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condition of dentition, the functional teeth being reduced to one pair of large cutting incisors situated close to the median line, and one great, trenchant, compressed premolar, on each side above and

FIG. 51.-Front view of skull of Thylacoleo carnifex, restored. natural size. From Quart. Journ. Geol. Soc. vol. xxiv. p. 312.

below. It was first described as a carnivorous Marsupial, and named, in accordance with its presumed habits,

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existing Phalangerida than to that of the Macropodidæ, and the genus may be provisionally regarded as the type of a distinct subfamily of the former.

Family MACROPODIDE.

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(0-1) c , p, m Incisors sharp and cutting,

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those of the lower jaw frequently having a scissor-like action against one another; upper canine, if present, small. Penultimate premolar shed with the fourth milk-molar, which is molariform and long persistent. Molars wide, and either transversely ridged or bluntly tuberculate. Premolars and molars moving forwards in the skull as the age of the animal increases, this being most marked in the larger species. Masseteric fossa of mandible hollowed out below into a deep cavity walled in externally by a plate of bone, and communicating with the inferior dental canal by a large foramen. Hind limbs usually larger than the anterior ones, and progression generally saltatorial. Fore feet with five digits; hind feet syndactylous, the fourth digit being very large and strongly clawed; hallux usually absent. Tail generally long and hairy,

occasionally prehensile; stomach sacculated.

opening forwards.

Pouch large and

The Macropodidae or Kangaroos, taken as a whole, form a very well-marked family, easily distinguished from the other members of the suborder by their general conformation, and

by peculiarities in the structure of their limbs,

teeth, and other organs.
They vary in size from
that of a sheep down to a small rabbit. The
head, especially in the larger species, is small,
compared with the rest of the body, and tapers
forward to the muzzle. The shoulders and fore
limbs are feebly developed, and the hind limbs
usually of disproportionate strength and magnitude,
which gives them a peculiarly awkward appearance
when moving about on all fours, as they occasion-
ally do when feeding. Rapid progression is, how-
ever, performed only by the powerful hind limbs,
the animal covering the ground by a series of
immense bounds, during which the fore part of the
body is inclined forwards, and balanced by the
long, strong, and tapering tail, which is carried
horizontally backwards. When not moving they
often assume a perfectly upright position, the tail
aiding the two hind legs to form a sort of support-
ing tripod, and the front limbs dangling by the
side of the chest. This position gives full scope
for the senses of sight, hearing, and smell to warn
of the approach of enemies, from which these
animals save themselves by their bounding flight.
The fore paws have five distinct digits, each armed
with a strong curved claw.

FIG. 52.-Skeleton

Kangaroo.

The hind foot (Fig. 52), as being a typical example of the syndactylous modification, may be noticed in some detail. It is extremely long and of right hind foot of narrow, and (with only one exception) without any hallux or great toe. It consists mainly of one very large and strong toe, corresponding to the fourth of the human or other typically developed foot, ending in a strong, curved, and pointed claw. Close to the outer side of this lies a smaller fifth digit, and to the inner side two excessively slender toes (the second and third), bound together almost to the extremity in a common integument. The two little claws of these toes, projecting together from the skin, may be of use in scratching and cleaning the fur of the animal, but the toes themselves must have quite lost all connexion with the functions of support or progression.

The dentition of the Kangaroos, functionally considered,

consists of sharp-edged incisors, most fully developed near the median line of the mouth, for the purpose of cropping the various kinds of herbage on which they feed, and ridged and tuberculated molars for crushing it, there being no tusks or canines for offensive or defensive purposes.

The number of vertebræ is-in the cervical region 7, dorsal 13, lumbar 6, sacral 2, caudal varying according to the length of the tail, but generally from 21 to 25. In the fore limb the clavicle and the radius and ulna are well developed, allowing of considerable freedom of motion of the hand. The pelvis has large epipubic or "marsupial" bones. The femur is short, and the tibia and fibula

[graphic]

FIG. 53. The Great Gray Kangaroo (Macropus giganteus).

are of great length, as is the foot, the whole of which is applied to the ground when the animal is at rest in the upright position.

The stomach is of large size, and very complex, its walls being puckered up by longitudinal muscular bands into a great number of sacculi, like those of the human colon. The alimentary canal is long, and the cæcum well developed. All the species have a marsupium or pouch formed by a fold of the skin of the abdomen, covering the mammary glands with their four nipples. In this pouch the young are placed as soon as they are born; there their growth and development proceeds; and to it they resort temporarily for the purpose of shelter, concealment, or transport, for some time after they are able to run and jump about the ground and feed upon the same herbage which forms the nourishment of the parent. During the early period of their sojourn in the pouch,

the blind, naked, helpless young creatures (which in the Great Kangaroo (Fig. 53) scarcely exceed an inch in length) are attached by their mouths to the nipples of the mother, and are fed by milk injected into their stomach by the contraction of the muscle covering the mammary gland.

The Kangaroos are all vegetable feeders, browsing on grass and various kinds of herbage, the smaller species also eating roots. They are naturally timid, inoffensive creatures; but the larger ones when hard pressed will turn and defend themselves, sometimes killing a dog by grasping it in their fore paws, and inflicting terrible wounds with the sharp claws of their powerful hind legs, sustaining themselves meanwhile upon the tail. A few aberrant forms are arboreal. The great majority are inhabitants of Australia and Tasmania, forming one of the most prominent and characteristic features of the fauna of these lands, and in the scenery of the country, as well as the economy of nature, performing the part of the deer and antelopes of other parts of the world, which are entirely wanting in Australia. Kangaroos were very important sources of food-supply to the natives, and are hunted by the colonists, both for sport and with a view to their destruction, on account of the damage they naturally do in consuming the grass, now required for feeding cattle and sheep. Notwithstanding this, they have in some districts increased in numbers, owing to the suppression of their former enemies, the aborigines and the Dingo or native dog. A few species are found in New Guinea and the adjacent islands, which belong, in the zoological sense, to the Australian region.

Before noticing the various generic types of the Macropodide, a few words are necessary in respect of the tooth-change, and we may here quote the observations of Mr. O. Thomas on this subject. "The full dentition of the members of this family consists, in the upper jaw, first of three incisors, then of a small canine (often, however, suppressed, as in Fig. 55), and then of six cheek-teeth, of which the second in the series is the only one which has a milk or deciduous predecessor, and is therefore the one to be regarded as the last premolar of the typical mammalian dentition. The special characteristics that render the development and succession of the teeth in the Macropodide, and especially in the genus Macropus, so puzzling to systematic zoologists, are: firstly, a general progression forwards in the jaw of the whole tooth-row, comparable to that found elsewhere only in the Elephants and some Sirenians; and, secondly, the fact that before the tooth-change the first tooth of the series (p 3) and the single milk-tooth (dm 4) placed next to it, both of which fall out at the change, are respectively so very similar in shape and size to the first and second teeth of the permanent series, viz. the permanent premolar (p 4) and the first

molar (m 1), as to be most naturally mistaken for, or compared with, them in specific descriptions. . . . The necessary knowledge as to the stage of dentition in which any skull may be, can often be gained only by cutting open the bone either above and behind the first tooth of the series to see if the true permanent p 4 be still buried there (in which case, of course, that first tooth is only p 3), or behind the last visible molar to see if there be yet another tooth behind it, showing it to be m 3 and not m 4. The first plan is, as a rule, the better, since p 4 is generally by far the most important tooth for diagnostic purposes, and its characters have, therefore, in any case to be taken into account."

The Macropodide are divided into three well-marked sections: (1) the true Kangaroos (Macropodina); (2) a group consisting of smaller animals, commonly called Rat Kangaroos, or (improperly) "Kangaroo Rats," or sometimes Potoroos; and (3) the Hypsiprymnodontine, now represented only by a single species.

Subfamily Hypsiprymnodontinæ.-Size very small. Claws small, feeble, and subequal. Hind feet with an opposable hallux. Tail naked and scaly. The fourth premolar twisted obliquely outwards, as in Phalanger. Other teeth as in the Potoroina.

This subfamily is now represented only by the genus Hypsiprymnodon, which is a form of great interest, as showing a structure of foot connecting that of the Kangaroos with that of the Phalangers. The single known species, H. moschatus, was described by Ramsay from specimens discovered in north-east Australia. It was described almost simultaneously by Owen under the name of Pleopus nudicaudatus. From the resemblance in the structure of the foot and the obliquity of the premolars to the Phalangers Mr. Thomas has some hesitation as to which family should receive this genus, but the macropine characters of the mandible preponderate in favour of the Macropodide.

Triclis.2-A lower jaw of a much larger form from the Pleistocene deposits of Australia apparently indicates another member of this subfamily, having the outwardly directed and grooved premolar characteristic of Hypsiprymnodon. It differs, however, from that genus, and also from all other known Macropodide, in having a small tooth between the incisor and fourth premolar, which apparently represents a canine, or perhaps an anterior premolar. This form indicates, therefore, a closer connexion between the Phalangeride and Macropodide than any other.

Subfamily Potoroinæ.-The second section or subfamily, the Potoroine, have the first upper incisor narrow, curved, and much exceeding the others in length (Fig. 54). Upper canines always persistent, flattened, blunt, and slightly curved. Premolars of both

1 Ramsay, Proc. Linn. Soc. N. S. Wales, vol. i. p. 33 (1876).
2 De Vis, Proc. Roy. Soc. Queensland, ser. 2, vol. iii. p. 8 (1888).

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