General Characters.—The Metatheria or Didelphia are represented at present by numerous species, presenting great diversities of general appearance, structure, and habits, although all united by many essential anatomical and physiological characters, which, taken altogether, give them an intermediate position between the Prototheria and the Eutheria.
Although the striking differences in external form, in many anatomical characters, and in mode of life of various animals of this section might lead to their division into groups equivalent to the orders of the Eutheria, it is more convenient on the whole to adhere to the usual custom of treating them all as forming one order called Marsupialia,[37] the limits of which are therefore equivalent to that of the subclass. The more essentially distinctive characters are as follows.
In the structure of the brain and the presence of epipubic bones they agree with the Prototheria, while in the structure of the ear-bones and the shoulder-girdle and the presence of teats on the mammary glands they resemble the Eutheria, the reproductive organs belonging to neither one nor the other type, but having a special character representing an intermediate grade of development. The ureters open into the base of the bladder. The oviducts are differentiated into uterine and Fallopian portions, and open into a long and distinct vagina, quite separate from the cystic urethra. The penis is large, but its crura are not directly attached to the ischia. The spongy body has a large bifurcated bulb. The young are born in an exceedingly rudimentary condition, and are never nourished by means of an allantoic placenta, but are transferred to the nipple of the mother, to which they remain firmly attached for a considerable time, nourished by the milk injected into the mouth by compression of the muscle covering the mammary gland. They are therefore the most typically mammalian of the whole class. The nipples are nearly always concealed in a fold of the abdominal integument or “pouch” (marsupium) which serves to support and protect the young in their early helpless condition.
Entering more fully into the characters of the subclass, which are also those of the order Marsupialia, it may be observed that the brain is generally small in proportion to the size of the animal, and the surface-folding of the cerebral hemispheres, though well marked in the larger species, is never very complex in character, and is absent in the medium-sized and smaller species. The arrangement of the folding of the inner wall of the cerebrum differs essentially from that of all known Eutheria, the hippocampal fissure being continued forward above the corpus callosum, which is of very small size. The anterior commissure is, on the other hand, greatly developed.
Fig. 34.—Teeth of upper jaw of Opossum (Didelphys marsupialis), all of which are unchanged, except the last premolar, the place of which is occupied in the young animal by a molariform tooth, represented in the figure below the line of the other teeth.
The teeth are always divisible, according to their position and form, into incisors, canines, premolars, and molars; but they vary much in number and character in the different families. Except in the genus Phascolomys, the number of incisors in the upper and lower jaws is never equal. The true molars are very generally four in number on either side of each jaw. The chief peculiarity in the dentition lies, however, in the mode of succession. Thus there is no vertical displacement and succession of the teeth, except in the case of a single tooth on either side of each jaw, which is always the hindermost of the premolar series, and is preceded by a tooth having more or less of the characters of a true molar (see Fig. 34); this deciduous tooth being the only one comparable to the “milk-teeth” of the diphyodont Eutheria. In some cases (as in Potorous) this tooth retains its place and function until the animal has nearly, if not quite, attained its full stature, and is not shed and replaced by its successor until after all the other teeth of the permanent series, including the posterior molars, are fully in place and use. In others, as the Thylacine, it is very rudimentary in form and size, being shed or absorbed before any of the other teeth have cut the gum, and therefore quite functionless. It must further be noted that there are some Marsupials, as the Wombat, Myrmecobius, and the Dasyures, in which no such milk-tooth, even in a rudimentary state, has yet been discovered, possibly in some cases from want of materials for observation at the right stage of development.
Epipubic or marsupial bones are present in both sexes of nearly all species. In one genus alone, Thylacinus, they are not ossified. The number of dorso-lumbar vertebræ is always nineteen, although there are some apparent exceptions caused by the last lumbar being modified into a sacral vertebra. The number of pairs of ribs is nearly always thirteen. The tympanic bone remains permanently distinct. The carotid canal perforates the basisphenoid. The lachrymal foramen is situated upon or external to the anterior margin of the orbit, and there are generally large vacuities in the bony palate. The angle of the mandible is (except in Tarsipes) more or less inflected. The hyoid bones have always a peculiar form, consisting of a small, more or less lozenge-shaped basihyal, broad ceratohyals, with the remainder of the anterior arch usually unossified, and stout, somewhat compressed thyrohyals. There are two anterior venæ cavæ,[38] into each of which a “vena azygos” enters. In the male the testes are always contained in a scrotum, which is suspended by a narrow pedicle to the abdomen in front of the penis. The vasa deferentia open into a complete and continuous urethra, which is also the passage by which the urine escapes from the bladder, and is perfectly distinct from the passage for the fæces, although the anus and the termination of the urethro-sexual canal are embraced by the same sphincter muscle. The glans is often bifurcated anteriorly. In the female the oviducts never unite to form a common cavity or uterus, but open separately into the vagina, which at least for part of its course is double. The mammæ vary much in number, but are always abdominal in position, having long teats, and in most of the species are more or less enclosed in a fold of the integument forming a pouch or marsupium, though in some this is entirely wanting, and the newly-born, blind, naked, and helpless young, attached by their mouths to the teat, are merely concealed and protected by the hairy covering of the mother’s abdomen. In this stage of their existence they are fed by milk injected into their stomach by the contraction of the muscles covering the mammary gland, the respiratory organs being modified temporarily, much as they are permanently in the Cetacea—the elongated upper part of the larynx projecting into the posterior nares, and so maintaining a free communication between the lungs and the external surface independently of the mouth and gullet, thus averting the danger of suffocation while the milk is passing down the latter passage.
Fig. 35.—Front view of skull of Sarcophilus ursinus, showing polyprotodont and carnivorous dentition (Quart. Journ. Geol. Soc. vol xxiv. p. 313).
Distribution.—The existing species of Marsupials are, with the exception of one family (the Didelphyidæ), limited in geographical distribution to the Australasian region,[39] forming the chief mammalian fauna of Australia, New Guinea, and some of the adjacent islands. The Didelphyidæ are almost purely Neotropical, one or two species ranging northwards into the Nearctic region. Fossil remains of members of this family have also been found in Europe and America in strata of the Eocene and early Miocene periods; and it is probable that at least many of the polyprotodont Mesozoic mammals noticed in Chapter IV. are referable to the Marsupialia.
Fig. 36.—Front view of skull of Koala (Phascolarctus cinereus), showing diprotodont and herbivorous dentition (Quart. Journ. Geol. Soc. vol. xxiv. p. 313).
Classification.—In dividing the Marsupials into minor groups, it may be observed that one of the most obvious distinctive characters among them is derived from the form and arrangement of the teeth. In certain species, as the Opossums, Dasyures, and Thylacine, the incisors are numerous, small, and subequal in size, and the canines large, as in the typical placental Carnivores (Fig. 35). To these the term “polyprotodont” is applied, and they are all more or less carnivorous in their habits. In others the central incisors are very prominent, and the lateral incisors and canines absent or subordinate in function (Fig. 36). These are called “diprotodont,” and they are all wholly or in great part vegetable feeders. In one group of these, the Wombats, there are but two incisors above and the same number below; but all the others, including the Kangaroos, Koalas, and Phalangers, have two functional incisors below and as many as six above, three on each side, but of these the first or central pair is the most fully developed.
Some hesitation has frequently been expressed as to whether the Polyprotodont and Diprotodont types are entitled to constitute distinct primary groups, owing to the presence of syndactylism among the Peramelidæ in the former, as well as in the latter; but if Mr. O. Thomas is right in regarding this feature as acquired independently in the two groups we may safely adopt such a division. Taking various combinations into consideration, the existing Marsupials readily group themselves into six very natural families, the leading characters of which may be summarised as follows:—
Order Marsupialia.
A. Polyprotodontia.—Incisors numerous, small, subequal. Canines larger than the incisors. Molars with sharp cusps.
α. Incisors ⁵⁄₄. Hind feet with the four outer toes subequal, distinct, and a well-developed opposable hallux. Didelphyidæ.
β. Incisors ⁴⁄₃. Hind feet with four outer toes distinct. Hallux small or rudimentary, rarely opposable. Dasyuridæ.
γ. Incisors ⁴⁻⁵⁄₃. Hind feet long and narrow. Fourth toe larger than the others. Hallux rudimentary or absent. Second and third toes very slender, and united in a common integument (syndactylous). Peramelidæ.
B. Diprotodontia.—Incisors not exceeding ³⁄₃, usually ³⁄₁ but occasionally ¹⁄₁. Central (first) upper and lower incisors large and cutting. Upper canines generally, and lower invariably, absent or small. Molars with bluntly tuberculated or transversely ridged crowns.
α. Teeth with persistent pulps. Incisors ¹⁄₁, large, scalpriform, with enamel on the outer surface only. No canines. Hind feet with four subequal outer toes, partially syndactylous, and with rudimentary hallux. Phascolomyidæ.
β. Teeth rooted. Three upper incisors and a canine. Hind limbs not disproportionately large. Feet syndactylous, broad, with four subequal outer toes, and a large opposable hallux. Phalangeridæ.
γ. Teeth rooted. Three upper incisors, and frequently a canine. Hind limbs disproportionately large, with syndactylous feet as in Peramelidæ. Macropodidæ.
The leading characters of this group are given in the foregoing schedule. This group is the only one represented at the present day, and so far as we know also in past epochs, beyond the confines of the Australasian region and adjacent islands.
Dentition: i ⁵⁄₄, c ¹⁄₁, p ³⁄₃, m ⁴⁄₄; total 50. Incisors very small and pointed. Canines large. Premolars with compressed pointed crowns. Molars with numerous sharp cusps. The last premolar preceded by a deciduous multicuspidate milk-molar, which remains in place until the animal is nearly adult (Fig. 34). Limbs of moderate development, each with five complete and distinct toes, all of which are provided with short, compressed, curved, sharp claws of nearly equal size, except the first toe of the hind foot or hallux (Fig. 37), which is large, widely separable from the others, to which it is opposed in climbing, and terminates in a dilated rounded extremity, without a nail. Tail generally long, partially naked and prehensile. Stomach simple. Cæcum of small or moderate size. Pouch generally absent, sometimes represented by two lateral folds of the abdominal integument, partially covering the teats, rarely complete. Vertebræ: C 7, D 13, L 6, S 2, C 19-35.
Fig. 37.—Skeleton of the right hind foot of the Virginian Opossum (Didelphys marsupialis).
The Didelphyidæ, or true Opossums, differ from all other existing Marsupials in their habitat, being peculiar to the American continent. They are mostly carnivorous or insectivorous in their diet, and arboreal in habits.
Opossums occur throughout the greater part of the American continent, ranging from the United States to Patagonia, the greater number of species being found in the warmer regions. In South America the opossums take the place of the Eutherian Insectivora, and the sharp cusps on their teeth are admirably adapted for crushing the insects on which they mainly subsist.
Chironectes.[40]—The family comprises two genera only, namely Didelphys, containing all the species, with the exception of the curious Yapock, which forms by itself the genus Chironectes and is distinguished from all other Opossums by its webbed feet, non-tuberculated soles, and peculiar coloration. Its ground colour is light gray, with four or five sharply-contrasted brown bands passing across its head and back, and thus giving it a very peculiar mottled appearance. It is almost wholly aquatic in its habits, living on small fish, crustaceans, and water insects. Its range extends from Guatemala to southern Brazil.
Didelphys.[41]—The type genus Didelphys is a very large one, containing, according to Mr. O. Thomas, twenty-three existing species. It may be divided into five groups, or subgenera, all of which have received distinct names. The typical group is represented only by the common or Virginian Opossum (D. marsupialis), of which the numerous varieties have received separate specific names. This species is of large size, with a long, scaly, prehensile tail, and long bristle-like hairs mingled with the fur. The pouch is complete. It ranges over all temperate North America, and is also found in central and tropical South America, where it is commonly known as the Crab-eating Opossum. This animal is extremely common, being even found living in the towns, where it acts as a scavenger by night, retiring for shelter by day upon the roofs of the houses or into the sewers. The female produces in the spring from six to sixteen young ones, which are placed in her pouch immediately after birth, and remain there until able to take care of themselves.
The second or Metachirine group includes three species found all over the tropical parts of the New World. They are of medium size, with short close fur, very long, scaly, and naked tails, and less developed ridges on their skulls than in the type species. As a rule there is no pouch adapted to carry the young, which commonly ride on their mother’s back, holding on by winding their prehensile tails round hers. The Philanderine group is closely allied to the preceding, but is readily distinguished by the woolly hair, and the brown streak down the middle of the face. The Woolly Opossum (D. lanigera), which is represented in the accompanying woodcut (Fig. 38) carrying its young in the fashion mentioned above, is one of the two species of this group. In the fourth or Micoureine group the numerous species are all smaller than in the preceding groups, and have short and close hair, and no dark streak down the face. The best known species is the Murine Opossum (D. murina), little larger than a House-Mouse, and of a bright red colour, which is found as far north as central Mexico, and extends thence right down to the south of Brazil. The last or Peramyne group contains several extremely shrew-like species, of very small size, with short, hairy, and usually non-prehensile tails, not half the length of the trunk, and with wholly unridged skulls. The most striking member of the group is the Three-striped Opossum (D. americana), from Brazil, which is of a reddish-gray colour, with three clearly-defined deep-black bands down its back, very much as in some of the striped mice of Africa.
Fig. 38.—The Woolly Opossum (Didelphys lanigera).
The numerous fossil species of Opossum found in the Upper Eocene and Lower Miocene of Europe are of especial interest from a distributional point of view, since they indicate how the Opossums of America may have been connected with the Australian Marsupials. These forms were originally referred to Didelphys, but have been subsequently described as Peratherium and Amphiperatherium. The characters of the molar teeth on which these genera are based do not appear to be sufficiently important to justify their separation from Didelphys. Allied forms occur in the Tertiaries of North America, which were originally described under the name of Herpetotherium, but have been subsequently referred to Peratherium. Remains of many of the existing species of Opossum are found in a fossil condition in the Pleistocene cave-deposits of Brazil.
Dentition: i ⁴⁄₃, c ¹⁄₁, p and m numerous, variable. Incisors small; canines well developed; molars with pointed cusps. Limbs equal. Fore feet with five subequal toes terminating in claws. Hind feet with the four outer toes well developed, and distinct from each other and bearing claws; the first (or hallux) clawless, generally rudimentary, sometimes entirely wanting. Stomach simple. No cæcum. Predatory carnivorous or insectivorous animals, inhabitants of Australia, Tasmania, and the southern parts of New Guinea and some of the adjacent islands. The aberrant genus Myrmecobius, though clearly a member of this family, is so sharply distinguished from all the others as to render a division into two subfamilies necessary.
Fig. 39.—The Thylacine (Thylacinus cynocephalus).
Subfamily Dasyurinæ.—This comprises the more typical Dasyuridæ, in which the premolars and molars never exceed the normal number of seven on either side of each jaw, and in which the tongue is not specially extensile.
Thylacinus.[42]—Dentition: i ⁴⁄₃, c ¹⁄₁, p ³⁄₃, m ⁴⁄₄ = 46. Incisors small, vertical, the outer one in the upper jaw larger than the others. Summits of the lower incisors, before they are worn, with a deep transverse groove dividing them into an anterior and a posterior cusp. Canines long, strong, and conical. Premolars separated from one another by intervals, with compressed crowns, increasing in size from before backwards. True molars in general characters resembling those of Dasyurus, but of more simple form, the cusps being not so distinct nor sharply pointed. Milk-molar very small, and shed before the animal leaves the mother’s pouch. Humerus with an entepicondylar foramen. General form very Dog-like. Head elongated. Muzzle pointed. Ears moderate, erect, triangular. Fur short and closely applied to the skin. Tail of moderate length, thick at the base and tapering towards the apex, clothed with short hair. Hallux (including the metacarpal bone) wanting. Vertebræ: C 7, D 13, L 6, S 2, C 23. Marsupial bones represented only by small unossified fibro-cartilages.
The only known existing species of this genus, T. cynocephalus (Fig. 39), though smaller than a common Wolf, is the largest predaceous Marsupial at present living. It is now entirely confined to the island of Tasmania, although fragments of bones and teeth found in caves afford evidence that a closely allied species once inhabited the Australian mainland. The general colour of the Thylacine is grayish brown, but it has a series of transverse black bands on the hinder part of the back and loins, whence the name of “Tiger” frequently applied to it by the colonists. It is also called “Wolf,” and sometimes, though less appropriately, “Hyæna.” Owing to the havoc it commits among the sheepfolds, it has been nearly exterminated in all the more settled parts of Tasmania, but still finds shelter in the almost impenetrable rocky glens of the more mountainous regions of the island. The female produces four young at a time. The pouch opens backwardly, and there are four mammæ. The figure of the skull exhibits the peculiar Dog-like form so characteristic of the genus.
Fig. 40.—Right lateral aspect of the skull of the Thylacine.
Sarcophilus.[43]—Dentition: i ⁴⁄₃, c ¹⁄₁, p ²⁄₂, m ⁴⁄₄. Upper incisors nearly equal, and placed vertically, the first not differentiated from the rest. Premolars rounded and closely crowded between the canine and molars, with broad crowns; molars broad and heavy, the last one without a distinct hind talon. Form thick and powerful; head disproportionately large for the body; muzzle short and broad; ears broad and rounded; tail of moderate length, and evenly hairy. Hallux wanting; soles of feet naked, without defined pads. Humerus with entepicondylar foramen.
This genus is now represented only by a single species (S. ursinus) found in Tasmania, where, from its ferocious and destructive habits, it is commonly known under the name of the “Devil.” A front view of the skull is shown in Fig. 35.
The prevailing colour of this animal is black, and the size about equal to that of an English Badger; its habits are fossorial, and it is very destructive to sheep. On account of the similarity in the number of its teeth this genus has been generally included in the next one, but in the structure of the teeth it is much nearer to Thylacinus. An extinct species is found in the Pleistocene deposits of the mainland of Australia.
It may be observed that the two premolars missing from the typical series of four in this and the next genus are the second and the fourth; the fourth milk-molar being likewise absent. In Thylacinus and other Polyprotodonts with three premolars it is the second that is missing.
Dasyurus.[44]—Dentition: i ⁴⁄₃, c ¹⁄₁, p ²⁄₂, m ⁴⁄₄; total 42. Upper incisors nearly equal, and placed vertically; first slightly longer, narrower, and separated from the rest. Lower incisors sloping forwards and upwards. Canines large and sharply pointed. Premolars with compressed and sharp-pointed crowns, and slightly developed anterior and posterior accessory basal cusps. True molars with numerous sharp-pointed cusps. In the upper jaw the first three with crowns having a triangular oral surface, the fourth small, simple, narrow, and placed transversely. In the lower jaw the molars more compressed, with longer cusps; the fourth not notably smaller than the others. Form viverrine. Ears long and narrow, prominent, and obtusely pointed. Hallux rudimentary, or absent; its metatarsal bone always present. Tail long and well clothed with hair. Humerus without an entepicondylar foramen. Vertebræ: C 7, D 13, L 6, S 2, C 18-20.
The Dasyures are small Civet-like animals with a gray or brown pellage profusely spotted with white; they are mostly inhabitants of the Australian continent and Tasmania, where in the economy of nature they take the place of the smaller predaceous Carnivora, the Cats, Civets, and Weasels of other parts of the world. They hide themselves in the daytime in holes among rocks or in hollow trees, but prowl about at night in search of the small living mammals and birds which constitute their prey. The species are not numerous, and include D. maculatus, about the size of a common Cat, inhabiting Tasmania and the southern part of Australia; D. viverrinus, Tasmania and Victoria; D. geoffroyi, nearly all Australia; D. hallucatus, North Australia; D. albopunctatus, New Guinea.
Remains referred to D. viverrinus occur in the Australian Pleistocene deposits.
Phascologale.[45]—This genus comprises a considerable number of small Marsupials, none of them exceeding a common Rat in size, differing from the Dasyures in possessing an additional premolar—the dentition being i ⁴⁄₃, c ¹⁄₁, p ³⁄₃, m ⁴⁄₄; total 46,—and having the teeth generally developed upon an insectivorous rather than a carnivorous pattern, the upper middle incisors being larger and inclined forwards, the canines relatively smaller, and the molars with broad crowns, armed with prickly tubercles. The muzzle is pointed. Ears moderately rounded and nearly naked. Feet broad and short. Fore feet with five subequal toes, having compressed, slightly curved, pointed claws. Hind feet with the four outer toes subequal, having claws similar to those in the fore feet; the hallux always distinct and partially opposable, though small and nailless. Tail long, very variable in its covering, being either bushy, crested, or nearly naked. Pouch represented merely by a few folds of skin. Mammæ varying from four to ten in number. The food of these animals is almost entirely insects; some species pursuing their prey among the branches of trees, while others are purely terrestrial. They are found throughout Australia, and also in New Guinea and the Aru and some of the adjacent islands.
P. cristicaudata, a species with a thick compressed tail ornamented upon its apical half with a crest of black hair, differs from the others by the very reduced size of the fourth premolar in the upper, and its complete absence in the lower jaw, thus forming an interesting transition in dentition towards Dasyurus. It constitutes the genus Chætocercus of Krefft, but is included by Mr. O. Thomas in Phascologale, the frequent absence of the fourth lower premolar in P. thorbeckiana indicating that the total absence of this tooth in the known specimens of this species cannot be regarded as of generic importance. All the members of this and the two following genera can be at once distinguished from Dasyurus by the absence of white spots on the fur.
Sminthopsis.[46]—The genus Sminthopsis includes several small species allied to Phascologale but characterised by the narrowness of the hind foot, and by the soles of the feet being either granulated or hairy, instead of naked.
Antechinomys.[47]—The last genus of the Dasyurinæ is Antechinomys, represented only by A. laniger of Queensland and New South Wales. This elegant little mouse-like creature, which has large oval ears and a long tail with the terminal part bushy, is distinguished from Sminthopsis by the absence of the hallux and the great elongation of the limbs. The tympanic bullæ of the skull are also unusually large, with the mastoid portion much swollen. A full account of the habits and anatomy of this animal, which appears to be of very rare occurrence, is given in the Proc. Zool. Soc. 1880, p. 454.
Subfamily Myrmecobiinæ.—Molars and premolars exceeding the normal number of seven on each side. Tongue, long cylindrical, and extensile.
Fig. 41.—Myrmecobius fasciatus. From Gould.
Myrmecobius.[48]—Dentition: i ⁴⁄₃, c ¹⁄₁, p ³⁄₃, m ⁵⁄₅ or ⁶⁄₆; total 52 or 56, being the largest number of teeth in any existing Marsupial. The distinction between the molars and premolars is founded not on a knowledge of the succession of the teeth, but on their form. The teeth are all small and (except the four posterior inferior molars) separated from each other by an interval. Head elongated, but broad behind. Muzzle long and pointed. Ears of moderate size, ovate, and rather pointed. Fore feet with five toes, all having strong, pointed, compressed claws, the second, third, and fourth nearly equal, the fifth somewhat, and the first considerably, shorter. Hind feet with no trace of hallux externally, but the metatarsal bone present. Tail long, clothed with long hairs. Fur rather harsh and bristly. Female without any pouch, the young when attached to the nipples being concealed only by the long hair of the abdomen. Vertebræ: C 7, D 13, L 6, S 3, C 23. A gland on the under surface of the body just in advance of the sternum.
Of this singular genus but one species is known, M. fasciatus (Fig. 41), found in western and southern Australia. It is about the size of an English squirrel, to which animal its long bushy tail gives it some resemblance; but it lives entirely on the ground, especially in sterile, sandy districts, feeding on ants. Its prevailing colour is chestnut red, but the hinder part of the back is elegantly marked with broad, white, transverse bands on a dark ground.
The special interest of this form lies in its apparent relationship to those Mesozoic mammals which possess a large number of true molars (see p. 114); and it is suggested by Thomas that it may eventually be found advisable to include some of the latter in the present subfamily.
Dentition: i ⁴⁻⁵⁄₃, c ¹⁄₁, p ³⁄₃, m ⁴⁄₄; total 46 or 48. Upper incisors small, with short broad crowns. Lower incisors moderate, narrow, proclivous. Canines well developed. Premolars compressed, pointed. Molars with quadrate tuberculated crowns. Fourth premolar preceded by a small molariform tooth, which remains in place until the animal is nearly full grown. Fore feet with two or three of the middle toes of nearly equal size, and provided with strong, sharp, slightly curved claws; the other toes rudimentary. Hind feet long and narrow; the hallux rudimentary or absent; the second and third toes very slender, and united in a common integument; the fourth very large, with a stout elongated conical claw; the fifth smaller than the fourth (see Fig. 43). The ungual phalanges of the large toes of both feet cleft at their extremities (as in Manis among the Edentata, but in no other Marsupials). Head elongated. Muzzle long, narrow, and pointed. Stomach simple. Cæcum of moderate size. Pouch complete, opening backwards. Alone among Marsupials they have no clavicles.
The Peramelidæ form a very distinct family, in some respects intermediate between the sarcophagous Dasyuridæ and the phytophagous Macropodidæ. In dentition they resemble the former, but they agree with the latter in the peculiar structure of the hind feet. In the construction of the fore feet they differ from all other Marsupials.
The Bandicoots, as these Marsupials are popularly termed, are of fossorial habits, and subsist either on an insectivorous or omnivorous diet. It has been generally considered that their syndactylous feet indicate direct affinity with the Diprotodonts, but owing to the essentially Polyprotodont character of the organisation—which extends even to their carpal and tarsal bones—Thomas dissents from this view, and concludes that their syndactylism is an independently acquired character, and that they are really a direct offshoot from the Dasyuridæ. Some individuals are remarkable for the presence of a longitudinal groove in the root of the canines, by which feature they approximate to some of the Mesozoic Polyprotodont forms. They may be divided into three genera.
Fig. 42.—Perameles gunni. From Gould.
Perameles.[49]—Anterior and posterior extremities not differing greatly in development. Fore feet with the three middle toes well developed, the third slightly larger than the second, the fourth somewhat shorter, provided with long, strong, slightly curved, pointed claws. First and fifth toes very short and without claws. Hind feet with hallux of one or two phalanges, forming a distinct tubercle visible externally; the second and third toes very slender, of equal length, joined as far as the ungual phalanges, but with distinct claws; the fifth intermediate in length between these and the largely developed fourth toe. Ears of moderate or small size, ovate, pointed. Tail rather short, clothed with short adpressed hairs. Fur short and harsh. Vertebræ; C 7, D 13, L 6, S 1, C 17. Skull long and narrow, with the bulla single, and its mastoid portion not inflated.
The animals of this genus are all small, and live entirely on the ground, making nests composed of dried leaves, grass, and sticks in hollow places. They are rather mixed feeders; but insects, worms, roots, and bulbs constitute their ordinary diet. The various species are widely distributed over Australia, Tasmania, New Guinea, and several of the adjacent islands, as Aru, Kei, and New Ireland. The best known are—P. gunni (Fig. 42), bougainvillei, nasuta, obesula, and macrura from Australia, and P. doreyana, raffrayana, and longicaudata from New Guinea.
Remains apparently referable to existing species are found in the cave-deposits of New South Wales.
Peragale.[50]—Molar teeth curved, typically with longer crowns and shorter roots than in the last. Hinder extremities proportionally longer, and hallux without claw. Muzzle much elongated and narrow. Fur soft and silky. Ears very large, long, and pointed. Tail long, its apical half clothed on the dorsal surface with long hairs which form a crest. Vertebræ: C 7, D 13, L 6, S 2, C 23. Skull distinguished from that of Perameles by the large size and double structure of the auditory bulla, of which the mastoid portion is inflated. There is also an abrupt contraction of the muzzle at the third premolar.
The type species of Rabbit-Bandicoot (P. lagotis), as these animals are called, is found in Western Australia, and also occurs fossil in the cave-deposits of New South Wales. It is the largest member of the family, being about the size of the common Rabbit, to which animal it bears sufficient superficial resemblance to have acquired the name of “Native Rabbit” from the colonists. It burrows in the ground, but in other respects resembles the true Bandicoots in its habits.
The smaller P. leucura has short-crowned molars, with distinct cusps, which are almost obsolete in the type species.
Fig. 43.—Skeleton of right hind foot of Chœropus castanotis. c, Calcaneum; a, astragalus; cb, cuboid; n, navicular; c³, ectocuneiform; II and III, the conjoined second and third digits; IV, the large and only functional digit; V, the rudimentary fifth digit.
Chœropus.[51]—Dentition generally resembling that of Perameles, but the canines are less developed, and in the upper jaw two-rooted. Limbs very slender; posterior nearly twice the length of the anterior. Fore feet with the functional toes reduced to two, the second and third, of equal length, with closely united metacarpals and short, sharp, slightly curved, compressed claws. First toe represented by a minute rudiment of a metacarpal bone; the fourth by a metacarpal and two small phalanges without a claw, and not reaching the middle of the metacarpal of the third; fifth entirely absent. Hind foot (Fig. 43) long and narrow, mainly composed of the strongly developed fourth toe, terminating in a conical pointed nail, with a strong pad behind it; the hallux absent or represented by a rudimentary metatarsal; the remaining toes completely developed, and with claws, but exceedingly slender; the united second and third reaching a little way beyond the metatarso-phalangeal articulation of the fourth; the fifth somewhat shorter. Tail not quite so long as the body, and covered with short hairs forming a slight crest. Ears large and pointed, and folded down when the animal is at rest. Fur soft and loose. Vertebræ: C 7, D 13, L 6, S 1, C 20. Skull short and wide, with a small and single bulla, and a contraction of the muzzle at the third premolar.
The only known species of this genus (Fig. 44), chiefly remarkable for the singular construction of its limbs, is an animal about the size of a small Rat, found in the interior of the Australian continent. Its general habits and food appear to resemble those of the other Peramelidæ. It was first described as C. ecaudatus by Ogilby from a mutilated specimen, but the specific name was afterwards changed, as being inappropriate, by Gray to castanotis.
For the leading characters of this group, see page 132.
Dentition: c ¹⁄₁, i ⁰⁄₀, p ¹⁄₁, m ⁴⁄₄ = 24. All the teeth with persistent pulps. The incisors large, scalpriform, with enamel only on the front surface, as in the Rodentia. The molars strongly curved, forming from the base to the summit about a quarter of a circle, the concavity being directed outwards in the upper and inwards in the lower teeth. The first of the series, or premolar, appears to have no milk-predecessor, and is single-lobed; the other four composed of two lobes, each subtriangular in section. Limbs equal, stout, and short. Fore feet with five distinct toes, each furnished with a long, strong, and slightly curved nail, the first and fifth considerably shorter than the other three. Hind feet with a very short nailless hallux, the second, third, and fourth toes partially united by integument, of nearly equal length, the fifth distinct and rather shorter; all four provided with long and curved nails. In the skeleton of the foot, the second and third toes are distinctly more slender than the fourth, showing a slight tendency towards the peculiar character so marked in the next two families. Tail rudimentary. Stomach simple, provided with a special gland situated near the cardiac orifice. Cæcum very short, wide, and with a peculiar vermiform appendage. Pouch present. The auditory bullæ of the skull are imperfect, open behind, with their anterior wall formed by a descending process of the squamosal, instead of the alisphenoid. Masseteric fossa of mandible with a perforation and a deep pit.
Fig. 44.—Chœropus castanotis. From Gould.
Phascolomys.[52]—The existing Wombats (Fig. 45) comprise three species, all of which are included in the one genus Phascolomys, and all of which date from the Pleistocene.
In the typical group we find the following characters. Fur rough and coarse. Ears short and rounded. Muffle naked. Postorbital process of the frontal bone obsolete. Ribs fifteen pairs. Vertebræ: C 7, D 15, L 4, S 4, C 10-12. The Wombat of Tasmania and the islands of Bass’s Straits (P. ursinus) and the closely similar but larger animal of the southern portion of the mainland of Australia (P. mitchelli) belong to this group.