Fig. 45.—Common Wombat (Phascolomys ursinus).
In the second group the characters are as follows. Fur smooth and silky. Ears large and more pointed. Muffle hairy. Frontal region of skull broader than in the other group, with well-marked postorbital processes. Ribs thirteen. Vertebræ: C 7, D 13, L 6, S 4, C 15-16. One species, P. latifrons, the Hairy-nosed Wombat of Southern Australia.
In their general form and actions the Wombats resemble small bears, having a somewhat similar shuffling manner of walking, but they are still shorter in the legs, and have broader, flatter backs than bears. They live entirely on the ground, or in burrows or holes among rocks, never climbing trees, and feed entirely on grass, roots, and other vegetable substances. They sleep during the day, and wander forth at night in search of food, and are shy and gentle in their habits generally, though they can bite strongly when provoked. The only noise the common wombat makes is a low kind of hissing, but the Hairy-nosed Wombat is said to emit a short quick grunt when annoyed. The prevailing colour of the last-named species, as well as of P. ursinus of Tasmania, is a brownish gray. The large wombat of the mainland is very variable in colour, some individuals being found of a pale yellowish brown, others dark gray, and some quite black. The length of head and body is about three feet.
It is noteworthy that P. mitchelli was first described from the evidence of fossil remains, the living form subsequently described as P. platyrhinus being found to be indistinguishable. Other extinct species occur in the Pleistocene of Australia.
Phascolonus.[53]—Remains of a large extinct Wombat, which must have nearly equalled the dimensions of a Tapir, occur in the Pleistocene of Queensland, and have been described as Phascolonus. It is probable that the expanded and flattened upper incisors from the same deposits upon the evidence of which the presumed genus Sceparnodon was founded, are likewise referable to the same form. The characters of both the upper and lower incisors distinguish Phascolonus from Phascolomys.
Dentition extremely variable, owing to the presence of minute rudimental teeth not constant in the same species, or even in the two sides of the jaws of the same individual; exclusive, however, of Tarsipes, the formula i ³⁄₁, c ¹⁄₀, p ²⁻³⁄₀₋₂, m ³⁻⁴⁄₃₋₄ represents fairly the general condition of the functional teeth. First incisors long and stout; the lower pair very large and pointed, but without the scissor-like action found in the existing Macropodidæ; second and third lower incisors minute and probably functionless. Fourth premolar generally secant; milk-molar generally minute and deciduous at an early period. Molars either with sharp cutting-crests or bluntly tuberculate; fourth sometimes absent. Mandible without pit, and at most a very minute perforation in the masseteric fossa. Limbs subequal. Fore feet with five distinct, subequal toes, furnished with claws. Hind feet short and broad, with five well developed toes; the hallux large, nailless and opposable; the second and third slender, and united by a common integument as far as the claws. Tail generally long, and frequently more or less prehensile. Stomach simple. Cæcum present (except in Tarsipes), and usually large. Pouch complete. Animals of small or moderate size and arboreal habits, usually feeding on a vegetable or mixed diet, inhabiting Australia and the Papuan Islands.
The homologies of the lower functionless teeth between the first incisor and fourth premolar are very difficult to determine, but it is probable that one represents a canine only when the largest known number is present; this tooth, according to Mr. Thomas, being the first to disappear.
Phalangers are small woolly-coated animals, with long, powerful, and often prehensile tails, large claws, and, as in the American opossums, with opposable nailless great toes. Their expression seems in the day to be dull and sleepy, but by night they appear to decidedly greater advantage. They live mostly upon fruit, leaves, and blossoms, although some few feed habitually upon insects, and all relish, when in confinement, an occasional bird or other small animal. Several of the Phalangers possess flying membranes stretched between their fore and hind limbs (Fig. 48), by the help of which they can make long and sustained leaps through the air, like the Flying Squirrels, but it is interesting to notice that the possession of these flying membranes does not seem to be any indication of special affinity, the characters of the skull and teeth sharply dividing the flying forms, and uniting them with other species of the non-flying groups. Their skulls (Fig. 47) are as a rule broad and flattened, with the posterior part swollen out laterally, owing to the numerous air-cells situated in the substance of the squamosal.
The Phalangers are interesting from an historical point of view, since the Gray Cuscus (Phalanger orientalis) was the first of the Marsupials of the eastern hemisphere brought to the notice of Europeans, having been described in a work published at Leyden in 1611, from an account of a specimen seen at Amboyna during the third expedition of Admiral Van der Hagen.
The present family corresponds to the Dasyuridæ among the Polyprotodonts as presenting, on the whole, the most generalised types of the suborder. The existing forms may be divided into three subfamilies.
Subfamily Tarsipedinæ.—Cheek-teeth almost rudimentary and variable in number. Tongue long, slender, pointed, and very extensile. Tail long. Cæcum absent.
Fig. 46.—Tarsipes rostratus. From Gould.
Tarsipes.[54]—So named from some supposed resemblance of its foot to that of the Lemurine genus Tarsius; but it must be remarked that it has none of the peculiar elongation of the calcaneum and navicular so characteristic of that genus. Head with elongated and slender muzzle. Mouth-opening small. The two lower incisors are long, very slender, sharp-pointed, and horizontally placed. All the other teeth are simple, conical, minute, and placed at considerable and irregular intervals apart in the jaws, the number appearing to vary in different individuals and even on different sides of the same individual. The formula, in a specimen in the Museum of the Royal College of Surgeons, is i ²⁄₁, c ¹⁄₀, p and m ³⁄₂ on one side, and ⁴⁄₃ on the other; total 20. Rami of the mandible extremely slender, nearly straight, and without coronoid process or inflected angle. Fore feet with five well-developed toes, furnished with small, flat, scale-like nails, not reaching to the extremity of the digits. Hind feet rather long and slender compared with those of the Phalangerinæ, having a well-developed opposable and nailless hallux; second and third digits syndactylous, with sharp compressed curved claws; the fourth and fifth free, and with small flat nails. Ears of moderate size and rounded. Tail longer than the body and head, scantily clothed with short hairs, prehensile. Vertebræ: C 7, D 13, L 5, S 3, C 24.
Of this singular genus but one species, T. rostratus (Fig. 46), is known, about the size of a common Mouse. It inhabits Western Australia, lives in trees and bushes, uses its tail in climbing, and feeds on honey, which it procures by inserting its long tongue into the blossoms of Melaleucæ, etc. One kept in confinement by Mr. Gould was also observed to eat flies.
Subfamily Phalangerinæ.—Teeth normal. One or more rudimentary teeth between the upper canine and fourth premolar, and between the first lower incisor and fourth premolar. Tongue of ordinary structure. No cheek-pouches. Stomach and ascending colon simple. Cæcum long, simple. Tail well-developed, generally prehensile.
A numerous group of animals, varying from the size of a mouse to that of a large cat, arboreal in their habits, and abundantly distributed throughout the Australian region. The members of this group are the typical representatives of the family, and are commonly known to the colonists as Opossums.
Phalanger.[55]—The typical genus Phalanger (Cuscus) presents the following characters. No flying membrane; size large or medium, and build stout and clumsy; fur thick and woolly. Ears short or medium, hairy externally, and in some cases also internally. Toes of fore feet subequal, their relative lengths in the order 4, 3, 5, 2, 1. Claws long, stout, and curved. Soles of feet naked and striated, with large ill-defined pads. Tail stout and markedly prehensile, with the proximal half furred like the body, and the terminal portion entirely naked. Four mammæ. Skull (Fig. 47) stout and strong, with large vacuities in the hinder half of the palate, and the auditory bullæ thick and inflated. Dentition usually i ³⁄₂, c ¹⁄₀, p ³⁄₃, m ⁴⁄₄. First upper incisor with nearly circular section, or only slightly flattened in front; canine more or less closely approximated to third incisor (which is very small), and situated partly in front of the suture between the premaxilla and maxilla. Fourth premolar large, secant, and placed obliquely to line of molars. Molars four-cusped, with the inner cusps of the upper ones crescentoid, and imperfect transverse ridges connecting each pair of cusps.
Fig. 47.—Left lateral view of skull of Gray Cuscus (Phalanger orientalis). After Peters.
The Cuscuses are curious sleepy-looking animals, inhabiting the various islands of the East Indian Archipelago as far west as Celebes, and being the only Marsupials found west of New Guinea. As already noted, it was a member of this genus, the Gray Cuscus (P. orientalis), a native of Amboyna, Timor, and the neighbouring islands, which was the first Australasian Marsupial known to European naturalists. There are altogether five species known, all of about the size of a large cat; their habits resemble those of other Phalangers, except that they are said to be somewhat more carnivorous.
Trichosurus.[56]—The members of the genus Trichosurus are of relatively large size, and are distinguished from Phalanger by the following characters. Ears more or less hairy behind. Relative lengths of toes of fore feet in the order 4, 3, 2, 5, 1. Hair on the soles of the hind feet beneath the heel, but not elsewhere. Tail thick, not tapering, covered with bushy hair up to the extreme tip, which is naked, but with a naked strip on the inferior surface in the distal third or half. A gland on the chest. Dentition usually i ³⁄₂, c ¹⁄₀, p ²⁄₂, m ⁴⁄₄. Upper incisors of nearly uniform length, the first much flattened in front. Canine situated some distance behind the third upper incisor, which it scarcely exceeds in size. Last premolar and molars very similar to those of Phalanger.
The true Phalangers comprise two species, of which the best known is the Vulpine Phalanger (T. vulpecula), so common in zoological gardens, where, however, it is seldom seen, owing to its nocturnal habits. It is of about the size and general build of a small fox, whence its name. In the typical variety the colour is gray, with a yellowish white belly, white ears, and a black tail. This variety is a native of the greater part of the continent of Australia, but is replaced in Tasmania by the closely allied Brown Phalanger (var. fuliginosa). Its habits are very similar to those of the Yellow-bellied Flying-Phalanger (Petaurus australis) described below, except that it is unable to take the flying leaps of that animal. Like all the other phalangers, its flesh is freely eaten both by the natives and the lower class of settlers.
Pseudochirus.[57]—The genus Pseudochirus agrees with the preceding in the absence of a flying membrane, and presents the following leading characters. Size large or medium. Fur comparatively short and woolly. Ears medium or short, hairy behind, although seldom closely furred over all this aspect. Claws medium. Fore toes subequal, the first two distinctly opposable to the other three. Soles of feet naked, with large, striated, round pads, and hair beneath the heels. Tail tapering, markedly prehensile, with its distal third and the whole of the under surface short-haired; tip naked underneath for a short distance. Four mammæ. No gland on chest. Skull with larger nasals than in the preceding genera; the posterior part of the palate in most cases fully ossified, and the auditory bullæ generally somewhat inflated. Dentition (at most) i ²⁻³⁄₂, c ⁰⁻¹⁄₀, p ³⁄₃, m ⁴⁄₄. Upper teeth nearly uniform in length, but the first incisor distinctly longer than second. Upper premolars variable. Molars with both inner and outer cusps distinctly crescentoid, and recalling those of the Selenodont Artiodactyle Ungulates.
Range.—Tasmania, Australia, and New Guinea.
There are about ten species of this genus known, of which the commonest is Cook’s Ring-tailed Phalanger (Pseudochirus peregrinus), an animal discovered by Captain Cook during his first voyage, at Endeavour river, North Queensland.
The complex and sub-selenodont character of the molars of this and the following genus readily distinguish them from the more typical Phalangers, and show an approximation to the type of dentition prevailing in Phascolarctus; according, however, to Mr. O. Thomas, a tendency towards the same structure is observable in unworn molars of young Cuscuses. The genus may be divided into three groups, of which the first, as typified by the common P. peregrinus, is restricted to Australia and Tasmania, while the third, as represented by P. canescens, is only found in New Guinea. P. albertisi may be taken as the type of the second group, which is represented by that species in New Guinea, and by P. archeri in Queensland. With the exception of P. peregrinus, the species have a more or less restricted range. Remains of Pseudochirus, probably referable to existing species, are found in the cave-deposits of New South Wales.
Petauroides.[58]—With the genus Petauroides, containing only the single species P. volans, we come to the first of the Flying-Phalangers, characterised by the possession of a living membrane along the flanks. The characters of this genus are as follows. Size large. Fur very long and silky. Ears large and oval, thickly furred on the back, but naked internally. Flying-membrane reaching from wrist to ankle, but very narrow along the sides of the forearm and lower leg. Fore toes subequal, their relative lengths in the order 4, 3, 5, 2, 1. Claws long, curved, and sharp. Tail long, cylindrical, and bushy, except near its tip, where it is naked and prehensile. Skull short and broad, with the nasals short, and not extending nearly as far forwards as the premaxillæ. Large vacuities in hinder part of palate. Auditory bullæ inflated and smooth. Dentition usually i ³⁄₂, c ¹⁄₀, p ²⁄₂, m ⁴⁄₄. General characters of teeth very similar to those of Pseudochirus, but the first upper incisor scarcely longer than the second.
The single species is found in Australia, from Queensland to Victoria, and is commonly known as the Taguan Flying-Phalanger. The structure of the skull and teeth indicates close affinity with Pseudochirus, although the external form is widely different in the two genera. This Phalanger seems, indeed, to be, so to speak, a very specialised Pseudochirus, in which the teeth have become somewhat further diminished and the flying membrane has been developed.
Dactylopsila.[59]—The genus Dactylopsila is one of the forms without any trace of a flying membrane, its characters being as follows. Size medium. Body striped black and white. Ears oval, nearly naked at the ends. Fore toes of very unequal length, the fourth being enormously elongated; fourth and fifth toes of pes also markedly elongated. Claws long, moderately curved. Tail long, cylindrical, and evenly bushy, with the extremity more or less naked below. Skull narrow, but with the zygomatic arches greatly expanded; palate fully ossified. Dentition: i ³⁄₃, c ¹⁄₀, p ³⁄₂, m ⁴⁄₄. Upper incisors very large, the third being directed horizontally forwards; canine small and approximated to the third incisor, which it resembles. The fourth premolar of moderate size, with its longer axis placed obliquely. First lower incisor longer than in any other genus. Molars oblong, with four cusps.
The typical D. trivirgata, or Striped Phalanger, inhabits the Papuan and North Australian sub-region; a second species (D. palpator), characterised by the still greater elongation of the fourth finger, occurring in South New Guinea. These animals are said to be of insectivorous habits, the elongated fourth finger, as in the analogous instance of the Lemuroid genus Chiromys, being apparently specially adapted for extracting insects and larvæ from their hiding places.
Petaurus.[60]—Size medium or small. Fur very soft and silky. A broad flying membrane extending from the outer side of the fifth digit of the manus to the ankle. Fore toes usually increasing regularly in length from the first to the fifth, but in some of the smaller species the fourth is the longest. Claws strong, sharp, and much curved. Tail long, evenly bushy to the extremity. Glands on the chest and between the ears. Skull short and wide, with the nasals expanded posteriorly, and usually two small palatal vacuities near the second molars. Auditory bullæ inflated, and variable in size. Dentition: i ³⁄₂, c ¹⁄₀, p ³⁄₃, m ⁴⁄₄. First upper incisors very large, and taller than canine. Molars with square crowns rounded at the angles, and four cusps, except in the last, which is triangular.
This genus, which ranges from New Ireland to South Australia, but is not found in Tasmania, contains three species, the largest of which is the Yellow-bellied Flying-Phalanger (P. australis), whose habits are recorded by Mr. Gould as follows. “This animal is common in all the brushes of New South Wales, particularly those which stretch along the coast from Port Philip to Moreton Bay. In these vast forests trees of one kind or another are perpetually flowering, and thus offer a never-failing supply of the blossoms upon which it feeds; the flowers of the various kinds of gums, some of which are of great magnitude, are the principal favourites. Like the rest of the genus, it is nocturnal in its habits, dwelling in holes and in the spouts of the larger branches during the day, and displaying the greatest activity at night while running over the small leafy branches, frequently even to their very extremities, in search of insects and the honey of the newly-opened blossoms. Its structure being ill adapted for terrestrial habits, it seldom descends to the ground except for the purpose of passing to a tree too distant to be reached by flight. When chased, or forced to flight it ascends to the highest branch and performs the most enormous leaps, sweeping from tree to tree with wonderful address; a slight elevation gives its body an impetus which, with the expansion of its membrane, enables it to pass to a considerable distance, always ascending a little at the extremity of the leap; by this ascent the animal is prevented from receiving the shock which it would otherwise sustain.”
A second species, P. sciureus, in some ways one of the most beautiful of all mammals, has been chosen for the accompanying woodcut.
Gymnobelideus.[61]—Like Petaurus in every respect, but without any trace of a flying membrane, and with the fifth digit of the manus slightly shorter than the third. This genus is represented only by G. leadbeateri of Victoria, and according to Mr. Thomas, may be regarded as the primitive form from which the specialised Petaurus has been developed.
Fig. 48.—Squirrel Flying-Phalanger (Petaurus sciureus).
Dromicia.[62]—Size small, and general appearance dormouse-like. Ears large and thin, almost naked, and without internal or basal tufts. No flying membrane. Digits of normal proportions, the relative lengths of those of the manus in the order 3, 4, 2, 5, 1; fore claws rudimentary, hind ones long and sharp. Tail mouse-like, cylindrical, furry at base, the remainder scaly, with fine hairs, except at the tip, which is naked and prehensile. Skull short and broad, with the hinder part of the palate incomplete, and the auditory bullæ large, much inflated, and transparent. Dentition: i ³⁄₂, c ¹⁄₀, p ³⁄₃, m ³⁻⁴⁄₃₋₄. First upper incisor spatulate, and much longer than either of the others. Canine large, placed at some distance behind the third incisor. Molars (except the last) with evenly rounded crowns, carrying four small smooth cusps.
This genus, which occurs in New Guinea, Western Australia, and Tasmania, is represented by four species. It seems to be intermediate between Petaurus and Acrobates, and it has apparently had to yield place to those more highly organised types in regions where they have come in contact with one another.
Distœchurus.[63]—Size small. Ears rather short, thinly covered with hair, but with small tufts at the base. No flying membrane. Digits of normal proportions, without expanded terminal pads. Claws curved and sharp. Tail, skull, and dentition as in Acrobates, with the exception that the fourth premolar is small in the upper, and absent in the lower jaw.
The one species of Feather-tailed Phalanger (D. pennatus) is found in New Guinea.
Acrobates.[64]—Size very small. Ears moderate, thinly covered with hair, but with small tufts round the base and on the internal prominences. A narrow flying membrane, fringed with long hairs, running from the elbow to the flank, and from the latter to the knee. Four mammæ. Digits furnished with expanded and striated terminal pads, the relative length of those of the manus being in the order 4, 3, 5, 2, 1. Claws sharp, although somewhat concealed by the terminal pads. Tail short-haired above and below, with a broad fringe on either side. Skull short, wide, and depressed. Posterior portion of palate very imperfectly ossified; anterior palatal vacuities almost confined to the maxillæ. Auditory bullæ low, rounded, and but slightly prominent. Dentition: i ³⁄₂, c ¹⁄₀, p ³⁄₃, m ³⁄₃. Teeth sharp, and of an insectivorous type. Upper canine long, and approximated to third incisor. The three upper premolars large, functional, and taller than the molars. Molars small and rounded, with smooth unridged cusps.
There is only one species in this genus, the beautiful little Pigmy Flying-Phalanger (A. pygmæus), not so big as a Mouse, which is found in Queensland, New South Wales, and Victoria, and feeds on the honey it abstracts from flowers, and on insects. Its agility and powers of leaping are exceedingly great, and it is said by Mr. Gould to make a most charming little pet.
Subfamily Phascolarctinæ.—Teeth large, normal; no rudimentary premolars before the last upper premolar, or any teeth between the first lower incisor and fourth premolar. Tongue of ordinary structure. Distinct cheek-pouches. Stomach with a special gland near the cardiac orifice. Cæcum very long, and (with the upper portion of the colon) dilated and provided with numerous longitudinal folds of mucous membrane. In many anatomical characters, especially the possession of a special gastric gland, this group resembles the Phascolomyidæ.[65]
Fig. 49.—Skeleton of right hind foot of Koala (Phascolarctus cinereus), showing the stout opposable hallux, followed by two slender toes, which in the living animal are enclosed as far as the nails in a common integument.
Phascolarctus.[66]—Dentition: i ³⁄₁, c ¹⁄₀, p ¹⁄₁, m ⁴⁄₄; total 30. Upper incisors crowded together, cylindroidal, the first much larger than the others, with a bevelled cutting edge (Fig. 36). Canine very small; a considerable interval between it and the premolar, which is as long from before backwards but not so broad as the true molars, and has a cutting edge, with a smaller parallel inner ridge. The molars slightly diminishing in size from the first to the fourth, with square crowns, each bearing four pyramidal cusps, with curved ridges radiating from them, and having a structure very similar to these of Pseudochirus. The lower incisors are semiproclivous, compressed and tapering, bevelled at the ends. Premolars and molars in continuous series, as in the upper jaw. Milk-tooth very minute, and almost functionless. Fore feet with the two inner toes slightly separated from and opposable to the remaining three, all with strong, curved, and much compressed claws. Hind foot (Fig. 49) with the hallux placed very far back, large and broad, the second and third (united) toes considerably smaller than the other two; the fourth the largest. No external tail. Fur dense and woolly. Ears of moderate size, thickly clothed with long hairs. Vertebræ: C 7, D 11, L 8, S 2, C 8. Ribs eleven pairs, a rare exception to the usual number (13) in the Marsupialia.
There is but one species, the Koala or Native Bear of the Australian colonists (P. cinereus), an animal of comparatively large size and heavy build (Fig. 50), found in the south-eastern parts of the Australian continent. It is about two feet in length, and of an ash-gray colour, an excellent climber, and residing generally in lofty Eucalyptus trees, on the buds and tender shoots of which it feeds, though occasionally descending to the ground in the night.
Numerous imperfect remains recently described by De Vis are regarded as indicating large extinct types of Phalangeridæ, but further evidence is required before all these determinations can be definitely accepted. Thus part of an upper jaw is provisionally referred to a large species of Pseudochirus, while part of a scapula is made the type of a genus Archizonurus which appears to be allied to the former. Another fragmentary scapula is considered to indicate a large Phalanger. Finally, part of a fibula, described under the name of Koalemus is regarded as affording evidence of the former existence of a large ancestral form allied to the Koala, and it is suggested that an upper jaw with teeth may belong to the same or an allied type.
Fig. 50.—The Koala (Phascolarctus cinereus). From Sclater, Proc. Zool. Soc. 1880, p. 355.
Thylacoleo.[67]—Dentition of adult: i ³⁄₁, c ¹⁄₀, p ³⁄₃, m ¹⁄₂; total 28. First upper incisor much larger than the others; canine and first two premolars rudimentary. In the lower jaw the two small anterior premolars are functionless, and often deciduous; posterior premolars of both jaws formed on the same type as those of Potorous, but relatively much larger; true molars rudimentary, tubercular. One species, T. carnifex. This animal presents a most anomalous 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 below. It was first described as a carnivorous Marsupial, and named, in accordance with its presumed habits, “as one of the fellest and most destructive of predatory beasts”; but, as its affinities are certainly with the Phalangeridæ and Macropodidæ, and its dentition completely unlike that of any known predaceous animal, this view has been called in question.
Fig. 51.—Front view of skull of Thylacoleo carnifex, restored. ¹⁄₃ natural size. From Quart. Journ. Geol. Soc. vol. xxiv. p. 312.
The dentition is nearer to that of the existing Phalangeridæ than to that of the Macropodidæ, and the genus may be provisionally regarded as the type of a distinct subfamily of the former.
Dentition i ³⁄₁, c ⁰⁻¹⁄₀, p ²⁄₂, m ⁴⁄₄. Incisors sharp and cutting, 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. Pouch large and opening forwards.
Fig. 52.—Skeleton of right hind foot of Kangaroo.
The Macropodidæ 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 occasionally do when feeding. Rapid progression is, however, 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 supporting 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 live distinct digits, each armed with a strong curved claw.
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 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 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.
Fig. 53.—The Great Gray Kangaroo (Macropus giganteus).
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 Macropodidæ, 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 Macropodidæ, 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 Macropodidæ are divided into three well-marked sections: (1) the true Kangaroos (Macropodinæ); (2) a group consisting of smaller animals, commonly called Rat Kangaroos, or (improperly) “Kangaroo Rats,” or sometimes Potoroos; and (3) the Hypsiprymnodontinæ, 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 Potoroinæ.
This subfamily is now represented only by the genus Hypsiprymnodon,[68] 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 Macropodidæ.
Triclis.[69]—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 Macropodidæ, 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 Phalangeridæ and Macropodidæ than any other.
Subfamily Potoroinæ.—The second section or subfamily, the Potoroinæ, 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 jaws always having large, simple, compressed crowns, with a nearly straight or slightly concave free cutting edge, both outer and inner surfaces usually marked by a series of parallel, vertical grooves and ridges, these teeth being either set in the same line with the molars, or slightly bent outwards. Molars with quadrate crowns, having a blunt, conical cusp at each corner, the fourth notably smaller than the third, sometimes rudimentary, and appearing early. Fore feet narrow; three middle toes considerably exceeding the first and fifth in length; their claws long, compressed, and but slightly curved. Hind feet as in Macropus. Tail long and hairy, sometimes partially prehensile, being used for carrying bundles of grass with which these animals build their nests.
Fig. 54.—Skull and teeth of Rat Kangaroo (Bettongia lesueuiri). c, Upper canine. The other letters as in Fig. 51.
The Potoroos or Rat Kangaroos are all small animals, none of them exceeding a common rabbit in size. They inhabit Australia and Tasmania, are nocturnal, and feed on the leaves of various kinds of grasses and other plants, as well as roots and bulbs, which they dig up with their fore paws. Nine species are known, presenting a considerable range of diversity in minor characters, and admitting of being grouped in four principal sections, which may be allowed the rank of genera. These are:
Potorous.[70]—Head long and slender. Auditory bullæ somewhat inflated. Ridges on premolars few and perpendicular. Large palatine foramina. Tarsus short. Muffle naked. Three species, viz. P. tridactylus, P. gilberti, and P. platyops; the last two being confined to West Australia.
Bettongia.[71]—Head comparatively short and broad. Ears short and rounded. Auditory bullæ generally much inflated. Large palatine foramina. Tarsus long. Ridges on premolars numerous and oblique. Tail more or less prehensile, thickly haired, and the hairs on the upper surface longer than those on the lower, and forming a crest. Muffle naked. Four species, viz. B. penicillata, B. cuniculus, B. gaimardi, B. lesueuiri.
Caloprymnus.[72]—Muffle naked, as in Bettongia, but the edge of the hairy part less emarginate backwards in the middle line. Ears short, rounded, and hairy. Auditory bullæ much inflated, and of large size. Nasals larger and wider behind than in the other genera. Very long anterior palatine foramina. Limbs as in Bettongia. Tail thin, cylindrical, evenly coated with short hair, without trace of a crest. Skull broad and flat, with a remarkably short and conical muzzle. The sole representative of this genus is C. campestris of South Australia, originally referred to Bettongia.