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Digitized by the Internet Archive
in 2007 with funding from
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http://www.archive.org/details/cambridgenatural10harmuoft
EWE:
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
VOLUME X
PE
MAMMALIA
By FRANK EVERS BEDDARD, M.A. (Oxon.), F.R.S. Vice-
Secretary and Prosector of the Zoological Society of
London.
London
ENC hibe IAN CAND: CO; Liver ED
NEW YORK: THE MACMILLAN COMPANY
1902
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 pereyng lynx, the lufar vnicorne
The fery tigere, full of felonye,
The dromydare, the standar oliphant.”
From The Kingis Quhair, JAMus I. (of Scotland).
FEB 15 1994
%, Ay
—eRsiry oF \OS
PREFACKH
INASMUCH as Sir W. 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
Rodentia 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 oncé 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 Handbuch 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.
Lonnon, February 28, 1902.
CONTENTS
PREFACE
SCHEME OF THE CLASSIFICATION ADOPTED IN THIS Book
CHAPTER I
INTRODUCTORY
CHAPTER II
STRUCTURE AND PRESENT DISTRIBUTION OF THE MAMMALIA
CHAPTER III
Tue PossipLE FORERUNNERS OF THE MAMMALIA
CHAPTER IV
Tue DAWN oF MAMMALIAN LIFE
CHAPTER V
Tue EXxIsTING ORDERS OF MAMMALS: PROTOTHERIA—MONOTREMATA
CHAPTER VI
INTRODUCTION TO THE Sup-CLAsSsS EUTHERIA
~
CHAPTER VII
EUTHERIA—MARSUPIALIA
PAGE
90
96
116
122
viii CONTENTS
CHAR TER “Vit
PAGE
EpENTATA—GANODONTA : Z : : , ; : ; : 4 AKI
@REACR ER aelexe
UNGULATA —CONDYLARTHRA — AMBLYPODA — ANCYLOPODA — TYPOTHERIA—
ToxoDONTIA—PROBOSCIDEA—HYRACOIDEA : : : é a 108)
CEPAGE AM Rex
UNGULATA (CONTINUED)—PERISSODACTYLA (ODD-TOED UNGULATES)—LI?0-
PTERNA . : F 3 3 ; 5 ; 2 : ; : 5 BY)
CHAPTER XI
UNGULATA (CONTINUED)—ARTIODACTYLA (EVEN-TOED UNGULATES)—SIRENIA 269
CHAPTER XII
CrraAcEA—WHALES AND DOLPHINS : : F ‘ : F ; 5 By?)
CHAPTER, Sot
CARNIVORA—FISSIPEDIA : ; ‘ ; ; 4 : : : + 3886
CHAPTER: XIV
CARNIVORA (CONTINUED)—PINNIPEDIA (SEALS AND WALRUSES)—CREODONTA 446
CHAPTER 2¥
RoDENTIA—TILLODONTIA ; , . 2 . : : : ? . 458
CHAPTER XVI
INSECTIVORA—CHIROPTERA . : 4 ; : ; : : : - 508
CHAPTER XVII
PRIMATES d 3 , ‘ , ; : : . : : ; 5 yas:
INDEX 5 : : : : : y i d : ‘ : 7 OOM
CHAPTER I
INTRODUCTORY
THE Mammalia form a group of vertebrated animals 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 gg B
N
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 crease in bulk accompanies specialis-
ation of structure. The huge Dinocerata when compared with
the ancestral Pantolambda 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-
I ‘ DE HENTTIONS OH EVE GLASS 3
acters might be 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 defined :—
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.
Cervical 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 by 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.’
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 have 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
' The degeneration of the hind-limb 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.”
CHAT GHEE: I
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 1s 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 Mai-
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 Reptiles, in which, however, the differ-
ences between the fore- and hind-limbs are less marked, indicat-
6 INTELLECT CHAP.
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 motionlessness, so familiar to
everybody in such a creature as the Crocodile, are unknown among
the more typical Mammala 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, 1s 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
II HAIR 7.
reduced to as few as two bristles on the lips. The term “ hair,”
however, 1s apt to be somewhat loosely applied; it has been
made use of to describe, for example, the slender processes of the
Sch
—~\
Fic. 1.—A, Section of human skin. Oo,
Dermis ; D, sebaceous glands; /, fat
in dermis ; G, vessels in dermis; GP,
vascular papillae ; H, hair ; NV, nerves
in dermis ; VP, nervous papillae ; Sc,
horny layer of epidermis ; SD, sweat
gland ; SD!, duct of sweat gland ; SJ/,
Malpighian layer. B, Longitudinal sec-
tion through a hair (diagrammatic). A),
Band of muscular fibres inserted into the
hair-follicle ; Co, corium (dermis) ; /,
external longitudinal ; /, internal cir-
cular, fibrous layer of follicle; /'%t, fatty
tissue in the dermis; GH, hyaline
membrane between the root-sheath and
the follicle; HBD, sebaceous gland ;
HP, hair-papilla with vessels in its in-
terior ; J/, medullary substance (pith)
of the hair; O, cuticle of root-sheath ;
FR, cortical layer ; Sc, horny layer of
epidermis ; Sch, Hair shaft ; SJ, Mal-
pighian layer of epidermis ; WS, WS},
outer and inner layers of root-sheath. (From Wiedersheim’s Comparative
Anatamy.)
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
8 DEVELOPMENT OF HAIR CHAP.
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
SUSE 7-200 homologue of une 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-
Fic. 2.—Four diagrams of stages in the development Jags feathers. Still
ofahair. A, Earliest stage in one of those mammals i :
in which the dermal papilla appears first ; B, C, D, later the knob of €pl-
three stages in the development of the hair in the dermic cells becomes
human embryo. 6/6, Hair-bulb ; crn, horny layer ;
of the epidermis ; fol/, hair-follicle; grm, hair- depressed into a tubu-
germ; h, hair, in D, projecting on the sur- Jar structure, which is
face ; muc, Malpighian layer of epidermis; pp, ~~ ; ?
dermal papilla ; seb, developing sebaceous glands; lined with cells also
sh.l, He inner and outer root-sheaths. (After digniaed er onic
Hertwig.) An
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. Meijerle’ has lately described in some detail the parti-
1 «Uber die Haare der Siiugethiere,”” Morph. Jahrb. xxi. 1894, is cules
II HAIR AND UNDER-FUR 9
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 ;+ i
in Ursus 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. Alhed 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. Meijerie’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 seales of the Armadillo, and which has led him to the view
that the ancestors of mammals were scaly creatures
supported by Professor Max Weber,’ and not in itself unreason-
able when we consider the numerous points of affinity between
the primitive Mammalia and certain extinct forms of reptiles.”
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 Ofaria wrsina 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
fo)
of hairs seems to be responsible for the scales of the Manis
a view also
1 << Bemerkungen iiber den Ursprung der Haare,” Anat. Anz. 1893, p. 415.
? See for this matter, p. 90. Dr. Bonavia has recently advanced (Studies 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, eg. 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-lmbs 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 flat 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
chiefly 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
II COLORATION Il
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. Next 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, e.g. the
Gorilla, or in the presence of tusks, e.g. various Boars, or of horns, as
in the Deer, ete. 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 Puma 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
[T2 CLYED 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 Mammalha 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
mamunals, with but a few exceptions. The structures that we are
now concerned with are agglomerations of these glands. The
mamiary 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
fo 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 J/yrmecobius.'
On the skin of the anterior part of the chest, just in front of the
! Proc. Zool. Soc. 1887, p. 527.
iI GLANDS OF TIIE 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 Didelphys 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 gregarious 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 Rhinoceros, 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, Rudi-
ments of this structure have, however, been recently discovered
in the higher inamimals, and, 2s Dr. Klaatsch’ 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 Uber Marsupialrudimente bei Placentaliern,” Morph. Jahrb. xx. 1893, p.
276,
Il POUCH I
un
In 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 asa temporary
harbour of refuge: from the pouch of female Kangaroos at the
Zoological Gardens may frequently be observed to protrude the tail
Fic. 3.—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 the secretion flows. 4.2m, Pouch ;
el, cloaca; g.m, groups of mammary glands. (From Wiedersheim’s Com ti
Anatomy, after 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 Monotremata (Gn Fehidna) there is a deep fold of
the skin which lodges the unhatched egg, and into which the
Inammary 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-
16 MARSUPIUM AND MAMMARY POUCH CHAP.
supials.' 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 Hehidna are to be looked upon as the equi-
valent of the commencing teat of the higher mammal.’ 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.’ 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 Lehidna. 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 ‘‘ Uber die Entstehung der Saugetiere, ”
Biol. Centraibl. viii. 1889, p. 8.
2. See Gegenbaur’s Llements of Comp. Anat. - Transl. by Bell, 1878, p. 421.
3 Uber die Beziehungen zwischen Mammartasche u. Marsupium,” Morph.
t=) r]
Jahrb. xvii. 1891, p. 483.
II POUCH OF MARSUPIALS ]
Py
‘
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
undifferentiated 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
Phascologale 1t is only pre-
sent in rudiment, and in
Myrmecobius it is entirely
obsolete. In the American
Fria. 4.—Diagram of the development of the nipple
Opossums the state of the (in vertical section). A, Indifferent stage, gland-
pouch is variable. “Gener- ular area flat ; B, elevation of the glandular oe
: with the nipple ; C, elevation of the periphery
ally absen t, sometimes of the glandular area into the false teat. a,
Periphery of the glandular area; 6, glandular
merely, composed of two area; gl, glands. (From Gegenbaur.)
lateral folds of skin separ-
ate at each end, rarely complete,” is Mr. Thomas’ summary in his
definition of the family Didelphyidae.t 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 Cc
18 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 ursinus 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 marsupium 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 marsupium 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 certaim 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. 1 6),7
is a secondary teat; hence there is no difficulty in the com-
parison from this point of view. A pouch contaiming 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 marsupium. 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 catia 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 lLmbs. 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 Vertebrata, 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 CHAP.
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.
Fia. 5.—Anterior surface of Fic. 6.—Side view of first
Human thoracic vertebra
(fourth). x2. az, Anterior
zygapophysis ; ¢, body or
centrum; J, lamina, and
p, pedicle, of the neural
arch ; 7c, neural canal ; ¢,
transverse process. (From
Flower’s Osteology of the
Mammalia. )
lumbar vertebra of Dog
(Canis familiaris). x 2.
a, Anapophysis ; az, an-
terior zygapophysis ; 1,
metapophysis ; pz, pos-
terior zygapophysis ;_ s,
spinous process ; ¢, 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. and the proximal ‘end of the left uterus is shown in
with a number of ioneeiainall section. jl.¢, Fallopian tube ; 71.1’, its peri-
fanciful names — toneal aperture ; /.ut, left uterus ; ut’, left os uteri ;
ov, ovary ; r.ut, right uterus ; 7.ut’, right os uteri; s,
the morsus diaboli. 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 Rat! 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.
? Beddard, Proc. Zool. Soc. 1900, p. 667.
TA: OVIDUCTS OF MARSUPIALS CHAP.
FiG. 48.—Female urino- genital
apparatus of various Marsu-
pials. A, Didelphys dorsigera
(young) ; B, Trichosurus ;
C, Phascolomys wombat. B,
Urinary bladder; (Cl,
“cloaca”; Fim, fimbriae ;
g, Clitoris ; N, kidney ; Od,
Fallopian tube; Of, aper-
ture of Fallopian tube; Ov,
ovary ; 7, rectum; Sp, sep-
tum dividing vagina; Sug,
urino-genital sinus; U7,
ureter; Ué, uterus; U¢?,
opening of the uterus into
the median vagina (VgB) ;
Vy, lateral vagina ; Vg’, its
opening into the urino-genital
sinus; + (in B), point of
approximation of uteri; + (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,
II THE BRAIN 7%
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
fimbriated 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 2 epicoracoid ;
epist, — epi-
sternum ;
ep.pb, epipubis ; fb,
fibula ; fem, femur ;
JSor.mag,foramenmag-
num; glen, glenoid a
cavity of shoulder-
joint ; glen, glenoid
cavity for mandible ;
hum, humerus;
in.cond, inner con-
dyle of humerus ;
inf.orb.for, points to
position of infra-orbi-
tal foramen ; 777 7.p70c,-
\/ G ‘J oe
ast |4 aS
cale Ma SS) a \E inferior processes of
ses ie CL DS PON, ' }-acelars caudal _ vertebrae :
scaph (Bg cod = “A int.rbs, intermediate
en£.curn lin cl.cun a ir "oH vn ribs ; ¢sch, ischium ;
metaltLT lalv. 7 S \\ mag, magnum of car-
\ \Y Ai i pus ; maz, maxilla ;
: \\\ CM H//}] max.for, maxillary
XS f yj foramen ; metat.T,
rh first metatarsal ;
Cs metat.V, fifth meta-
ae tarsal; vas.cart, nasal
% cartilage ; obt, obtu-
rator foramen ; ol,
olecranon ; ovt.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 ; vad, radius ; scap, scapula ; scaph, scaphoid of tarsus ;
scaph.dun, scapho-lunar ; ses, sesamoid bones of wrist and ankle ; sp, tarsal horny spur ;
sq, Squamosal ; 7/2, tibia ; trd, trapezoid ; trm, trapezium ; fym.c, tympanic cavity ;
uln, Wina ; unc, unciform ; vom, vomer ; ~, dumb-bell shaped bone: zyg, zygomatic
arch ; /-V, digits oY manus ; V, foramen for fifth nerve. (From Parker's Zoology.)
CHAP. V DR. SEMON’S OBSERVATIONS Hi
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
fifty 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, /n 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 molluse 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.
CHAP Val
INTRODUCTION TO THE SUB-CLASS EUTHERIA
SuB-CLASS I1—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-
rhynchus and Eehidna from all other mammals. In his well-
known memoir upon the arrangement of the Mammalia,’ 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 Prototherian type are (1) the presence of epipubes ;
(2) the small corpus callosum; (3) the absence of an allantoic
placenta.
The last of these can be dismissed, im consequence of the
recent discovery of an allantoic placenta in Perameles. The first
character is apparently a valid distinction between the Marsupials
l Proc. Zool. Soc. 1880, p. 649.
CHAP. VI EU THERIA—CORPUS CALLOSUM Val bi 7/
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. 77) being small,
that seems to be not more than a slight difference of degree.!
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 ee aa
single successional tooth. It
seems at the present moment (tT
to be fairly clear that Marsu- CN
pials have a milk dentition i= Ca)
like other Eutherians, but
that only one of these teeth,
the fourth premolar, comes
to functional maturity. That emam |opl\ tubolf
it is really one of a complete wentsa coljforn
milk series is evidenced by Fie. 57.—Brain of Lchidna aculeata ; sagittal
the fact that this tooth is section. ant.com, Anterior commissure ;
cbl, cerebellum ;— ¢.mam, corpus mammil-
differentiated contemporane- lare ; col.forn, column of the fornix ; c.qu,
ously with another series corpora quadrigemina : gang. hab, ganglion
habenulare ; hip.com, hippocampal com-
formerly held to belong to missure ; med, medulla oblongata ; mid.com,
Pay ee ar C middle commissure ; o/f, olfactory lobe ;
the so-called prelacteal denti- opt, optic chiasma; tub.olf, tuberculum
tion.” There still remains, of olfactorium ; vent. 3, third ventricle. (From
course, the actual fact that Parker and Haswell’s Zoology.)
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 ins many other mammals admittedly Eutherian.
Huxley regarded the pecuharities in the reproductive organs
* Moreover, the ‘‘ corpus callosum and the anterior commissure... in... Hrin-
aceus and Dasypus are almost Monotreme-like.”’
2 See Wilson and Hill, Quart. J. Mier. Sci. xxxix. 1899, p. 427.
118 MOLARS OF EUTHERIA : CHAP.
of the Marsupials as “singularly specialised characters,” im no
way intermediate in character. This view apples 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
dividing them from
the higher mammals
than was formerly the
case. ‘The total denti-
tion of the group is
on the whole com-
posed of more numer-
ous individual teeth
than in the typical
cmam vents Eutheria; but we have
Fic. 58.—Sagittal section of brain of Rock Wallaby exceptions like the
(Petrogale penicillata). ant.com, Anterior commis- yy7},.7,. wees
sure ; cb/, cerebellum ; ¢.mam, corpus mammillare ; Whales, the Arma-
e.qv, corpora quadrigemina ; erwr, crura cerebri dillo Priodontes, and
epi, epiphysis, with the posterior commissure im- fei Maine ; ;
mediately behind it ; fmon, position of foramen of 1€ Manatee, ol
Monro ; hip.com, hippocampal commissure, consist- better, because free
ing here of two layers continuous behind at the oe ip
spleneium, somewhat divergent in front where the from the suspicion of
Segums lucidum extends between them ; hypo, hypo- secondary multiplica-
physis ; med, medulla oblongata ; mid.com, middle — ,
commissure ; olf, olfactory lobe ; opt, optic chiasma ; tion, Otocyon and ocea-
Cae ventricle. (From Parker and Haswell’s sionally (according to
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.
it is at least double the width of the second or third finger; the
pollex is very slender. _ In the little Cycloturus 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.
Cycloturus is by far the smallest of the Anteaters. It has
170 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 Bradypus and
Fic. 96.—Unau, or Two-toed Sloth. Choloepus didactylus. x 4.
(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
VIII PROTECTIVE COLOUR OF SLOTH el
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 hoffmanni (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." 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
fT
Cer
Fic. 97.—Skull of Three-toed Sloth. Bradypus tridactylus. Wateral view. j7, Frontal ;
ju, jugal ; ler, lachrymal ; maz, maxilla ; nas, nasal; par, parietal ; s.oc, swpra-
occipital ; ty, tympanic. (From Parker and Haswell’s Zoology.)
out by Dr. Siemann,” who observed that a species occurring in
Nicaragua “has almost exactly the same ereyish-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
uumicry between the sloth’s hair and the 7%llandsia, 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.
" 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.
NH 2 OSTEOLOGY OF SLOTH CHAP.
The zygoma is incomplete, though the part connected with the
frontal has a strong downward process like that found in Diprotodon
Bradypus tridactylus, (After de Blainville. )
Skeleton of Three-toed Sloth.
98.
Fic.
and some other mammals. There is, moreover, a process from the
squamosal, though it does not reach the anterior part and thus
VIII TEP OR ARMADIMIE@OS 7
ios)
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 pecuharity of the Sloths is the enormous number of dorsal
vertebrae. There are twenty-three of these in Choloepus 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
Choloepus, shows another structural feature which does not bear
out this conclusion. The trapezoid and the os magnum of the carpus
are united, while in Choloepus 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 Chlamydophorus. 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
' This name is written ‘‘ Prionodos”” by Gray, which might lead to a confusion
with the Carnivore Prionodon.
yA 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' The premaxillaries are
Fie. 99.—Skull of Armadillo. Dasypus sewcinctus. 3%. ex.oc, Exoccipital ; 77,
frontal ; maz, maxilla; nas, nasal; pai, parietal; pert, periotic ; p.max, pre-
maxilla ; s.oc, supraoccipital ; sq, squamosal; ¢y, tympanic. (From Parker and
Haswell’s Zoology.)
small, but are larger in Dasypus than in 7atusia. 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 Zatusia (but not in Dasypus) the “short thick pterygoids add
somewhat to the hard palate.” This is clearly a beginning or :
remnant of the quite crocodilian character of the palate of
Myrmecophaga. 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
' For the anatomy of several forms, see Garrod, Proc. Zool. Soc. 1878, p. 222,
who quotes other memoirs.
vIn 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
Fria. 100.—Bones of the right Fic. 101.—Bones of the manus of
manus of the Hairy Armadillo. the Great Armadillo. Priodon
Dasypus villosus. x 3. Gs giganteus. x4. a, An acces-
Cuneiform ; /, lunar ; m, mag- sory carpal ossicle in front of
num ; p, pisiform ; £&, radius ; the pisiform, which is not seen
s, seaphoid ; ¢d, trapezoid ; tm, in the figure. Other letters as
trapezium ; wv, unciform; U, in Fig. 100. (From Flower’s
ulna; J-V, digits. (From Osteology.)
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 Tolypeutes the first digit has vanished ;
on the other hand, in Priodon, the fifth has become rudimentary
176 PGE - Gls GOAN Ome al. CHAP.
and the third enormously enlarged. This latter fact recalls
the arrangement characteristic of Myrmecophaga. The pelvis is
greatly attached by the
ischium to the verte-
bral: coltmaney aie
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, ete.
The little Pichi-
chago (or, more cor-
Fra. 102.—Pelvis and sacrum of Armadillo. Dasypus ,,, oe ats .
5 ene ie o si eG A sNny - c1eg
sexcinctus. ac, Acetabulum; 7/, ilium ; isch, ischium ; rectly, Pichy c1ego),
obt,for, obturator-foramen ; pect.tub, pectineal tuber- Chlamydophorus, which
cle; pub, pubis. (From Parker and Haswell’s ike Geos fo alos &
Zoology.) only grows to abou
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 arma-
ture is continuous and nowhere ringed.
The genus Tolypeutes, of which the best-known species 1s 7.
tricinctus, the Apar (there are two other species in the genus), can
roll itself up into a ball like the Puill-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-
VII 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-banded Armadillo or Apar. Tolypeutes tricinctus. x 4.
The VPeludo, Dasypus sexcinctus, is, like other Armadillos, an
x4. (After Vogt and Specht.)
Fic. 104.—Peludo Armadillo. Dasypus sexcinctus.
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 novemeincta 18 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.!
The genus Yenurus contains several species, the best known of
which is inaptly named 1. wnicinctus. 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 Vatusia, which 1s 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 alhance between the genera Yenuwrus and Dasypus
and their difference from Zatusia, 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
Dasypus and Tolypeutes.’ 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. minutus
' Flower, Proc. Zool. Soc. 1886, p. 419.
VIII GIANT ARMADILLO 179
5:1, with a large intestine of no less than 7 feet; D. vellerosus
4°35 and 66.
Priodon is the giant of its race. This Armadillo may
reach a leneth 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! S. bruneti, 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. Peltephilus
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
Hutatus 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 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 Ihinoceros.
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. Mus. vii. 1871, p. 177.
2 See especially Lydekker, An. Mus. La Plata, Pal. Arg. iii. 1894.
180 EXTINCT XZWVARTHRA 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
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, Scelidotheriwm
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.’
Fic. 105.—Mylodon robustus. (Restoration, after Owen.)
1 Dr. Moreno and Mr. A. Smith Woodward in Proc. Zool. Soc. 1899, p. 144 ;
Wiss. Ergeb. Schwed. Exped. Magellansland. li. 1899, p. 149.
VIII NEOMYLODON OR GLOSSOTHERIUM ISI
The well-known naturalist of La Plata, Sefior Moreno, engaged
in studies connected with the political boundary line between
Chil 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. Lonnberg 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 Glossotheriwm. “Ancient chroniclers inform us
that the indigenous inhabitants recorded the existence of a
182 HAIR OF NEOMYLODON CHAP.
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 Rio 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 beheving
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 “ Veomylodon ”
must be referred to Glossotherium or Grypotheriwm, as it 1s
sometimes termed. This skull is perforated on the roof in such
a way as could only have been effected (an 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 Glossotheriwm is, like
that of other extinct “ Ground-sloths” (e.g. Mylodon), filled with
small and irregular ossicles. But in JMJylodon, 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 MZylodon and the existing Armadillo (Dasypus).”
Now Gilossotherium 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 Glossotheriwm would be also inter-
mediate, or at least like that of one of the two genera Myrme-
cophaga and Bradypus. But microscopical investigation has
VIII MEGATHERIUM Kos
negatived this supposition. It has shown that the Armadillos
are in this matter the nearest relatives of Glossotheriwm. 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. Glossotheriwm 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-lke
tooth, in both jaws.
Megalonyx and its allies are sometimes placed in a distinct
family, Megalonychidae. JJegalonyx 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. the inner).
184 ARMOURED EDENTATES CHAP.
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 4.
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
Fic. 106.—Glyptodon clavipes. x js. (After Owen.)
1
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
VIII 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
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-limhs,
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.
Propalaeohoplophorus is, unlike the great Armadillos that we
have hithérto 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
186 PELTEPHILUS CHAP.
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. Pe/te-
philus differs from other Armadillos in having teeth in the front
of the jaws. The total number of teeth is twenty-eight, 7.e. seven
in each half of each jaw.
SuB-OrDER 2. NOMARTHRA.
As already explained, the Old-World Edentates differ from
the New-World forms in having normal dorsal vertebrae, that 1s
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.
VIII AARD VARK 187
This group contains but one family, the Orycteropodidae, of
which there is but a single genus.
The Aard Vark (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.
ANG
Fic. 107. —Aard Vark, or Cape Anteater. Orycteropus capensis. x 3}
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. 162, the Aard Vark 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 twe
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 Ray Myliobates. According to Mr. Oldfield
188 DEE TE OF AAR DRVARK CHAP.
Thomas! 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
Fic. 108.—Section of lower jaw with the teeth of Orycteropus. x2. (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,
O. capensis, 18 more hairy than the northern, O. aethiopicus.
O. gaudryi is a Pliocene species from the Island of Samos and
from Persia, described by Dr. Forsyth Major and Dr. Andrews.’
It closely resembles the existing O. aethiopicus.
Of the Scaly Anteaters, Group SguaMATA 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.” The external form
of these animals is fairly well known, the remarkable scales dis-
tinguishing the Pangolins from other animals. Between the
scales he 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,’ 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. 198.
4 Weber, Zool. Ergebnisse einer Reise in Niederl. Ost Indien, 1892. See also
Romer, in Jen. Zeitschr. xxxi. 1896, p. 604, and Reh, cbid. xxx. 1895, p. 187.
Vl THE PANGOLINS 189
tures, as he does the scales of other maminals, 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. Jetia 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 thé 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 lkenesses which Janis shows
I90 ANCIENT, EDENTATES CHAP.
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 Janis, is
small in Myrmecophaga ; the clavicle is absent and again small or
rudimentary in the Anteaters; it is large in other Edentates.
Fie. 109.—Manis. Janis gigantea. x 4)5.
The third trochanter is absent, as in IMJyrmecophaga (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 Janis.
The facts are not a httle remarkable.
Order III. GANODONTA.'
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.
Vill PSITTACOTHERIUM IOI
logical order. The following genera are included by Wortman in
his family Stylinodontidae.
The earliest type of the Ganodonta is the genus Hemiganus,
with but one species, H. ofariidens. 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 Kocene 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 lke that of Hemz-
ganus—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 ot
Ground Sloth, Megalonyx (but it must be remembered that the
same characteristic is not unknown in Rodents). 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,
192 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 Hemiganus having
vanished in this later form. The tooth pattern of the molars is
hike that of Hemiganus. 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-lmbs 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 Reodent-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 Psittacothervwm ; they are, when unworn,
quadricuspidate, with accessory cusps; when more worn, the teeth
VII 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. SS. 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 lke that of Psittacotherium.
Tn 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—OW YCHODECTES CHAP. VIII
facial portion, and is thus more hke 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.
Cie AGE NBR: EX
UNGULATA——-CONDYLARTHRA——AMBLYPODA—ANCYLOPODA—
TY POTHERIA——_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 peculharities 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 Hyraz. There is, too, as we pass from the ancient
types to the modern, a gradual perfection of the hmbs as running
196 PHENACODUS CHAP.
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* is highly
remarkable. The tail
too, an organ which is
long in some of the
early forms, gets short
in their modern deriva-
tives. .
Coupled with the in-
creasing perfection of the
foot as an organ used
merely for the support
of the body, certain in-
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-
(After Osborn. )
1
12
x
Phenacodus primaevus.
Fia. 110.—An early Ungulate.
' This creature is, however, sometimes referred to the neighbourhood of the
Rodents.
Pix 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
F
Fic. 111.—Series of metacarpals and metatarsals of Camelidae, to show secular and
progressive increase in size. From left to right the species are Protylopus
petersont, Poebrotherium labiatum, Gomphotheriwm sternbergi, Procamelus occt-
dentalis. F, Fore-foot ; H, hind-foot ; III, IV, third and fourth metapodials.
(After Wortman.)
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 CHAP.
Matthew.’ 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 hving 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
Fic. 112.—Bones of the manus A, of the Indian Elephant, Hlephas indicus. x
B, of the Cape Hyrax, Hyrax capensis. x1. c, Cuneiform; cc, centrale; /
lunar; m, magnum ; p, pisiform; R, radius ; ¢d, trapezoid ; tm, trapezium; s
scaphoid ; w, 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 flatter crushing sur-
faces exhibited by the lophodont teeth of later Ungulates. Strong
1 Bull. Amer. Mus. Nat. Hist. ix. 1897, p. 321.
IX 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
Fic. 113.—Bones of the manus A, of Rhinoceros, Rhinoceros sumatrensis. x. B, of
Pig, Sus scrofa. x}. 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 néarer than in quite
early Tertiary times. The order Condylarthra seems to be the
starting-point of both these sub-divisions. Huprotogonia has been
considered to be an ancestor of the Perissodactyle branch, and
Protogonodon or Protoselene of the Artiodactyla. If this be true,
200 HORNS CHAP.
the likenesses which 7Z%tanotherium 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
ageressive 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, 7.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
IX HORNS AS A SEXUAL CHARACTER 201
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 Antzlocapra, 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, 7.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,’ 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.
ZO) CONDYLARTHRA CHAP.
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, Sumotherium, of which the male alone had a pair
of short horns, while the skull of the female was entirely hornless ;
the modern Giraffu, 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 Diceratheriwm apparently inherited and
unproved 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.
SUBZORDER Ly CONDYLARTHERA:
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
1X PHENACODUS AND ‘THE CREODONTS 203
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 Kocene 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 le 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,’ 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 Kuropean 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.
2 Marsh, Amer. Journ. Sci. xliii. 1892, p-. 447.
204 CONDYLARTHRA—PROTOGONIA 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 Luprotogonia, or Protogonia ' as it has been called. The best-
known species is HL. 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 16 or 17 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 1s 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 Huprotogonia 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. Nat. Hist. ix. 1897, p. 303.
IX CEPA TOn TARE YUNG ULATES 205
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
Meniscotherium, tor 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.
“ Astragal 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 Uneulates. 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 1s 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-OrRDER 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-linb, are well developed. The bones
206 AMBLYPODA—_DINOCERAS AND ITS ALLIES CHAP.
in the carpus are alternating in position. The toes are five in
both feet, and are very short. There is a hint of commencing
“ pyerissodactylism” 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
oT ee oe
le
Fic. 114.—Skull of Protolambda bathmodon. x. e.a.m, External auditory meatus ;
m, mastoid ; m.f, mastoid foramen. (After Osborn. )
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.". 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
! Or perhaps rather to the primitive Ungulates Condylarthra. It is especially
compared with Periptychus of that group.
1X COR VYPHODON ZOV,
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-lmbs, and it had a long tail. It was
apparently plantigrade, and would have had not a little hkeness
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 Coryphodon, 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.' 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 Huprotogonia. 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 Huprotogonia 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
R. Owen CT 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. Amer.
Mus. Nat. Hist. x. 1898, p. 169.
oly
temains 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, 7. elatum ; lower figure, 7. platyceras. (After Osborn.)
266 : HORNS OF 777TANOTHERIUM 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-lhmb is un-
doubtedly Perissodactyle in the arrangement of its component
parts, the fore-limb shows a hint of an Artiodactyle mode of
structure. This hmb 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, 7. 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 7itanotheriwm 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. '
Palaeosyops is somewhat lke a Tapir in build, the skull
especially resembling that of the Tapir. As in TVitanotheriwm
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. Mus. Nat. Hist. vii. 1895,
>
p. 333, and Osborn, ibid. viii. 1896, p. 157.
XK CAMEL-LIKE PERISSODACTYLES ZOY.
three-toed. It was 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 Zitanotheriwm.
Related to Palaeosyops is another primitive Titanothere, the
genus Velmatotherium. 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 7elmatotherium, 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 7. cornutwm the horns are chiefly borne upon the very
long nasals, whose size contrasts with the same bones in the
more highly-developed Zitanotherium. It appears to be quite
possible that Titanothertwm was evolved from the genus
Telmatotherium.' ¢
SuUB-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 Macrauchenia 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 Burmeister’ 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-lke
than Horse-lhke. On the other hand, the proportions of femur
to tibia are more Horse-lhke. 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 TZhoatherium,
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, Protorotheriwm, we have
the two lateral toes diminishing just as in Anchithervum.
1 N. Acta Acad. Caes. Leop. Car. xxvii. 1885, p. 238.
. > |
CHAPTER XI
UNGULATA (continued)—ARTIODACTYLA (EVEN-TOED
UNGULATES )—SIRENIA
SuB-OrpEer 10. ARTIODACTYLA.
Tue Artiodactyle or “Even-toed” Ungulates are to be dis-
VY i
Fic. 138.—Bones of the Manus—A, of Pig (Sus scrofa). 4%. B, of Red Deer (Cervus
elaphus). x%. (©, of Camel (Camelus bactrianus). x. c, Cuneiform ; /, lunar ;
m, magnum ; m7, m°, second and fifth metacarpals; R, radius; s, scaphoid ; ¢d,
trapezoid ; wv, unciform; U, ulna; JJ-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
ZO SOLID-HOOFED PIGS CHAP.
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.” 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 hoot”
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 spht 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, ete. 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
' See Bateson, Materials for the Study of Variation, London, 1894, p. 387.
XI CARPUSTAND? DARSUS OF ARDTIODAGD VEES 27) Mt
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, ze. Me IT with
trapezium, trapezoid, and magnum; Me III
with magnum and unciform; Me IV and V
wath unciform. This would appear to give 5, 139.— Dorsal surface of
greater solidity and consequently greater right tarsus of Red Deer
strength to the foot. Peale ede
a, Astragalus; c, cal-
The carpal bones of the Artiodactyla caneum ; ¢’, cuneiform ;
: : ; ‘ A te cb, cuboid 5; mIIT, m1V,
alternate in their articulation; the primi- TWIN 2) Gin HEN
tive state of affairs’ is not retained even ar. | (From Flower’s
: : Osteology.)
in the earliest types. The femur has no
third trochanter, so prevalent in the Perissodactyles. In the
hind-foot the caleaneum 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
* See, however, p. 196, for a discussion as to which is the more primitive
arrangement.
yz, 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." The number of ribs varies from twelve (Camelus,
Hydropotes) through thirteen (Cervus, Gazella) to fourteen in
Dicotyles, Giraffa, ete.
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 Anthracotheridae.
While the stomach of the Perissodactyles is always a simple
sac, 1t is complicated, or shows signs of complication, in the
Artiodactyles. That of the Hippopotamus is divided into two
chambers; there are three in 7ragulus, and four in the typical
Ruminants such as Cervus, Ovis, ete.
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 Vitanotheriwm (see p. 266) is exceptional.
xl ARTIODACTYLES OF MADAGASCAR BGP E
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
Peceary 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.'
Group I.—SUINA.
Fam. 1. Hippopotamidae.—The family Hippopotamidae con-
tains of existing genera only Hippopotamus, for the Liberian dwart
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. Flower? 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 imhabited 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
de 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 qt
2 4 HIPPOPOTAMUS CHAP.
jaw, and they were distinguished as a genus Hexaprotodon, con-
trasting with Zetraprotodon, until intermediate conditions were
observed. Choeropsis, as already observed, was a still further
reduction of the tetraprotodont type. The molars (the formula is
Pm # M8) when worn show a double trefoil pattern. The
orbital cavity is encircled by bone. As with many other aquatic
mammals the kidneys are lobulated.
Fia, 140.—Hippopotamus. Hippopotamus amphibius. x 3.
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."
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 1t 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.
XI ANATOMY OF PIGS 25
ean 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, differ 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-
Fria. 141.—Wild Boar. Sus scrofa. x 4s.
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! 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.
=
Elaphodus' contains probably two species, #. cephalophus of
Milne-Edwards and /. michianus 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.
XI THE MUNTJACS 295
species has a dark iron-grey pelage, and the late Mr. Consul
Swinhoe described it as very Goat-lke 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, 1s 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
Fie, 153.—Mule Deer. Cariacus macrotis. xs. (From Nature.)
are small NQeer 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
2096 SIMPLE ANTLERS CHAP.
are spotted. The antlers are occasionally very simple ; in C. rufus
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 Pudua, the vomer is prolonged back-
wards and divides the posterior nares into two. The bulk of the
species are South American.
Fic. 154.—Chilian Deer. Cariacus chilensis. x+y. (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’ and Mr. Forbes” found the rudi-
1 Proc. Zool. Soc. 1877, p. 789. 2 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. ie
un
A
Ae
Fic. 155.—Water Deer. Hydropotes inermis. x 5. (From Nature.)
~ 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 lencth. 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. Alces machlis. x #5.
Rangifer tarandus, the Reindeer, 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 Reindeer is circumpolar. As is well known, during the
Pleistocene period the Reindeer extended its range as far as the
South of France. Even in the historic period it is said to have
been hunted in Caithness.
teindeer, like so many other particularly Arctic animals,
have regular migrations. In Spitzbergen, for instance, the animal
migrates in the summer to the inland region of the island, and in
XI 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 +}.
Sub-Fam. 2. Moschinae.—JMoschus moschiferus' 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
moschiferus),”’ Proc. Zool. Soc. 1875, p. 159; Garrod, loc. cit. 1877, p. 287; and
l. Jeffrey Bell, Proc. Zool. Soc. 1876, p. 182.
300 ANATOMY OF MOSCHUS ‘ CHAP.
metacarpals, are of the more ancient type represented in ).
are highly characteristic. The horns are bovine in appearance,
standing outwards and then curving upwards.’ There are three
species of Gnu, all from South Africa. They are C. gnu, C. taurinus,
and C. albogulatus.
Of the Cephalophine section there are two genera :—
Cephalophus 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 Cervulus.
Messrs. Sclater and Thomas allow thirty-eight species. The
1 They are straight in the young.
Siu© WATERBUCKS AND REEDBUCKS 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. gum; ¢ straight of these plates in the mouth is
edge of baleen plate; d, e, frayed
out surface of baleen plates. (After Very great. AS many as OU
ene) 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
XI 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 Lalaenoptera includes
the Rorquals, which are Whalebone Whales of large size, differing
from the Right 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 Right Whales, and as a
consequence the plates of baleen are shorter. The hand is only
four-fingered. The cervical vertebrae are for the most part all
free. One of the earhest 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 cf 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 finns, a picked snout, and a mouth so wide that
divers men might have stood upright in it; no teeth, but suck’d
shme 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 given! 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 Rorquals; 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.
350 RORQUALS CHAP.
symphysis of the jaws to the middle of the belly is, as in other
species, marked by furrows, forty to fifty-eight im 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
Jinmarehicus. 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 (4. borealis) seems to be a_ perfectly
inoffensive beast; 1t 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 reduced ' 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. patachonica differ specifically
from this.”
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 1t to beat itself, the surrounding water, and, more
playfully, its mates. The general outline of this Cetacean is
more clumsy than that of Lalaenoptera. The most important
internal difference is in the form of the scapula, which has at
most a shght acromion and coracoid process. These are rather
more pronounced, according to Messrs. van Beneden and Gervais,”
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 Ostéographie des Cétacés, Paris, 1880, p. 130.
XII THE CALIFORNIAN WHALE 257
in the southern form of the genus, which is known as JZ. lalandit.
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
Seammon, who observed many “ gams” or herds of these Whales,
remarked! that he had extreme difticulty in finding any two
individuals precisely ahke! The best-known species in any case
is the northern Jf. Jongimana, which occurs on our own coasts.
The genus is, ike 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 J/. lalandii (with which
in that case, perhaps, JZ. capensis and MW. novae zelandiae will be
synonymous). Quite recently M. Gervais has insisted upon a
Megaptera indica from the Persian Gulf. Jlegaptera 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.
Rhachianectes, with but one species, &. glaucus, 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 Rorqual, 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-lhke; 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 Rorquals, and the sternum is quite as in that group.
The scapula has more the shape of that of Lalaena.
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 North-West Coast of North America, 1874.
2 Cf. Secammon, Joc. 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. Rhachianectes 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 Right Whales of the genus
Balaena are to be distinguished from Neobalaena and from
the Rorquals 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-hmb 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 Right Whales coming from different parts of the
world. In Dr. Gray’s catalogues we find the following allowed,
viz. Balaena, Hubalaena, 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 Right
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 “ Right 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
XII GREENLAND WHALE . 25/6)
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 Rorqual, 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 Right Whale, B. australis,'
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 6. 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. biscayensis ; 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 Guetaria,’ writes Sir Clements Markham,’
“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. Rondeletius
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, 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 hilla
tower from which they could see “the Balaines which pass, and
perceiving them coming partly by the loud noise they make, and
1 Proc. Zool. Soc. 1881, p. 969.
* 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 carease 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 Neobalaena 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 Right
Whales and the Rorquals. 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 outhne is not Rorqual-like in
spite of its sumilar proportions. The whalebone is long. The
throat is not grooved. Neobalaena 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 Rorquals than that of the Right Whales ;
3262 TOOTHED WHALES CHAP.
that is to say, it is long and not very high. The skull is most
like that of Balaena, 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 marginata.'
Sus-OrpDER 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 :—AIl 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
(?exe. 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. yan 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. vii. 1876, p. 251; and
3eddard, Trans. Zool. Soc. xv. 1901, p. 87.
XII 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 cominence 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. No distinct lachrymal
bone. Stomach with only four compartments (?7as to Kogic).
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 leneth 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,’ 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 it ?—
that the Sperm Whale turns over upon its back to bite. The
blow-hole is single, and shaped like the sound-hole 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 UV 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. ainbergris. 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.’ 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 hes the spermaceti already
referred to. The skull, as in Toothed Whales generally, is ex-
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 1s in the Ziphioids. The ptery-
voids 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:—TI 1, II 5,
[EE 53 Vi 4.53;
‘
1 See Pouchet, ‘Contribution a’ lhistoire du spermaceti,” Bergens Musewms
Aurbog for 1893, No. 1.
XII PEROCIDY 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 1t 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 coms 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 lbs. 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 enough to swallow a man
(the Whale is credited with being that which swallowed Jonah),
does not usually admit fishes larger than Bonitos and Albacores.
The ferocity of the Cachalot has been denied and affirmed.
It certainly has great strength, for it can 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 HIGH-FINNED CACHALOT CHAD.
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 Weganeuron krefti. The “ High-
finned Cachalot” rests mainly upon the suggestions of Sir Robert
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
inuseum 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 im 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 Aogia (sometimes written Cogia), the so-called
“Pyomy Sperm Whale,’ is a southern form of much smaller
dimensions than its gigantic ally just described. Aogia does
not exceed 15 feet or so in length. It differs from Physeter
also in the well-marked and faleate 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 Aogia all the
cervical vertebrae are ankylosed together; the skull is short,
though equally asymmetrical; the ribs are as many as twelve or
1 Yule, Zravels of Marco Polo, ii. London, 1874, p, 281.
xu 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 difliculty 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
(EBuphiysetes, Callignathus), but there do appear to be reasons for
allowing two species, if the accounts of their osteology are to he
depended upon. One of these is AY 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 A. stmws, 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
L 2311-5, TIl.4, IV 4, V2.
A Californian species has been called AY jlowert, whose teeth
seem to be particularly long and recurved. And the New Zealand
K. pottst 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.
All the Dogs have a caecum! of simple cylindrical form. In
C. cancrivorus, C. jubatus, and Nyctereutes procyonides this organ
is straight or only very faintly curved; in other Dogs it is
eolled into an §-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 J/eficyon contains but one recent species, the Bush
Dog CL. venaticus, Lund) of British Guiana. The animal has a
somewhat Paradoxure-hke, 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. Jeticyon 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
eurved. 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 Ieticyon resemble the primitive dogs.”
A genus Nyctereutes is usually separated from Canis for the
inclusion of V. procyonides only. The separation is based upon
1 Flower, Proc. Zool. Soc. 1879 2 766.
* Proc. Zool. Soc. 1880, p. 7
XII DENTITION OF OZOCYON 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 Raccoon,’ especially in the black patches below
Fre. 206.—Raccoon-like Dog. Nyctereutes procyonides. x &.
the eyes, whence of course the scientific name and the pseudo-
vernacular “ Raccoon-like Dog.” It inhabits China and Japan.
As to structure, there is hardly anything that justifies its ex-
clusion from the genus Canis. Garrod, however, mentions the
unusually large size of the Spigelian lobe of the liver.
Wortman and Malkens” have instituted a genus Nothocyon for
Dr. Mivart’s species C. wrostictus® and C. parvidens, which are
both South American forms.
The genus Ofocyon contains but one species, O. 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 ~ Moreover the carnassial- teeth are not so pro-
4
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 thrs point of external likeness.
2 Bull. Amer. Mus. Nat. Hist. xii. 1900, p. 109.
3 Proc. Zool. Soc. t890, 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-lke 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 2. The one species is L. pictus, the Cape Hunting Dog. It is
singularly like a Hyaena’ in general appearance; the ochraceous
grey ground-colour with black markings and the iong ears pro-
duce this lhkeness. 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 ZL. 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, Zycaon” 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, Zycaon 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 Canis. But the numerous
1 Temminck, its original describer, placed it in the genus Hyaena.
2 See Garrod, Proc. Zool. Soc. 1878, p. 373.
XIII ALOPECOIDS AND THOOIDS AI7
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. xt.
series he termed the “ Alopecoid ” or Fox-like, and the “ Thooid ”
or Wolf-lke. It was suggested that the generic name Vulpes be
Fic. 208.—Prairie Wolf or Coyote. Canis latrans. xt
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 25
_
418 CHARACTERS OF FOXES CHAP.
the Dogs belonging to genera that have already been separated
off. Thus Zycaon is distinctly Thooid. The characters in
question are these:—JIn 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.
Fic. 209.
Japanese Wolf. Canis hodophylax. «4. (From Nature.)
To the Fox series belong among others the species C. lagopus
(Arctic Fox), C. zerda (the Fennec), C. chama (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. hodophylaz),
the Red Wolf of America (C. jubatus), the Jackals (C. aureus, C.
anthus, ete.), the Prairie Wolf (C. /atrans), and a number of
American forms, such as C. azarae, its close ally C. cancrivorus
(=C. rudis), C. antarcticus, C. magellanicus, ete., are decidedly
Wolves rather than Foxes.
= =
XIII 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
Red Crag, a deposit of Plocene times. Its prevalence now is no
doubt due to its preservation as a beast of chase. It lves 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 teeyas with its accomplishments. The worthy
Archbishop of Upsala, Olaus Magnus, figured Foxes dipping their
tails in the streams, and then pulling out inquisitive crayfishes
420 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 ':—
CO. anthus, C. variegatus, C. mesomelas, C. 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,
Fic. 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
A
1 Proc. Zool. Soc. 1899, p. 533.
_-
ra
=
Z
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, fearmg 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. pallipes, C. chanco, and C. laniger, are
hardly, if at all, different from C. lupus. Professor Huxley has
remarked upon the likeness of C. pallipes 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
A 2 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
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
Fic. 211.—Dingo. Canis dingo. x.
way into time, and may thus still have imported this companion
with him. Anyhow it is quite a wild creature now. Dr. Nehring,
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 CL miki
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
XIII 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 Zycaon (L. anglicus) has been
met with in caves in Glamorganshire, while Jcticyon of South
America appears to be congeneric with Speothos 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 Plocene.
The earlier types of Dogs have been placed in different genera.
Cynodictis is an Kocene 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, Simocyoninae. 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. Hnhydrocyon is an intermediate form; it has lost one
premolar in each jaw.
Amphicyon, forming the type of another sub-family, Amphi-
eyoninae, though usually placed among the Dogs, presents us with
424 THE BEAR TRIBE CHAP.
many Bear-lke 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. giganteus, 1s 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
microdont modern Dogs. In accordance with its age the cerebral
conyolutions of this Dog are much simpler than in existingDogs,
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 im 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 t-inch from the posterior
margin of the bulla. In the Raccoon the bullae are much more
swollen, and the paroccipital processes are closer to them. In
the Marbled Polecat, Putorivs sarmaticus, the bullae are fairly
XIII 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 /etonyx 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 WVandinia and