C. pyrenaica, 325 ; C1. sinaitica,

325

Capreolus, 295 ; (7. capraea, 295
Capromys, 489, 490 ; (7. melanurus, 490 ;

(7. pilorides, 490
Capuchin Monkey, 557
Capybara, 491
Cardinal vein, 68
Cariacus, 295 ; <7. chilensis, 296 ; C*.

maerotis, 295 ; C'. rufus, 296
Carnivora, 386 f. ; C. Fissipedia, 387 f. ;

C. Pinnipedia, 446 f.
Carpomys, 473 ; (7. melanurus, 473 : C.

phaeurus, 473
Carterodon, 489
Castor, 467 ; (7. canadensis, 468 ; C. ./Jder,

468

Castoridae. 467
Castoroides, 506
Cat, Domestic, 10, 14, 392, 400 ; -tribe,

390 ; coloration of, 392 ; Wild, 399
Catarrhines, 555, 562
Cart«, 493 ; C. porcellus, 493
Caviidae, 491

Cavy, 493 ; Patagoniau, 492
Cebidae, 557
Cfe6ws, 557

Celaenomys, 474 ; C. silaceus, 474
Cenoplacentalia, 103

VOL. X

Centetes, 511 ; C'. ecandatus, 511 ; skull

of, 512

Ceutetidae, 511
Cephalogale, 423
Cephalophus, 309
Cephaltii-hynchiis, 380
Cephtdotes, 526
Ceratorhinvs, 254, '257
Cercocebus, 564
Cercolabes, 498 ; C. rillosus, 42, 499 ; (7.

insidiosus, 498
Cercolabidae, 497

Cercoleptes, 429 ; C. caudivolrula, 430
Cercomys, 489
Cercopithecidae, 562
Cercopithecus, 565 ; C. diana, 565 ; C*.

ln/diidii, 565 ; (7. moloneyi, 565 ; C.

stairsi, 566 ; C'. fa1«i>»<it, 565
Cerricapra, 311 ; C'. iscdtellina, 18
Cervidae, 291
L'emdits, 295
Ce/Ti/*, 293 ; C. rf«»»«, 293 ; C. rfan'rft-

«««*, 293 ; C. diir<ni,;-n, 291 ; C.

elaphus, 293 ; (7. luehdwffi, 294 ; C.

sedrjivicki, 301 ; C'. */£«, 291
Cetacea, 339 f. ; primitive position of, 120
c/i>i' iiohyus, 280
Chaetomys, 499
Chalicotherium, 211
Chalinolobus, 530
Chamois, 326
Cheetah, 400
Chestuuts of Horse. 240
Chevrotain, 282
Chillingham Cattle, 321
Chilmxys, 480
('liiiimrrogcde, 518, 519
Cliimpnnzee, 576 ; brain of, 577 ; species

of, 579

rhhK-hilla. 496 ; rr. lanigent, 496
Chinchillidae, 496
Chirogale, 543 ; C. coquereli, 544
Chiromyiclae, 548

ciiiromys, 548; (7. madayascarieims, 548
Chironectes, 156
t'ltiri'/imfoiiiyii, 473
Chiroptera, 521 f.
Chiru, 311
C/iirummys, 472
Chlai»>/>/"jih"r>i*, 173, 176
Choeropotamvs, 280
Choeropsis, 273
Choeropus, 158 ; C.castannti*, 158 : nianus

of, 157
rW<»y/w«, 170; C'. didactylus, 170; '.

hiifinanni, 171 ; scapula of, 164
Chriacidae, 552
C/iriiK-its, 602

(.'limtiiinus, 474 ; C. ichiteheadi, 474
Chrysochloridae, 514
Chrysochlvris, 514 ; C. trevelyuni, 515
Chrysothrix, 559 ; C'. sciiireu, 559

2 Q

594

INDEX

Civet Cat, 406

Clavicle, 38

Claws, on tip of tail of Lion and Leopard,

41 ; see also Nails
Coaitas, 557
Coassus, 296
Coati, 430
Cobus, 310 ; (7. ellipsiprymnus, 310 ; C.

maria, 311 ; (7. unctuosus, 310
Codogenys, 493 ; 0. paca, 493 ; C. taczan-

oicskii, 494

Coelops, 527 ; C. frithii, 527
Coendou, 498
Cogia ; see Kogia
Colobus, 567
Condylarthra, 202 f.
Condylura, 518
Conepatus, 439
ConUurus, 473
Connochaetes, 309 ; C. albogulahis, 309 ;

(7. gwtt, 309 ; (7. taurinus, 309
Conoryctes, 193
Conoryctidae, 193
Coracoid, 38
Cormura, 531
Corpus callosum, 77, 125
Corypfwdon, 207 ; C. eocaenus, 207 ; (7.

radians* skeleton of, 208
Cotton Rats, 479
Coypu ; see Myopotainus
Crateromys, 474
Craurothrix, 473
Creodonta, 449, 455 f.
Cricetomys, 472

Cricetus, 479 ; (7. frumentarius, 479
Crocidura, 518

Crocuta, 411 ; (7. maculata, 412
Crossarchus, 410
Crossopus, 518 ; C.fodiens, 519
Crumen, of Antelopes, 13 ; of Deer, 299
Crunomys, 474 ; C.fallax, 474
Cryptophractus, 178
Cryptoprocta, 404 ; (7. ferox, 405
Ctenodactylidae, 490
Ctenodadylus, 490 ; (7. gundi, 490
Ctenomys, 488

Cuniculus, 478 ; (7. torquatus, 478
Ciiscus, 140

Cydopidius, 331 ; (7. simus, 331
Cydoturus, 167 ; 0. didactylus, bones of

manus of, 169
Cynaelurus, 400 ; (7. brachygnatha, 401 ;

(7. jubatus, 400 ; (7. lanigera, 401
Cynictis, 410 ; C. penicillata, 410
Cynocephalus, 566 ; (7. anubis, 566 ; C.
hamadryas, 566 ; (7. maimon, 566 ; (7,
mormon, 566 ; (7. porcarius, 566 ; (7.
tfurfA, 566
Cynodesrmis, 424
Cynodictis, 423
Ci/nogale, 409
Cynomys, 465 ; (7. ludovicianws, 465

Cynopithecus niger, 566, 567

Cynopterus, 526

(72/o»i, 416 ; (7. dukkunensis, 416 ; (7.

primaevus, 416
Cystophora, 453 ; C. cristata, 453

Dacrytherium, 333

Dactylomys, 488 ; Z). dactylinvs. 488

Dactylopsila trivirgata, 141

Daedicurus, 185

Damaliscus, 309

Daphaenus, 424

Dasymys, 472

Dasypodidae, 173

Dasyprocta, 494 ; Z>. aguti, 495

Dasyproctidae, 493

Dasypiis, 173 ; 7). minutus, 178 ; Z).

sexcinctus, skull of, 174 ; pelvis of, 176 ;

X>. villosus, 178 ; manus of, 175
Dasyure, 151
Dasyuridae, 149

Dasyuroides, 154 ; Z). bi/rnei, 154
Dasyurus, 151 ; skull of, 152 ; />. geoffroyi,

152 ; 1>. hallucatus, 153 ; Z>. macu-

latus, 152 ; Z>. iirsinus, 18 ; Z>. viver-

rinus, 152
Deer, 11, 291 f.
Ddphinapteriis, 372 ; Z>. kingii, 373 ; Z).

leucas, 373
Delphinidae, 372
Ddphinus, 377 ; Z). ddphis, 377 ; Z).

longirostris, 377 ; Z). roseiventris, 377
Dendrohyrax, 234
Dendrolagus, 135 ; 7). bennetti, 136 ; Z).

inustus, 136
Dendromys, 472, 476 ; Z). mesomelas,

476

Deomys, 476 ; Z). ferrugineus, 476
Dermal plates, in Armadillos, 173 ; in

Whales, 342
Dermoptera, 520
Desman, 518
Desmatippus, 248
Desmodus, 532 ; Z>. rvfns, 532
Diabolus, 151 i

Diaphragm, 69
Diceratherium, 259
Dichobune, 333
Dichodon, 330
Dididurus, 531
Dicotyles, 279 ; Z). tajafu, 278
Dicotylidae, 278
Dicroceras, 301
Didelphyidae, 155
Didelphys, 155 ; Z). crassicaitdata, 156 ;

Z>. dimidiata, 13 ; Z). virginiana, 155
Dinictis, 401
Dinoceras, 210
Dinocerata, 210
Dinomyidae, 495

Dinomys, 495 ; Z>. branickii, 496
Dinotheriidae, 231

INDEX

595

Dinotherium, 231 ; D. gigauteum, skull of,

231

Diphylla, 532

Diphyodonty, definitiou of, 51
333

, 519
Dipodidae, 484

IiilKxImHijs, 484 ; D. merriami, 484
Mprotodon, 146 ; Z>. australis, 147
Diprotodontia, 128 f.
Dipus, 485; Z>. acyyptius, 485; pes of,

485 ; Z>. hirtipe*, 484
Jjistaechu-ftts, 141
Distribution, geographical, 78 ; in the past

83

Dogs, 413 ; Raccoon-like, 415
Doliclwtis, 493 ; D. patachonica, 492
Dolphin, 372

Donuthenum ; see Hyomoschus
Jjorcopsis, 135 ; Z>. luctuosa, 135, 136 ;

Z>. madeani, 136 ; X>. mitelleri, 136
Doi-cotm<jus, 313 ; A*, megalotis, 313
Dorinou.se, 470
Doroucouli, 560
JSremotherium, 301
, 98

141 ; Z> nana, 141
Dromiciops, 156
Drydestes, 99
Dryopitkeeiu, 585
Duck-billed Platypus ; see Platypus and

Or/i ithorhyuckus
Dugong, 336 ; skeleton of, 336
Duplicidentata, 502 f.
Dymecodon, 518

Echidna, 110 ; egg of, 72 ; JE. uculeata,
111 ; skull of, 108 ; shoulder girdle of,
109 ; brain of, 110 ; E. hystrix ( = K.
aculeata), 16 ; shoulder girdle of, 37

Echidnidae, 110

Echiiufmys, 488

Eckinops, 512

Echiolhrix, 473

Edentata, 161 f.

Eland, 316

Elaphodus, 294 ; E. cephalophus, 294 ; E.
michianus, 294

Elatnwgnathus, 251

Elasmothcrivm, 259

Elephant, 217 ; observations of Pliny and
Aristotle upon, 225

Elephantidae, 217

Elephas,2lS ; E.ajn^mi'.-i, 221.222 ; -skull
of, 218 ; E. antiquus, 229 ; E.falconeri,
229 ; E. indicus, 221 , 223, 224 ; foot
of, 198 ; E. inelilensis, 229 ; E. meri-
dionalis, 229 ; E. planifrons, 229 ; E.
prim {genius, 226 ; E. priscus, 226

Elifjmodontia, 480

Eliomys, 471

EUurus, 480

Elk, 297

Ellvbius, 479

Elotherium 11 intense, 279

SmbaUoHttm, 530

Eniballonuridae, 530

Enhydriilun, 440

Enhytlris, 439

Enhydrocyoii, 423

Eohippus, 248

Eonycteris, 526 ; .£. spdaea, 526

Epauorthidae, 145

Kji'iinifthits, 148

Epicondylar foramina, 39

Epipubes of Monotreines and Marsupials,

116

Episternum, 33, 34. 35
EjMjiiii>i>hiiri'it, 525
E<iuidae, 237
Eqi'Hs, 239 ; J?. africantis, 243 ; A1. •

243 ; A:. ioeAwii, 245 ; A', burchelli. 244,

245 ; £. cabidlus, 239 ; A', jrreryi, 244 ;

Jt'. hemionus, 24'2 ; £. hemippus, 242 ;

JS1 onager, 242 ; £. przeioalskii, 241 ;

A. quagga, 244 ; A', sivalensis, 246 ;

A*, somalicus, 244; A', sttnunis, 246;

jB. taeniopus, 243 ; A'. 2e&ra, 244
Erethizon, 498
Ergot, of Horse, 240
Ericidus, 512 ; A\ setosus, 512
Erinaceidae, 509

Ei-iit'K-eun, 510 ; A', europaeus, 510
Ermine, 436
Eschatia, 290
-, 507
a, 358
tes, 485

Enjyetawnis, 467 ; A^. cincrens, 467
Euphyseter, 367
Eupleres, 403 ; £. goudotii, 403
Knprotogonia, 204 ; E. puercensis, 204
EurhinodtlpJiis, 383
Eusmilus, 402
Eutatus, 179

Entheria, 116 ; earliest forms of, 102
Evotomys. 479

Fallow Deer, 293

Felidae, 390

/Ww, 391 ; F. feu%y, 397 ; /: cn/ra,
400 ; /•'. fanadensis, 397 ; A', ca^iw,
399 ; K concolor, 399 ; .F. <ryra, 399 ;
/'. &•<>, 393 ; /'. /.»//<j-, 397 ; /'. manieu-
/nfii, 400 ; F. nebulosa, 396 ; /'. OMCO,
398 ; /". pardalis, 398, 399 ; A', pardina,
397 ; A", pardus, 395 ; A'. ri//a, 397 : /•'.
/(<//•/'«, 394 ; A', u/ia'a 396 ; /'. r«i-«n'na,
397

Fere*ia, 377

FiAw, 477 ; A', osoyoosensis, 478 ; /'.
zibethicu-s, 478

Flying Fox, 524 ; skeleton of, 523

Flying Squirrel, 466

596

INDEX

Fore-limb, 39

Fossa, 405

Fossa, 407 ; F. daubentoni, 407

Fossil mammals, 96

Fox, 419 ; Arctic, 419

Fur Seals, 451

furia, 531

Galago, 543 ; G. crassicaudatus, 543 ; G.

gametti, 543 ; G. senegalensis, 543
Galeopithecidae, 520
Galeopithecus, 521 ; G. volans, 520
Galera, 434
Galeriscus, 434

Gcdictis, 433 ; G. barbara, 433
Galidia, 404 ; G. elegans, 404
Galidictis, 404 ; G. vittata, 404
Ganodonta, 190 f.
Gaur, 317
Gayal, 318

Gazella, 312 ; G. loderi, 312
Gdada, 566
Gelocits, 283
Gemsbok, 315
Genet, 406

Genetta, 406 ; (?. vulgaris, 406
Geogcde, 513 ; (?. aurita, 513
Geomyidae, 483
Geomys, 483
Georhychus, 481
Gerbillus, 475 ; (?. aegyptius, 475 ; 6-'.

pyramidum, 475
Gibbon, 570
Giraffa, 301 ; (?. attica, 3T5 ; Cr. australis,

303 ; (?. camelojMrdalis, 302 ; (?. sival-

ensis, 305
Giraffidae, 301
Glands, of skin, 12 ; of Lemurs, 12, 537 ;

of Marsupials, 13 ; of Skunk, 13, 439 ;

on feet of Rhinoceros, 13, 254 ; tem-
poral, of Elephant, 12 ; musk, 13, 300 ;

mammary, 16
Glauconycteris, 530
Gliridae, 470
Globicephalus, 374 ; G. brachypterus, 375 ;

G. indicus, 375 ; G. melas, 375 ; hand

of, 345 ; G. scammoni, 375
Glossophaga, 531
Glossotherium, 180
Glutton, 435
Glyptodon, 184 ; G. clavipes, skeleton of,

184

Glyptorlontidae, 184
Gnu, 309

Goats, species of, 324
Golunda, 473

Gomphotheriuin sternbergi, foot of, 197
Goral, 327
Gorilla, 572, 573 ; brain of, 575 ; habits

of, 576

Gorilla, 572 : G. gorilla, 572
Grampus, 375

Grampus, 375 ; G. griseus, 375

Graphiums, 471

Gravigrada, 179

Greenland Whale, 358

Grisonia, 433 ; G. allamandi, 434 ; G.

vittata, 434
Ground Sloths, 179
Grypotherinm, 182
Grysbok, 310
Guinea-pig, 493
Gulo, 435 ; G. luscus, 435
Gymnobelideus, 141
Gymnura, 509 ; G. rafflesii, 509
Gymnuromys, 480

Habrocmna, 487 ; H. bennetti, 487

Hair, structure of, 6 ; sensory, 10 ; tuft

of elongated, on wrist, 10 ; of Whales,

341

Halichoerus, 452 ; H. grypvs, 452
Halicore, 336 ; H. australis, skeleton of,

336 ; H. dugong, 337 ; H. indicus, 33
Halitherium, 335
Hallomys, 480
Hamster, 479

Hapale, 556 ; H. pygmaea, 556
Hapalemur, 541 ; H. griseus, 541 ; arm

of, 537 ; H. olivaceus, 541 ; H. simus,

541

Hapalidae, 556
Hapalomys, 473
Hapalotis, 473
Haploceros montanus, 326
Haplodon, 469 ; H. major, 469 ; //. rufus,

469

Haplodontidae, 469

Hare, 504 ; Cape Jumping, 486 ; -Kan-
garoo, 134 ; Variable, 504
Harnessed Antelopes, 315
Harpyia, 526
Harpyionycteris, 526
Hartebeest, 309

Heart, 65 ; of Monotremata, 66
Hedgehog, 510
Helaletes, 250

Helictis, 438 ; H. subaurantiaca, 438
Hdiphobius, 481

Helladotherium, 306 ; H. duvernoyi, 306
Helogale, 410
Ilemicentetes, 512
Hemigale, 408 ; IT. hardwicki, 408 ; /f.

Aosei, 408
Hemigalidia, 404

Hemiyanus, 191 ; ^T. otariidens, 191
Herpestes, 409 ; #. albicauda, 409, 410 ;

#. ichneumon, 409 ; /T. wrva, 409
Heierocephalus, 481 ; ZT. ^ader, 481 ; #.

phillipsii, 481
Heterodont, 47
Heteromyidae, 484
JJeteromys, 484
Hexaprotodon, 274

597

Hind-limb, 42

Hipparion, 250

Hippopotamidae, 273

ffippopotamtu, 273 ; H. amphibius, 273,

274

Hipposiderus, 527
Hippotigris. 246
Hippotragus, 314 ; H. equinus, 314 : //.

leucophaeus, 314; 77. w<V/«', 313 f.
Hodomys, 480
ffolochilus, 480
Holmneniscus, 290
flomacod&n, 280 ; 77. vayans, 280
Homalodontotherium, 212, 216
Hominidae, 585
77o»«0, 585 ; skull of, 586 ; foot of, 587 ;

palate of, 589 ; H. sapiens, 585
Homocamelus, 290
Homodout, 47
Homunculus, 570
Hoolock, 571
Hoplophoneus, 402
Horns, 200, 307
Horse, 239 ; Wild, 240 ; striping of, 240 ;

Domestic, 241 ; ancestry of, 247
Howling Monkeys, 558
Humpback Whale, 356
Hunter i us, 358
Hunting Dog, 416
Hutia, 490
Hyaena,, 411 ; H. crocuta, 412 ; H. spelaea,

413 ; H. striata, 412
Hyaenarctos, 445
Hyaenictis, 413
Hyaenidae, 411
Hyaenodon, 455
Hybrid Oxen, 317
Hydraspotheriitm, 306
Hydrockoenis, 492 ; H. capybara, 491
Hydrcnnys, 474 ; H. chrysogaster, 474
Hydropotes, 296 ; //. inermis, 296, 297
Hylobates, 570; 77. oflri^w, 571; H.

hainanns. 571 : H. hoolock, 571
Hylomys, 509 : 77. suillus, 509
Hyomoschus, 283 ; //. aquaticus, 283
Hyopotamus, 329
Hyotherium, 280
Hyperoodon, 23, 41, 370 ; 77. planifrons,

'371 ; 77. rostratum, 371
Hypogenmys. 480
Hypselodont. 48
Hypsiprymninae, 131
ffypsip-rymnodon, 138 ; J/. moschatus,

138

Hypsiprymnodontinae, 138
Hymchyus, 262 ; 77. agrarius, 262
Ht/racodon, 146, 262; 77. nebrascensis,

260

Hifmcodmitotherium, 333
Hyracoi.lea, 232 f.
Hyrricops, 203
J/yraffitherium. 247

i Hyrax, 232 ; 77. capensis, 233 ; manus of,
198

Hystricidae, 499
i Hystricomorpha, 487

Hystrix, 499 ; 7/. cristaia, 499, 500

Ibex, 325

Ichneumon, 409

Jr/ifJiyomys, 480 ; 7. hydrobates, 480 ; 7.

stolzmanni, 480

A-/ /<•//«/( . 414 ; 7. venaticus, 414
Ictimyx, 438
Idivrus, 463 ; 7. macrotis, 463 ; 7. zenlceri,

463

Incisor teeth, 48

Jndris, 538 ; 7. brericaudatn, 538
Jndrodon, 552 ; 7. malaris, 552
7«m, 381 ; 7. geo/rensis, 382
Iniopsis caiicasica, 383
Insectivora, 508 f.
Intercentia. 24

Intestine, (52 ; large and small, 64
fsectolophns, 250
Ismnys, 473
Ivory, of Elephant, 227

Jackal, 420
Jaguar, 398
Jerboa, 484
Jerboa Rat, 473

Kanchil, 283

Kangaroo, 132 ; Tree-, 136

Kerivoula, 529 ; A', ^icto, 529

Kiang, 242

Kidney, 68

Killer " Whale, 341, 375

Kinkajou, 430

Klipspringer, 310

Koala, 142, 143

. 366 ; /f. brericeps, 367 ; JST. ftoiceri,
367 ; A'. j;o««t, 367 ; A". «/»»<«, 367
Kudu, 316

Lagenorhynchus, 378 ; 7>. acutus, 378 ;

7>. albirostris, 378
Lagidium, 496 ; Z. curieri, 496
Lagomyidae, 505
Lagomys, 505 ; 7,. ladacensis, 505 ; Z.

roylei, 505
Lagorchestts, 134 ; Z. conspicillatus, 134 ;

7x. lrjK)roid(s, 135
LngostoMtix, 496 ; X. trich<«l«ctylus, 496,

497

Lagostrophus, 137 ; L.fasciatus, 137
Lagothrix, 557 ; Zr. humbcldti, 557
Lagotis, 496
Zawia, 286 ; Z. huanacot, 286, 287 ; Z.

vicugna, 286
Ungur, 568
7x*/«w, 439
Ltggada, 472

593

INDEX

Lemming, 478

Lemur, 541 ; L. catta, 542 ; L. leucomys-

tax, 542 ; L. macaco, 542 ; L. niger-

rimus, 54li ; L.' rufipes, 542 ; L. rarius,

12, 542

Lemur, Gentle, 540
Lemuridae, 538
Lemuroidea, 534

Leopard, 395 ; Snow, 396 ; Hunting, 400
Lepilemnr, 540 ; L. mvstdinus, 540
Leporidae, 502

Leptarctus, 431 : L. primaemts, 431
Lepta/uchenia, 331
Leptomeryx, 284
Leptonyx, 453 ; L. wcdddli, 453
Lepus, 502 ; L. americanus, 504 ; L.

crawshayi, 504 ; L. cumculus, 502 ;

brain of, 77 ; Z. europaeiis, 504 ; Z.

timidus, 504 ; Z. whytei, 504
Lestodon, 183
Limgxomys, 476
Limnofelis, 456

Limnogale, 512 ; Z. mcrgulus, 513
Linsang, 406
Lion, 393
Liponyx, 526
Listriodon, 279
Lithocranius, 313
Litopterna, 267
Liver, 64

Loncheres, 489 ; Z. guianae, 489
Lophiodon, 250
Lophiodontidae, 247
Lophiomys, 476 ; Z. imhausi, 476
Lophodont teeth, definition of, 51
LopJiatragus michicmus, 294 n.
Lophuromys, 472
Zori's, 23, 547 ; Z. gracilis, 547
Lungs, 69

Zwira, 440 ; Z. vulgaris, 441
Lycaon, 416 ; /,. anglicus, 416 ; Z.

pictws, 416
Lycyaena, 413

Lyncodon, 437 ; Z. luganensis, 437
Lynx, 397
Lytta, a rod formed of connective tissue

within the tongue, 416

Macacus, 563 ; JZ cynomolgus, 564 ; JZ

I'WMMS, 563 ; JZ leoninus, 564 ; J/.

nemestrinus, 564 ; Jf. priscus, 569 ; J/.

rAesws, 564 ; J/. silenus, 564 ; 3/.

sinicus, 564 ; Af. speciosm, 564 ; JZ

tcheliensis, 563, 564
Machaerodontidae, 401
Machaerodus, 402 ; Af. pohnidens, 402
Macleayius, 358
Macravchenift, 267
Macraucheniidae, 267
Macroglossus, 526
Macropodidae, 129
Macropodinae. 131, 132

Macropus, 132 ; Jf. brunii, 133 ; JZ
giganteus, 132; Jf. MTOW, 133.; J-Z
rufus, 132

Macrorhinus, 453 ; JZ leoninus, 454
Macroscelidae, 515
Macroscelides, 515
Macrotherium, 211
Malacmnys, 472
Malacothrix, 476

Mammals, position in Vertebrate series, 1 ;
characters of, 3 ; coloration of, 10, 11 ;
Reptilian ancestors of, 90 ; earliest
known forms, 96
Mammoth, 226
Man, 585
Manatee, 335

Manatus, 335 ; JZ inunguis, 336
Mandrill, 566
Mangabeys. 564
Manidae, 188
Afanis, 188 ; JZ gigantea, 190 ; JZ

macrura, 25 ; JZ tricuspis, 31
Manteoceras, 267

Marine Mammals ; see Sirenia, Cetacea
Marmosa, 156
Marmoset, 556
Marmot, 465 ; Prairie-, 465
Marsupial Mole, 159
Marsupials, 122 ; imperfectly born young

of, 124

Marsupium or Pouch, 14 ; of Monotremata,
15 ; of Marsupials, 17 ; rudiments of,
in higher Mammals, 23, 18
Marten, 435
Massoutiera, 490
Mastacomys, 473
Mastodon, 230
Meerkat, 410, 411
Megachiroptera, 524 f.
Megaderma, 528 ; JZ lyra, 528
Megaladapididae, 554
Megaladapis insignis, 554 ; JZ. madagas-

cariensis, 554
Megaloglossus, 526
Megalomys, 480
Megalonychidae, 183
Megalonyx, 183
Megamys, 506
Meganeuron kre/ti, 366
Megaptera, 356 ; JZ capensis, 357 ; JZ
indica, 357 ; JZ. lalandii, 357 ; JZ
longimana, 357
Megatherium, 183
J/efes, 432 ; JZ anakuma, 432 n. ; JZ

taxtts, 432

Mellivora, 437 ; JZ capensis, 438
Melonycteris, 526
Melursus, 443 ; JZ labiate, 443
Meniscoe-ssus, 101
Meniscotherium, 203
Menodus, 264
Mephitis, 439

INDEX

599

Meriones, 475
Mermaids, 337
Merychyus, 331
Merycochoerus, 330
Merycopotmiius, 329 ; M. nanus, 329
Mesohippus, 248
Mesomys, 489
Mesonyx, 456
Mesopithecus pentelici, 569
Mesoplacentalia, 103

Mesoplodon, 369 ; J/. Strews, 369 ; If.
europaeus, 369 ; Jl/. liectm'i, 369 ; JdT.
layardi, 369 ; J/. stejneyeri, 369
Mesoreodon, 330
Metamynodon, 264 ; J/. planifrons,

skeleton of, 263
Metatheria, 116
Microbiot/ierium, 160
Microcebus, 544 ; J/. smithii, 564
Mierochiroptera, 526 f.
Microdipodops, 484
Microgale, 512 ; J/. longicaiulata, 25
Microlestes, 98 ; J/. antiquus, 98
Microtus, 477 ; -fl/. agrestis, 477 ; J/.

amphihius, 477 ; 3/. glareolus, 477
Midas, 556 ; J/. bicolm; 557 ; Jf. Zeo-
ninus, 556 ; J/. nigricollis, 556 ; .V.
rosalia, 556

Mimicry, in Bassaricymi, 428 ; in Tupaiu,
511

Miniopterus, 529 ; J/. scheibersi, 529
Mioclaenidae, 205
Mioclaenus, 205
Miohippus, 248
Miopitliecus, 565

Mixocebus, 540 ; J/. caniceps, 540

Molar teeth, 51

Mole, 517 ; sternum of, 31 ; fore-foot of,
40 ; Golden, 515

Mole Rat, 481

Molossus, 531

Monachus, 453 ; J/. albiventer, 453 ; A/.
tropicalis, 453

Mongoose, 409

Monkeys, 554 f.

Monodon, 373 ; J/. monoceros, 373

Monophyodonty, definition of, 53

Monotremata, 105 f.

Moose, 297, 298

Moropus, 211

Morse, 451

Moschinae, 299

Moschus, 299 ; J/. moschifenis, 299, 300

Mouflon, 324 ; Cyprus, 322

Mouse, 472 ; Pharaoh's, 409

Multituberculata, 98

Multituberculy, definition of, f>9

Muntjac, 295

Muridae, 471

J/«s, 471 ; Jlf. decumanus, 472 ; ^/.
nmsculus, 472 ; J/. minutus, 472 ; J/.
raltus, 472

Muscardinus, 470 ; J/. m-ellanarius, 470
Musk, 300 ; Deer, 12, 13, 299 ; Kangaroo,

138 ; Ox, 327
Musquash, 477
Mustela, 435 ; J/. foina, 435 ; jl/. inartes,

435 ; Af.pntorins( = Putoriusfoetidus),

436 ; J/. zibeUina, 435
Mustelidae, 431
Mycetes, 34, 558

Mydans, 433 ; J/. meliceps, 433

Mylodon, 179 ; JV/. robustus, 179 ; skele-
ton of, 180

Mylodon tidae, 179

Myocastor, 489

Myodes, 478 ; J/. lemmies, 478

Myogale, 518

Myomorpha, 469

Afyopotamus, 489

Myoscalops, 481

Myoswex, 518

Myoxus, 470 ; J/. #/t's, 470

Myrmembius, 154 ; M. fasciatus, 155

Myrmecophaya, 166 ; J/. jubata, 166 :
skull of, 167, 168 ; hand of, 169 ;
vertebrae of, 163

Myrmecophagidae, 166

Mystacoceti, 353 f.

Myxopoda, 529

Nail-tailed Wallaby, 134

Nails, 14

Nandinia, 408

Nannosciurns, 464

Narwhal, 373 ; " horn " of, 49

Nasalis, 569

Nasuo., 430 ; JV. nasica, 431 ; Ar. r^/a,
430, 431

Natalus, 529

Native, Bear, 142 ; Cat, 152

Necrodasypus, 164 11.

Necrogymnura, 510

Nectogale, 518 ; J\r. elegans, 520

Nectomys, 480

Nelomys, 488

Nelsonia, 480

Nemorrhaedus, 326 ; iV. crispvs, 326 ; ^V.
grora?, 327

Neobalaena, 361 ; ^V. marginata, 362

Neofiber, 478 ; iV. o/toit, 478

Neogaea, 85

Nemneris, 374 ; A', phocaeiwldes, 374

Neomylodon listai, 181

Neotoma, 480

Neotragns pyijmaens, 310

Nesodon, 215

Nesokia, 472

Nesonycteris, 526

Nesopithecus, 553 ; .A", australis, 553

New Zealand, absence of terrestrial mam-
mals from, 85

Nilgai, 316

Nimravvs, 401 ; *V. gomphodu*, 401

6oo

INDEX

Noctilio, 530 ; X. leporinus, 530

Nomarthra, 186 f.

A'othocyon, 415

Notiomys, 480

Notiosorex, 518

Notogaea, 85

Notopteris, 526

Notoryctes typhlops, 159

Notoryctidae, 158

Nototherium, 148 ; X. mitcMli, 148 ;

skull of, 148

Nyctereutes, 414 ; X. procyonides, 415
Nycteridae, 527

Nycteris, 528 ; X. javanica, 528
Xydicebns, 545 ; -AT. tardigradus, 546 ;

foot of, 537
Nycticejus, 530
Nyctinomus, 531
Nyctipithecus, 560
Nyctophilus, 529

Ocapia johnstoni, 305
Oceanic islands, mammalian fauna of, 81
Ocelot, 398, 399
Ochotona, 505

Octodon, 487 ; 0. <%MS, 487
Octodontidae, 487
Odobaenus, 451

Odontoceti, 362 f. ; fossil forms of, 383
Oginorhinus, 453
Okapi, 305

Ommatophoca, 453 ; 0. rossi, 453
Onohippidium, 247 n.
Onychodectes, 193 ; 0. tissonensis, 193
Onychogale, 134
Onychomys, 479
Opolemur, 544

Opossum, 155 ; Thick-tailed, 156 ; Vir-
ginian, 155

Orang Utan, 580, 581, 582, 583. 587, 589
Orca, 375 ; 0. gladiator, 341, 375
Orcella, 376 ; 0. brevirostris, 376 ; 0.

fluminalis, 376
Oreinomys, 475
Oreodon, 330
Oreodontidae, 330
Orenpithecus bambolii, 569
Oreotragus scdtator, 310
Orias, 316 ; 0. canna, 316 : 0. liviny-

stonii, 316
Oruithodelphia, 106
Ornithorhynchidae, 112
Ornithorhynchus, 112 ; 0. anatimts, 112 ;

shoulder girdle of, 34 ; skeleton of, 114
Orohippiis, 248
Orycteropodidae, 187
Orycteropus, 187 ; teeth of, 188 ; 0.

aethiopicus, 188 ; 0. capensis, 187 ; 0.

yaudryi, 188
Or//:/;, 314 ; 0. beatrix, 314 ; 0. fteisa,

315 ; 0. leucoryx, 314
Oryzomys, 480

Oryzoryctes, 513

Otaria, 451 ; 0. gillespiei, 451 ; 0. hookeri,
451 : 0. jubata, 448 ; 0. nigrescens,
451 ; 0. pusilla, 450 ; 0. ursina, 451

Otariidae, 450

Otocyon, 415 ; 0. megalotis, 415

Otomys, 475 ; 0. unisulcatus, 475

Otonycteris, 529

Otter, 441 ; Sea, 439

Ouakari Monkey, 560 ; Red-faced, 560

Oulodon grayi, 380

Ounce, 396

Ourebia, 310

Ova, 71 ; of Mouotremes, 72

Ovaries, 70

Ovibos moschatus, 327

Oviduct, 73 ; of Marsupials, 74

OM'S, 321 ; 0. blan/ordi, 323 ; 0. burrhel,
322 ; 0. montana, 322 ; 0. musimon,
324 ; 0. nahura, 324 ; 0. qphi&n, 322 ;
0. ^o^', 321 ; 0. tragelaphus, 323 ; 0.
vignei, 321

Oxen, 317

Oxymycterns, 480

Paca, 493

Pachynohphus, 248

Pachyrucos, 213

Pachyuromys, 475

Palaeochoerus, 280

Palaeoerinaceus, 510

Palaeonictis, molar teeth of, 56

Palaeosyops, 266

Palaeotragus, 306

Palate, 61

Palm Civet, 407

Pancreas, 64

Panda, 431

Pangolin, 188

Pangolin gigantesque, 211

Panochthns, 185

Panther, 395

Pantholops, 311

Pantolambda, 206 ; P. bathmodon, 207 ;

skull of, 206
Pantolestidae, 328
Paradoxurns, 407 ; P. grayi, 407 ; P.

niger, 407

Patriofelis, 449, 456
Peccary, 278
Pecora, 290 f.

Pectinator, 490 ; P. spekii, 491
Pectoral girdle, 35
Pedetes, 486 ; P. c«/e?-, 486
Pedetidae, 486

/-We«, 311 ; P. capreolus, 311
Peltephilus, 186
Peludo, 177
Pelvis, 41
Peraceras, 262

Peragale, 156 ; P. lagotis, 157
Perameles, 158

INDEX

60 I

Peramelidae, 156

Peramus, 99

Peramys, 156

Perascalops, 518

Pr.riptychHs, 204 ; P. rhabdodon, 205

Perissodactyla, 235 f.

Perodicticus, 547

Perodipus, 484

Perognathus, 484

Peromyscus, 479

Petauroides, 142

Petaurus, 141 ; P. breticeps, 142

Petrodromus, 516 ; P. tetradactyliis, 516

Petrogale, 134 ; P. penicillata, 134 ; P.

xanthopus, 134
Petromys, 488
Phacochoerus, 277 ; P. aethiopicus, 277 ;

P. africanus, 277

Phalanger, 140 ; P. macidatus, 140
Phalanger, 138 ; Flying, 141 ; Striped,

141 ; King-tailed, 141
Phalaiigeridae, 138
Phalangista vulpina, 140
Phascolarctos dnereus, 142, 143
Phascologale, 153 ; P. Virginia, 153
Phascolomys, 144; skull of, 145; P.

latifrons, 145 ; P. mitchdli, 145 ; P.

ursitius, 145 ; P. tcomlmt, 144
Pkascolonus, 148

PfuuGolotheritm, 99 ; P. bucklandi, 99
Phatagin, 188

Phenacodus, 202 ; P. primaerus, skele-
ton of, 196 ; molar teeth of, 56
Phenacomys, 479
Phlaeomys, 473
Phoca, 452 ; P. groenlandica, 452 ; P.

hispida, 452 ; P. vitulina, 452 ; skeleton

of, 447
Phocaena, 374 ; P. commnnis, 374 ; P.

dallii, 374 ; P. spinipennis, 374 ; P.

tuberculifera, 342
Phocidae, 452
Pholidotus, 188
Phyllorhina, 527
Phyllostomatidae, 531
PhyUotis, 480
Physeter, 363 ; P. macrocephalus, 363 ;

P. tamo, 366
Physeteridae, 362
Pkysodmi, 383
Pichy-ciego, 176
Tig, 275 ; sternum of. 32 ; foot of, 199 ;

solid-hoofed, 270
Pika, 505
Pinnipedia, 446 f.
Pithecanthropus erectus, 584
Pithechirus, 473
Pithecia, 562 ; P. Minasa, 561 ; P.

cheiropotes, 562 ; P. satanuy, :<<'<-
Placenta, 125
Playiaulax, 99
Plafjiodontia, 489

Platacanthomys, 471 ; P. lasitmis, 471

Platanista, 380 ; P. gangetica 381

Platanistidae, 380

Platycercomys, 486

Platypus, 113

Platyrrhines, 555, 556

Plecotvs, 529 ; 7J. auritus, 529

Pli-xiomeryx, 329

J'linj>i(kecus, 585

Poebrotherium, 288 ; P. hillntnui. foot of,

197 ; P. trilsoni, skull of, 288
Poecilogale, 437 ; P. "///////"•//«, 437
Poephaijux yrnnniens ; see .Bwt grunniens
Pogonodon, 401
Poianu, 407
Polecat, 436
Polycladus, 301
Polyprotodontia, 149 f.
Pontoporia, 382 ; P. blainvUlii, 382
Porcula salmnia, 276
Porcupine, 499, 500 ; Tree, 498 ; Brush-
tailed. 501
Porpoise, 374
PotaniocJwerus, 278
Potamogale, 513 ; P. reto, 513
Potamogalidae, 513
Potoroiuae, 131, 137
J'otorous, 138
Potto, 547

Pouch ; see Marsupium
Prairie-dog, 465
I'fUfjins, 178
Prepollex, 41
Primates, 533 f.
Priodon, 173, 179 ; P. giganteus, hand of

175

Prionodon, 406 ; P. pardicolor, 406
Priscotlelphinus, 384
Proaelurtis, 401
Probosci<lea, 216 f.
Procamelus, 289 ; P. angustidens, 290 ;

P. occidentaiis, foot of, 197
Procavia, 232
Procyon, 427 ; P. eancrmtrus, 4'27 ; P.

/,,/,„•, 427 ; P. nigripes, 427
Procyonidae, 426
Prodelphinus, 377
/'roer/iidna, 111; P. bruijnii, 111; P.

iiliiroaculeata, 111
Pronghorn, 306
I'ropalaehfiplophoru-s, 185
/'ro/iithecus, 539 ; P. coquereli, 540 ; /'.

verreaiixi, 540

Prarastoma, 336 ; P. rerunenx, 337
ProsquaJoflon, 384
Proteles, 413 ; P. cristatu, 413
Protelvtkerium, 280
Prothylacinus, 160
Protoceras, 284
Protoceratidae, 284
Protuchriacns, 553
204

602

INDEX

Protohippus, 249

Protolabis, 289

Protoreodon, 332

Protorotlieriidae, 268

Protorotherinm, 268

Protoselene, 205

Prototheria, 105 f.

Protylopus, 287 ; P. peter soni, feet of,

197

Protypotherium, 213
Psammomys, 475
Pseudamphicyon, 424
Pseiulochirus, 141
Psetidorca, 376 ; P. crassidens, 376 ; P.

merulionalis, 376
Pseudorhynchocyon, 516
Psittacotlierium, 191 ; P. muttifragum,

191

Pteralopex, 526 ; P. atrata, 526
Pteromys, 466 ; P. alborufus, 466 ; P.

petaurista, 466
Pteroneura brasiliensis, 441
Pteropodidae, 524
Pteropus, 524 ; P. edulis, 524 ; P. fuscus,

skull of, 524 ; P. nicobaricus, 526 ; P.

polioccplialus, 525
Ptilocerctis, 511 ; P. Jew/, 511
Ptilodus, 101
Pudua, 296
Puma, 399
Putarius, 435 ; P. brasiliensis, 437 ; P.

erminea, 436 ; P. foetidus ( = Mustela

putorins), 436 ; P. hibernicus, 437 ;

P. vulgaris, 437

Pygmy, Hog, 276 ; Right Whale, 361
Pyrotherium, 232

Quagga, 244, 246

Rabbit, 502 ; skull of, 503 ; oviduct of,

73 ; brain of, 77
Raccoon, 427

Rangifer tarandus, 298, 299
Raphiceros, 310
Rasse, 406

Rat, 472 ; -Kangaroo, 137
Ratel, 437, 438
Red Deer, 293
Reed Buck, 311
Reindeer, 292, 298, 299
Reithrodon, 480
Reithrodontomys, 480
ReithrosciuTus macrotis, 464
Reproductive organs, 70
Respiratory organs ; se« Lungs
Rhackianectes, 357 ; /?&. glancus, 357
Rhinoceros, 254 ; /jL bicwnis, 257, 258 :

.R/4. indicu.1 ( = Rh. unicomis), 265 ;

femur of, 43 ; .R/t. lasiotis, 257 ; /ta.

schleiermacheri, 258 ; 7i/i. SI'WIMS, 257 ;

.R/i. sondaicus, 256 ; .R&. smnatrensis,

256 ; hand of, 199 ; «/(. tichorhinus, 259

Rhinoceros, White, 257 ; Black, 257

Rhinocerotidae, 253

Rhinogale, 410 n.

Rhinolophidae, 527

Rhinolophus, 527 ; Rh. ferrum equinum,

527 ; Rh. hipposiderus, 527
Rhinonycteris, 527
Rhinopithecus, 569
Rhinopoma, 530
Rhipidomys, 480
Rhizomys, 482 ; ^JA. badius, 483 ; /fA..

pruinosus, 482 ; .R/t. sumatrensis, 482
Rhynchocyon, 516 ; PA. chrysopygus,

516

Rhynchogale, 410

Rhynchomys, 474 ; /2A. soricioides, 474
Rhynchonycteris, 531
Rhytina, 338
Ribs, 29 ; single - headed, of Whales and

Monotremes, 30
Risfco's Dolphin, 375
River Hog, Red, 278
Roan Antelope, 314
Rodentia, 458 f. ; brain of, 461 ; teeth of,

459

Roe Deer, 295
Romerolagus, 504
Rorqual, 355

Ruminantia, 280 f. ; stomach of, 281 f.
Riipicapra, 326

Sable, 435 ; Antelope, 313

Saccopteryx, 531

Saccostomus, 472

Sacrum, 24

Sagmatias, 376

Saiga, 311 ; & tartarica, 311

Saki monkey, 562

Salivary glands, 64

Samotherium, 305 ; S. boissieri, 306

Sarcophilus ursirius, 151 ; skull of, 149

Scales, 9 ; of Anomaluridae, 463 ; of
Mauidae, 188

Scalops, 518

Scapasius, 518

Scapteromys, 480

Scaptonyx, 518

Scapula, 36 ; of Whales, 37

Scelidotlierium, 180

Schizodon, 488

Schizotherium, 211

Sciuridae, 463

Sciuropterus, 467 ; <ST. volucella, 467

Sciurus, 463 ; <S. castaneiventris, 464 ; &
laticaudatus, 511 ; <S. maximus, 464

Sderopleura, 179 ; £ bruneti, 179

Scotophilus, 529

Scotozous, 530 ; AS dormeri, 530

Seal, 452 ; Bladder -nosed, 454 ; com-
pared with Zeuglodonts, 385 ; Elephant,
454 ; Eared, 450 ; Hooded, 453

Sea-Lion, 450 ; see Otaria

INDEX

603

Sealskin, 9

Sea Serpent, 368

Selenodont, 272

Selenodontia, 280

Semnopithecus, 568 ; S. entellus, 568 : X.

schistaceus, 569
Sewellel, 469
Sheep, Barbary, Blanford's, and other

species of, 321 f.
Shoulder girdle, 35; of Ontthorhynckus,

34 ; of Echidna, 37
Shrew, 518 ; Elephant-, 515
Siamang, 571
Sibbald's Rorqual, 2, 356
Sigmodon, 479
>SY«i if sat urns, 580, 581 ; skeleton of, 582 :

skull of, 583 ; nest of, 583
Simiidae, 570
Simocymi, 423
Simplicidentata, 462 f.
Sing-sing Antelope, 310
Siphneiis, 479
Sirenia, 333 f.
Ximtherinm, 306
Size, of Mammals, 2 ; progressive secular

increase in, 103 ; of Whales, 342
Skull, 25 ; of Human embryo, 27 ; of adult,

586 ; of Dog. 26, 29 ; comparison of,

with that of Anomodontia, 28
Skunk, 439

Sloth, 170 ; Ground-, 180
Sinilodon, 402 ; S. neogaeus, 402
Smiiithopsis, 153 ; S. virginiae, 153
Sminthus, 486
Smutsia, 188

Solenodon, 514 ; S. cubanus, 514
Solenodontidae, 513
Sorex, 518 ; S. alpinus, 510 : >'. mi'itnf".-;

519 ; S. vttlgaris, 519
Soricidae, 518
Soricuhis, 518
Sotalia, 378 ; S. gadamu, 379 ; S. gui-

anensis, 378 ; S. lentiginoso, 379 ; S.

pallida, 382 ; S. sinensis, 378 ; S. teaszii,

378

Sonslik, 464
Spalacidae, 482

Spalacopus, 487 ; S.pofjipigi, 487
Spalacotkerium, 99
Spalax, 482 ; S. typhlv.s, 482
Sparassodonta, 160
Speke's Antelope, 315 f.
Speothos, 423
Sperm Whale, 363
Spermaceti, 363
Spermophilus, 464 ; .S'. tmli'1-imlinfatus,

466

Sphingitrns, 498 ; S. jtrthensilis, 498 f.
Spider Monkey, 658
Spiny Anteater ; see Echidna
Springbok, 312
Hqualodon, 384

Squalodontidae, 384

Squamata, 188

Squirrel, 463 ; Pygmy, !64 : Ground, 464 ;

Flying, 466
fSleatomys, 473, 476
StegodoH, 2-29 ; X ganet .
Steinbok, 310
Steller's Sea-cow, 338
AVf««, 37!': >'. i» 'ii/ii/'-i-, 379

XffHoi'rlpIiix, 382

ifi, 441
*, 545
Stenorhynchus, 453
Stentor, 558
Stereognathus, 99
Sternum, 31 ; of Whales, 32, 33, 352 : of

Dugong, 33
Stoat, 436
Stomach, 62 ; of various mammals, 63 :

of Whales, 347
Strepsicerns, 316 ; N. imltfrhix, 316 : ."•'.

kudu, 316
Stylacodon, 99
>•///////<«//»«, 19:1 ; X. f)/fhi<ln/i-i; 193 ; X

minis, 193
Stylinodontidae, 191
Sublingua, 61
Suidae, 275

Suricata, 410 : .Sf. tetrado.ct >/hi , 411
istis, 275; S. barbatus, 276; >'. .•/•///»"/'-

thius, 279 ; S.salranin, 276 ; .V. *<•/•"/'</

275 ; £ sennaariensis, 275 : »S¥. m -

rucosiis, 276 ; & vittatus, 276
Synaptomys, 479
Synetheres, 498

Synotus, 529 ; .S'. barbastellus, 622, 529
Bystemodon, 250

Tachyoryrte.i, 483

Talpa, 517 ; T7. eurnjKiea, 517

Talpidae, 516

Ti'miiiiilini, 167 ; sternum of, 168

Tamarin, 556

TII in I UK, 464

Taphozoiis, 531

Tapir, 11

Tapiridae, 250

Tapirvs, 2oO ; 7". bairdi, 251 ; 7*. rfoict,

251 ; 7". emadvrensis, 251 11. ; 7". indicus,.

252 ; 7*. lencoyetiys, 251 n. ; 71. roulini,

251 : 7*. terrestri.i, 251
Tarsiidae. 550
Tarsipedinae, 140, 145
Tarsipes, 145

Tarsius, 550 ; J. sj>fi-(rinti, foot of. 651
Tasmanian. Devil, 151 : Wolf, 150
Tatusia, 173 ; 7". nnrfmcinctn, 178
Tau-rotragvs oryx, 316 n.
Taxideu, 438
Tayra, 433
Teeth, 43 ; minute structure of, 44 :

development of, 62 ; brachyodont (with

604

INDEX

short roots), 48 ; bunodout, 272 ;

iliphyodont, 51 ; heterodont, 47 ; homo-

dont, 47 ; hypselodont (with long roots),

48 ; lophodont, 51 ; mouophyodout,

53 ; selenodont, 272
Teiedu, 433

Teleoceras, 259 n. ; T.fossiger, 261
Telmatotherium, 267 ; T. cornutum,

267

Temperature of Monotremata, 112
Tenrec, 511
Testes, 75
Tetraceros, 310
Tetraprotodon, 274
T/uilassarctos, 443
Theropithecus, 566
Thoatherium, 268
Thomomys, 483

Thrinacodus, 489 ; T. albicauda, 489
Thrynomys, 489 ; T. swindeniianus, 489
Thylacinus, 150 ; T. cynocepludus, skull

of, 150
Thylacoleo, 147 ; T. carnifex, skull of,

147

Thyroptera, 529
Tiger, 394
Tillodoutia, 506 f.
Tillotherium, 507 ; T. fodiens, 507 ; skull

of, 507
Tinoceras, 210
Titanops, 264
Titanotheriidae, 264
Titanotherium, 264 ; T. coloradense, 264 ;

T. datum, 265 ; T. Jieloceras, 264 ; T.

platyceras, 265, 266 ; T. ramosum, 266 ;

T. trigonoceras, 265
Tolypeutes, 173 ; T. tricinctus, 176,

177

Tomitheriiim, 552
Tongue, 61
Toxodon, 214
Toxodontia, 214 f.
TrcKjelaphus, 315 ; 71. grahts, 316 ; 71.

spe&ii, 315 ; T7. sylvaticus, 316
Tragulidae, 282
Tragulina, 281
Trayulus, 282 ; 7*. meminna, 282 ; T7.

Jiffpif, 283 ; T7. staiileyanus, 283
Tree- Porcupines, 498 ; Kangaroos, 136
Triaenops, 527
Trichechidae, 451
Trichechus, 451
Trichomys, 488

Trichosurus, 140; 7. vulpecula, 140
Tricky s, 501 ; T7. lipura, 501
2Vjcto, 149 ; 7". oseillans, 149
Triconodon, 99
Trituberculata, 99
Trituherculy, definitiou of, 56
TrUyliMlon, 98
Trtylodytes, f>76 ; 7*. «»tiryi, 579 ; T7.

kooloo-kaipta, 579 ; T7. snvagei, f>7t>

Troffontherium, 469

Tubulidentata, 186

Tuco-tuco, 488

Tupaia, 511 ; T7. belangeri, 511 ; T.

ta««, 511
Tupaiidae, 511
7Vm0, 379 ; T. borealis, 380 ; 71. peronii,

380
Tursiops, 379 ; T7. abusalam, 379 ; 71.

tursio, 379
Tusks, 49
Tylomys, 480
Tylopoda, 285 f.
Typhlomys, 471
Typotheria, 212 f.
Typotberium, 213

Uintacyon, 424

Uintatlierium, 210

Unau, 170, 171

Ungulata, 195 f.

Urinary organs, 68

Uromys, 473

UropsUus, 518

Urotrichus, 518

Ursklae, 442

Urson, 498

Ursus, 442 ; £7. arctos, 442 ; {/. fossilis,

442 : 6r. malayamis. 443 ; f/. ornatus.

443 ; £7. pruinosus, 443 ; t7! spelaeus,
442, 444 ; 6T. tibetanus, 442

Urus, 321

Vampire, 532

Vampyrus, 531 ; T*. spectrum, 531

Vandeleuria, 473

Vansire, 404

Veins, 67 ; anterior abdominal, in Echidna,

68

Vena cava, double in Elephants, 68
Vertebrae, 19 ; cervical, 21, 22 ; dorsal,

20, 23 ; lumbar, 20, 23 ; caudal, 24
Vespertilio, 529 ; F. murinus, 529
Vespertiliouidae, 528
Vesperugo, 528 ; F. discolor, 528 ; F.

feWm', 528 ; F. noctula, 528 ; F.

pipistrellus, 528 ; F. serotinus, 528
Vicuna, 286
Vishnutherium, 306
Ft'remz, 405 ; F. civetta, 406
Viverricula, 406
Viverridae, 403
Vizcacha, 497
Vole, 477
Vulpine Phalanger, 140

Wallaby, 129 ; skeleton of, 130 ; Nail-
tailed, 134
Walrus, 451
Wapiti, 292
Wart Hog, 277
Waterbuck, 310

INDEX

605

Water Chevrotain, 283

Weasel, 437

Whalebone, 354

Whalebone Whales, 354

Whales, 339 ; hunting of, 360

White Whale, 372

Wild Boar of Europe, 276

Wild Cat of Europe, 399

Wisent, 318

Wolf, 420

Wombat, 144 ; skull of, 145

Wrist, 40 ; tactile hairs of, 10

Xenarthra, 166 f.

Xenomys, 480

Xenurus, 173, 178 ; X. unicinctus, 178

Xeromys, 474

Xertis, 464

Xiphodon, 3'29

Xiphodontidae, 329
Xiphodontotherium, 330

Yak, 320
Yapock, 156

Zaglossus, 111 n.

Zamicrus, 184

Zapus, 486

Zui-rhachis, 384

Zebra, 244 ; Burchell's, 245 ; Common,

244 ; Grevy's, 244
Zenkerella, 463 n.
Zeuglodon, 384
Zeuglodontidae, 384
Ziphius, 367, 370 ; Z. carirostris, 370 ;

Z. novae zelandiae, 370
Zoological regions, 84
Zorilla, 438

THE END

Printed by R. & R. CLARK. LIMITED, Edinburgh.

THE CAMBRIDGE NATURAL HISTORY

Edited by S. F. HARMER, Sc.D., F.R.S., Fellow of King's College,
Cambridge, Superintendent of the University Museum of Zoology,
and A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge,
University Lecturer on the Morphology of Invertebrates.

To be compkted in Ten Volumes. Svo. Price 17s. net each.
[Now Ready.]

MAMMALS

VOLUME X
By F. E. BEDDARD, M.A. Oxon., F.R.S., Prosector to the Zoological Society,

WORMS, LEECHES, ETC.

VOLUME II
Second Impression

Flat Worms. By F. W. GAMBLE, M.Sc. Viet., Owens College.— Nemertines.
By Miss L. SHELDON, Newnham College, Cambridge. — Thread-worms, etc.
By A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge. — Rotifers.
By MARCUS HARTOG, M.A., Trinity College, Cambridge, D.Sc.Lond., Professor
of Natural History in the Queen's College, Cork. — Polychaet Worms. By
W. BLAXLAND BENHAM, D.Sc., Hon. M.A. Oxon., Professor of Biology in
the University of Otago. — Earth-worms and Leeches. By F. E. BEDDARD,
M.A. Oxon., F.R.S., Prosector to the Zoological Society, London. — Gephyrea
etc. By A. E. SHIPLEY, M.A., Fellow of Christ's College, Cambridge. —
Polyzoa. By S. F. HARMER, Sc. D. , F. R. S. , Fellow of King's College, Cambridge.

SHELLS

VOLUME III

Molluscs and Brachiopods
By the Rev. A. H. COOKE, M.A., A. E. SHIPLEY, M.A., and F. R. C. REED, M.A.

TIMES. — "There are very many, not only among educated people who take
an interest in science, but even among specialists, who will welcome a work of
reasonable compass and handy form containing a trustworthy treatment of the
various departments of Natural History by men who are familiar with, and
competent to deal with, the latest results of scientific research. Altogether, to
judge from this first volume, the Cambridge Natural History promises to fulfil all
the expectations that its prospectus holds out."

INSECTS AND CENTIPEDES

VOLUME V
Second Impression

Peripatus. By ADAM SEDGWICK, M.A., F.R.S.— Myriapods. ByF. G.SINCLAIR,
M.A.— Insects. Part I. By DAVID SHARP, M.A. Cantab., M.B. Edin., F.R.S.
FIELD. — "Although written for the student and the specialist, the book is not
the less adapted to all intelligent readers who wish to make themselves thoroughly
acquainted with the habits, structure, and the modern classification of the animals
of which it treats. To such it cannot be recommended too strongly. "

ENTOMOLOGIST'S MONTHLY MAGAZINE.— " We venture to think the
work will be found indispensable to all who seek to extend their general knowledge
beyond the narrowing influence of exclusive attention to certain orders or groups,
and that it will take a high position in ' The Cambridge Natural History' series."

INSECTS— PART II

VOLUME VI

Second Impression
Hymenoptera continued (Tubulifera and Aculeata), Coleoptera, Strepsiptera,

Lepidoptera, Diptera, Aphaniptera, Thysanoptera, Hemiptera, Anoplura.

By DAVID SHARP, M.A., M.B., F.R.S.

SATURDAY REVIEW. — "Dr. Sharp's treatment is altogether worthy of the
series and of his own high scientific reputation. But in a work of this sort it is
not only necessary that information should be accurate, but also that it shall be
presented to the eye, so far as illustrations and printing are concerned, in such a
way as to render its matter as easily intelligible as possible, and readily usable
for purposes of refe. anee. Under both these heads we have nothing but commenda-
tion for Mr. Sharp's treatise. The illustrations are indeed beautiful, and the use
of the heavy type for the headings of the various sections and leading paragraphs
materially helps the reader in the progress of his study. Certainly this is a book
that should be in every entomologist's library."

AMPHIBIA AND REPTILES

VOLUME VIII

By HANS GADOW, M.A., Ph.D., F.R.S.

FIELD. — "The work is worthy of the series in which it appears, and we can-
not give it higher praise."

SCIENCE GOSSIP.— " More than maintains the high scientific reputation of
this series. The herpetologists, or students of the Amphibia and Reptiles, have now
a standard work of the highest class."

BIRDS

VOLUME IX.— Second Impression
By A. H. EVANS, M.A., Clare College, Cambridge. With numerous Illustrations

by G. E. LODGE.

IBIS. — •" Mr. Evans has produced a book full of concentrated essence of informa-
tion on birds, especially as regards their outer structure and habits, and one that
we can cordially recommend as a work of reference to all students of ornithology."

The following volumes which are in the Press or in active preparation will

complete the series : —

VOLUME I

Protozoa, MARCUS HARTOG, M.A., D.Sc., Trinity College (Professor of Natural
History in the Queen's College, Cork) ; Sponges, W. J. SOLLAS, Sc.D., F.R.S.,
St. John's College (Professor of Geology in the University of Oxford) ; Jelly-
fish, Sea-Anemones, etc., S. J. HICKSON, M.A., F.R.S., Downing College
(Beyer Professor of Zoology in The Owens College, Manchester) ; Star-fish,
Sea-Urchins, etc., E. \V. MACBRIDE, M.A., St. John's College (Professor
of Zoology, McGill University, Montreal). [In active preparation.

VOLUME IV

Spiders, Mites, etc.,C. WARBURTON, M. A., Christ's College (Zoologist to the Royal
Agricultural Society) ; Scorpions, Trilobites, etc., M. LAURIE, B.A., King's
College, D.Sc. Edinb. (Professor of Zoology in St. Mungo's College. Glasgow);
Pycnogonids, etc., D'ARCY W. THOMPSON, C.B., M.A., Trinity College
(Professor of Zoology in University College, Dundee) ; Crustacea, W. F. R.
WELDON, M.A., F.R.S., St. Jolin's College (Linacre Professor of Comparative
Anatomy in the University of Oxford). [In active preparation.

VOLUME VII

Balanoglossus, etc., S. F. HARMER, Sc.D., F.R.S., King's College ; Ascidians
and Amphioxus, W. A. HERDMAN, D.Sc. Lond., F.R.S. (Professor of Natural
History in University College, Liverpool); Fishes, T. W. BRIDGE, Sc.D.,
Trinity College (Professor of Zoology in Birmingham University), and G. A.
BOCLENGER, F.R.S. [Inthe Press.

MACMILLAN AND CO., LTD., LONDON.

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