Marsupialia
MARSUPIALIA (from Lat. marsupium, a "pouch," or "bag"), the group of mammals in which the young are usually carried for some time after birth in a pouch on the under-surface of the body of the female. The group, which has also the alternative title of Didelphia, is by some authorities regarded as a sub-class of the mammalia of equal rank with the Monotremata, while by others it is brigaded with the placentals, so that the two together form a sub-class of equal grade with the one represented by the monotremes. There is much to be urged in favour of either view; and in adopting the former alternative, it must be borne in mind that the difference between monotremes and marsupials is vastly greater than that which separates the latter from placentals. In elevating the marsupials to the rank of a sub-class the name Metatheria has been suggested as the title for the higher grade, with Marsupialia as the designation for the single order by which they are now represented. It is, however, less liable to cause confusion, and in many other ways more convenient to employ the better known term Marsupialia in both senses.
Marsupials may be defined as viviparous (that is non- egglaying) mammals, in which the young are born in an imperfect condition, and almost immediately attached to the teats of the mammary glands; the latter being generally enclosed in a pouch, and the front edge of the pelvis being always furnished with epipubic or " marsupial " bones. As a rule there is no allantoic placenta forming the means of communication between the blood of the parent and the foetus, and when such a structure does occur its development is incomplete. In all cases a more or less full series of teeth is developed, these being differentiated into incisors, canines, premolars and molars, when all are present; but only a single pair of teeth in each jaw has deciduous predecessors.
The pouch from which the marsupials take their name is supported by the two epipubic bones, but does not correspond to the temporary breeding-pouch of the monotremes. It may open either forward or backwards; and although present in the great majority of the species, and enclosing the teats, it may, as in many of the opossums, be completely absent, when the teats extend in two rows along the whole length of the under-surface of the body. Whether a pouch is present or not, the young are born in an exceedingly imperfect state of development, after a very short period of gestation, and are immediately transferred by the female parent to the teats, where they remain firmly attached for a considerable time; the milk being injected into their mouths at intervals by means of a special muscle which compresses the glands. In the case of the great grey kangaroo, for instance, the period of gestation is less than forty days, and the newly-born embryo, which is blind, naked, and unable to use its bud-like limbs, is little more than an inch in length.
As additional features of the subclass may be mentioned the absence of a corpus callosum connecting the right and left hemispheres of the brain, 1 and of a fossa in the septum between the two auricles of the heart. In the skull there are always vacuities, or unossified spaces in the bones of the palate, while the " angle," or lower hind extremity of each half of the lower jaw is strongly bent inwards so as to form a kind of shelf, and the alisphenoid bone takes a share in the formation of the tympanum, or auditory bladder, or bulla. Didelphia, the alternative name of the group was given in allusion to the circumstance that the uterus has two separate openings; while other features are the inclusion of the openings of the alimentary canal and the urino-genital sinus in a common sphincter muscle, and the position of the scrotum in advance of the penis. The bandicoots alone possess a placenta. Lastly the number of trunk- vertebrae is always nineteen, while there are generally thirteen pairs of ribs.
As regards the teeth, in all cases except the wombats the number of upper incisors differs from that of the corresponding lower teeth. As already stated, there is no vertical displacement and succession of the functional teeth except in the case of a single tooth on each side of each jaw, which is the third of the premolar series, and is preceded by a tooth having more or less of the characters of a molar (see fig. i). In some cases (as in rat-kangaroos) this tooth retains its place and function until the animal has nearly, if not quite, 1 The presence or absence of the corpus callosum has been much disputed ; the latest researches, however, indicate its absence.
attained its full stature, and is not shed and replaced by its successor until after all the other teeth, including the molars, are in place and use. In others, as the thylacine, it is rudimentary, being shed or absorbed before any of the other teeth have cut the gum, and therefore functionless. It may.be added that there are some marsupials, such as the wombat, koala, marsupial ant-eater and the dasyures, FIG. I. Teeth of Upper Jaw of Opossum (Didelphys marsupialis), all of which are unchanged, except the third 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.
in which no such deciduous tooth, even in a rudimentary state, has been discovered. In addition to this replacement of a single pair of functional teeth in each jaw, it has been discovered that marsupials possess rudimentary tooth-germs which never cut the gum. According to one theory, these rudimentary teeth, together with the one pair of functional teeth in each jaw that has vertical successors, represent the milk-teeth of placental mammals. On the other hand, there are those who believe that the functional dentition (other than the replacing premolar and the molars) correspond to the milk-dentition of placentals, and that the rudimentary tooth-germs represent a " prelacteal " dentition. The question, however, is of academic rather than of practical interest, and whichever way it is answered does not affect our general conception of the nature and relationships of the group.
Unfortunately the homology of the functional series does not by any means end the uncertainty connected with the marsupial dentition ; as there is also a difference of opinion with regard to the serial homology of some of the cheek-teeth. For instance, according to the older view, the dental formula in the thylacine or Tasmanian wolf is i. J, c, \, p. |, m. 4 =46. On the other hand, in the opinion of the present writer, this formula, so far as the cheek-teeth are concerned, should be altered to p. J, m. f , thus bringing it in accord, so far as these teeth are concerned, with the placental formula, and making the single pair of replacing teeth the third premolars. It may be added that the formula given above shows that the marsupial dentition may comprise more teeth than the 44 which form the normal full placental complement.
As regards geographical distribution, existing marsupials, with the exception of two families, Didelphyidae and Epanorthidae, are mainly limited to the Australian region, forming the chief mammalian fauna of Australia, New Guinea, and some of the adjacent islands. The Didelphyidae are almost exclusively Central and South American, only one or two species ranging into North America. Fossil remains of members of this family have also been found in Europe in strata of the Oligocene period.
History. The origin and evolution of the Australian marsupials have been discussed by Mr B. A. Bensley. In broad contrast to the views of Dr A. R. Wallace, this author is of opinion that marsupials did not effect an entrance into Australia till about the middle of the Tertiary period, their ancestors being probably opossums of the American type. They were then arboreal; but they speedily entered upon a rapid, although short-lived, course of evolution, during which leaping terrestrial forms like the kangaroos were developed. The short period of this evolution is at least one factor in the primitive grade of even the most specialized members of the group. In the advance of their molar teeth from a tritubercular to a grinding type, the author traces a curious parallelism between marsupials and placentals. Taking opossums to have been the ancestors of the group, the author considers that the present writer may be right in his view that marsupials entered Australia from Asia by way of New Guinea. On the other hand there is nothing absolutely decisive against their origin being southern.
Again, taking as a text Mr L. Dollo's view that marsupials were originally arboreal, that, on account of their foot-structure, they could not have been the ancestors of placentals, and that they themselves are degenerate placentals, Mr Bensley contrasts this with Huxley's scheme of mammalian evolution. According to the latter, the early monotremes which became specialized into modern monotremes, gave rise to the ancestors of the modern marsupials; while the modern placentals are likewise an offshoot from the ancestral marsupial strck. This phylogeny, the author thinks, is the most probable of all. It is urged that the imperfect placenta of the bandicoots instead of being vestigial, may be an instance of parallelism, and that in marsupials generally the allantois failed to form a placental connexion. Owing to the antiquity of both placentals and marsupials, the arboreal character of the feet of the modern forms of the latter is of little importance. Further, it is considered that too much weight has been assigned to the characters distinguishing monotremes from other mammals, foetal marsupials showing a monotreme type of coracoid, while it is probable that in the long run it will be found impossible to maintain the essential dissimilarity between the milk-glands of monotremes and other mammals.
Another view is to regard both marsupials and placentals as derivates from implacental ancestors more or less nearly related to the creodont carnivora, or possibly as independently descended from anomodont reptiles (see CREODONTA). Finally, there is the hypothesis that marsupials are the descendants of placentals, in which case, as was suggested by its discoverer, the placenta of the bandicoots would be a true vestigial structure.
Classification.
Existing marsupials may be divided into three main divisions or sub-orders, of which the first, or Polyprotodontia, is common to America and Australasia; the second, or Paucituberculata, is exclusively South American; while the third, or Diprotodonts, is as solely Australasian inclusive of a few in the eastern Austro-Malayan islands.
i. Polyprotodonts. The Polyprotpdonts are characterized by their numerous, small, sub-equal incisors, of which there are either five or four pairs in the upper and always three in the lower jaw, (fig. 2) and the generally strong and large canines, as well as by the From Flower, Quart. Jour. Ccol. Sac.
FIG. 2. Front View of Skull of the Tasmanian Devil (Sarcophilus ursinus) to exhibit polyprotodont type of dentition.
presence of from four to five sharp cusps or tubercles on the crown of the molars. The pouch is often absent, and may open backwards. For the most part the species are carnivorous or insectivorous.
The first family is that of the true or American opossums Didelphyidae, in which there are five pairs of upper incisors, while the feet are of the presumed primitive arboreal type, the hind foot having the four outer toes subequal and separate, with the first opposable to them all. With the exception of the water-opossum, forming the genus Chironectes, all the living members of the family may be included in the genus Didelphys. The latter may, however, be split up into several sub-generic groups, such as Metachirus, Philander, Marmosa (Micoureus or Grymaeomys), Peramys, Dromiciops, etc. The small South American forms included in Marmosa, which lack the pouch, and have numerous teats, and molar teeth of a primitive type, are doubtless the most generalized representatives of the group (see OPOSSUM; and WATER-OPOSSUM).
Nearly allied is the Australian family Dasyuridae, characterized by the presence of only four pairs of upper incisors, the generally small and rudimentary condition of the first hind toe, which can but seldom be opposed to the rest, and the absence of prehensile power in the tail; the pouch being either present or absent, and the fore feet always five-toed. The stomach is simple, and there is no caecum to the intestine, although this is present in the opossums.
The largest representative of the family is the Tasmanian wolf, or thylacine, alone representing the genus Thylacinus, in which the dentition numbers i. |,c. \, p. }, m. } = 46; with the incisors small and vertical, the outer one in the upper jaw being larger than the others. Summits of the lower incisors, before they are worn, with a deep transverse groove, dividing it into an anterior and a posterior cusp. Canines long, strong and conical. Premolars with compressed crowns, increasing in size from before backwards. Molars in general characters resembling those of Sarcophilus, but of more simple form, the cusps being less distinct and not so sharply pointed. Deciduous FIG. 3. The Tasmanian Wolf, or Thylacine (Thylacinus cynocephalus).
molar very small, and shed before the animal leaves the mother's pouch. General form dog-like, with the head elongated, the muzzle pointed, and the ears moderate, erect and 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. First hind toe (including the metacarpal bone) absent. Vertebrae: C. 7, D. 13, L. 6, S. 2, Ca. 23. Marsupial bones unossified. The gradual passage of the thick root of the tail into the body is a character common to the Tasmanian wolf and the aard-aark, and may be directly inherited from reptilian ancestors (see THYLACINE).
The next genus is represented solely by the Tasmanian devil, Sarcophilus (or Diabolus) ursinus, a medium-sized animal with a dental formula similar to that of the dasyures, but with teeth (fig. 2) approximating to those of the thylacine, though markedly different in details. The first hind toe is absent.
In the " native cats," or dasyures, constituting the genus Dasyurus, the dental formula is i._j, c. \, p. I, m. f : total 42. The upper Incisors are nearly equal and vertical, with the first slightly longer, narrower, and separated from the rest. Lower incisors sloping forward and upward. Canines large and sharply pointed. First two premolars with compressed and sharp-pointed crowns, and slightly developed anterior and posterior accessory basal cusps. Molars with numerous sharp-pointed cusps. In the upper jaw the first two with crowns having a triangular free surface; the last small, simple, narrow and placed transversely. In the lower jaw the molars more compressed, with longer cusps; the last not notably smaller than the others. Ears of moderate size, prominent and obtusely pointed. First hind toe rudimentary, clawless or absent; its metatarsal bone always present. Tail generally long and well clothed with hair. Vertebrae: C. 7, D. 13, L. 6, S. 2, Ca. 18-20 (see DASYURE).
The genus Phascologale comprises a number of small marsupials, none exceeding a rat in size, differing from the dasyures in possessing an additional premolar the dentition being i. }, c. \, p. J, m. | : total 46 and in having the teeth generally developed upon an insectivorous rather than a carnivorous pattern, the upper middle incisors being larger and inclined forward, the canines relatively smaller, and the molars with broad crowns, armed with 'prickly tubercles. The muzzle is pointed. Ears moderately rounded, and nearly naked. Fore feet with five sub-equal toes, with compressed, slightly curved pointed claws. Hind feet with the four outer toes sub-equal, with claws similar to those in the fore feet; the first toe alrrost always distinct and partially opposable, though small and nailless, sometimes absent.
In some respects intermediate between the preceding and the next genus is Dasyuroides byrnei, of Central Australia, an animal of the size of a rat, with one lower premolar less than in PhascologaU, without the first hind toe, and with a somewhat thickened tail. The pouch is incomplete, with two lateral folds, and the number of teats six.
Sminihopsis includes several very small species, with the same dental formula as Phascologale, but distinguished from that genus by the narrowness of the hind foot, in which the first toe is present, and the granulated or hairy (in place of broad, smooth and naked)
soles. A pouch is present, and there are eight or ten teats. Nearly allied is the jumping Antechinomys laniger, of East Central Australia, an elegant mouse-like creature, with large oval ears, elongated limbs, a long and tufted tail and no first hind toe. In connexion with the large size of the ears is the excessive inflation of the auditory bulla of the skull.
From all other members of the family the marsupial, or banded, ant-eater (Myrmecobius fasciatus) differs by the presence of more than seven pairs of cheek-teeth in each jaw, as well as by the exceedingly long and protrusile tongue. Hence it is made the type of a distinct subfamily, the Myrmecobiinae, as distinct from the Dasyurinae, which includes all the other members of the family. From the number of its cheek-teeth, the banded ant-eater has been regarded as related to some of the primitive Jurassic mammals; but this view is disputed by Mr Bensley, who regards this multiplicity of teeth as a degenerate feature. On the other hand, it is noteworthy that this marsupial retains in its lower jaw the so-called mylo-hyoid groove, which is found in the aforesaid Jurassic mammals. Myrmecobius has a total of 52 or 54 teeth, which may be classed as i. f, c. \, p. + m. |^|. 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 first toe externally, but the metatarsal bone is present. Tail long, clothed with long hairs. Fur rather harsh and bristly. Female without pouch, the young when attached to the nipples being concealed by the long hair of the abdomen. Vertebrae: C. 7, D. 13, L. 6, S. 3, Ca. 23.
From Gould.
FIG. 4. The Marsupial or Banded Ant-eater (Myrmecobius fasciatus).
The single species, which is a native of western and southern Australia, is about the size of an English squirrel, to which 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 marked with broad, white, transverse bands on a dark ground.
With the bandicoots, or Peramelidae, we come to a family of polyprotpdpnts which resemble the diprotodonts in the peculiarly specialized structure of their hind limbs; an adaptation which we must apparently' regard as having been independently acquired in the two groups. The dentition is i. |,.c. }, p. f, m. jf; total, 48; the upper incisors being small, with short, broad crowns; the lower incisors moderate, narrow, proclivous; canines well developed. Premolars compressed, pointed; and the molars with quadrate tuberculated crowns. Deciduous premolar preceded by a minute 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 first toe rudimentary or absent; the second and third 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. 6). The terminal phalanges of the large toes of both feet cleft at their extremities. Head elongated, with the muzzle long, narrow and pointed. Stomach simple. Caecum of moderate size. Pouch complete, generally opening backwards. Alone among marsupials bandicoots have no clavicles. More remarkable still is the development of a small allantoic placenta.
In the true bandicoots of the genus Perameles (fig. 5) the fore-feet have 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 one or two phalanges, in the first toe forming a distinct tubercle visible externally; the second and third toes very slender, of equal length, joined as far From Gould.
FIG. 5. Gunn's Bandicoot (Perameles gunni).
as the terminal phalange, 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 depressed hairs. Fur short and harsh. Pouch opening backwards. Vertebrae: C. 7, D. 13, L. 6, S. I, Ca. 17. (see BANDICOOT.)
The rabbit-bandicoot, Peragale (or Thylacomys) represents a genus in which the cheek-teeth are curved, with longer crowns and shorter roots than in the last. Hind extremities proportionally longer with inner toe represented only by a small metatarsal bone. 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. Pouch opening forwards. Vertebrae: C. 7, D. 13, L. 6, S. 2, Ca. 23.
The one species, from Western Australia, is the largest member of the family, being about the size of a rabbit, to which 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 bandicoots in habits.
In the pig-footed bandicoot (Choeropus castanotis) the dentition generally resembles 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 long and narrow, mainly composed of the strongly developed no/is. fourth toe, terminating in a conical pointed nail, c - calcanium ; with a strong pad behind it; the first toe repre- J,bo?d; T 15 ; na risented by a rudimentary metatarsal; the remaining cuiar ; '<:>, ' cctocutoes completely developed, with claws, but exceed- neiform ; II. and ingly slender; the united second and third reaching b$* *$!* a little way beyond the metatarso-phalangeal articu- dj g i ts ; iv. the lation of the fourth; the fifth somewhat shorter, large and only func- Tail not quite so long as the body, and covered tio " al <*'' : v -JEf . , _.?_ r^i j.^j j rudimentary nitn with short hairs. Ears large and pointed, and folded down when the animal is at rest. Fur soft and loose. Pouch opening backwards. Vertebrae : C. 7i D- I3> L. 6, S. i, Ca. 20.
The only species of this genus is about the size of a small rat, found in the interior of Australia. Its general habits and food appear to resemble those of other bandicoots. A separate family, Notoryctidae, is represented by the marsupial mole (Nptoryctes typhlcps), of the deserts of south Central Australia, a silky, golden-haired, burrowing creature, with a curious leathery muzzle, and a short, naked stumpy tail. The limbs are five-toed, with the third and fourth toes of the front pair armed with enormous digging claws; F '5y. 6 j~:
there are no external ear-conchs ; and the dentition includes four pairs of upper, and three of lower, incisors, and distinctly tritubercular cheek-teeth. The small pouch, supported by the usual epipubic bones, opens backwards. In correlation with its burrowing habits, some of the vertebrae of the neck and of the loins are respectively welded together. The eyes have degenerated to a greater extent than From Gould. FIG. 7. The Pig-footed Bandicoot (Choeropus castanotis).
those of any other burrowing mammal, the retina being reduced to a mass of simple cells, and the cornea and sclerotic (" white ") to a pearshaped fibrous capsule enclosing a ball of pigment. The reason for this extreme degeneration is probably to be found in the sandy nature of the soil in which the creature burrows, a substance which would evidently irritate and inflame any functional remnant of an eye. The portion of the lachrymal duct communicating with the cavity of the nose has, on the other hand, been abnormally developed, apparently for the purpose of cleansing that chamber from particles of sand which may obtain an entrance while the animal is burrowing. (See MARSUPIAL MOLE.)
2. Paucituberculates. The second suborder of marsupials, the Paucituberculata, is exclusively South American, and typically represented by the family Epanorthidae, the majority of the members of which are extinct, their remains being found in the probably Miocene Santa Cruz beds of Patagonia, although one existing genus (Caenolestes) survives in Ecuador and Colombia. One of the two living species was, indeed, described so long ago as the year 1863, under the preoccupied name of Hyracodon, but attracted little or no attention, as its affinities were not fully recognized. Externally Caenolestes has a shrew-like appearance. The elongated skull (fig. 8) has four pairs of upper incisors and long upper canines, while in the lower jaw there is a single pair of procumbent incisors.
Alter Thomas.
FIG. 8. Skull of Caenolestes obscurus.
followed by several small teeth representing the canine and earlier premolars. The three pairs of molars in each jaw are, like the last premolar, quadritubercular oblong teeth. The five-toed feet are of normal structure, and the rat-like tail is prehensile towards the tip. The female has a small pouch. The extinct members of the family are represented by the genera Epanorthus, Acdestis, Garzonia, etc. In a second family Abderitidae also from the Patagonian Miocene, the penultimate premolar is developed into an enormous tooth, with a tall, secant and grooved crown, somewhat after the fashion of the enlarged premolar of Plagiaulax. From the structure of the skull, it is thought probable that Abderitfs had an elongated snout, like that of many Insectivora. As a sub-order, the Paucituberculata are characterized by the presence of four pairs of upper and three of lower incisor teeth ; the enlargement and forward inclination of the first pair of lower incisors, and the presence of four or five sharp cusps on the cheek-teeth, coupled with the absence of " syndactylism ' in the hind limbs.
3. Diprotodonts. The third and last sub-order of marsupials is the Diprotodontia, which is exclusively Australasian and includes the wombats, koala, cuscuses, kangaroos and their relatives. There are never more than three pairs of upper and one of lower incisors, of which the middle upper and the single lower pair are large and chisel-like (fig. 9); the canines are small or absent; the cheek-teeth have bluntly tuberculate or transversely-ridged crowns in most cases; and the hind-feet are syndactylous. With one exception, the intestine has a caecum, and the pouch is large and opens forwards. It should be added that Professor Elliot Smith has pointed out a certain peculiarity in its commissures whereby the brain of the diprotodonts differs markedly from that of the polyprotodonts From Flower, Quart. Journ. Geol. Soc.
FIG. 9. Front view of Skull of the Koala (Phascolarctus cinereus) to exhibit Diprotodont type of dentition.
and approximates to the placental type. Dr Einar Lonnberg has also recorded certain adaptive peculiarities in the stomach. Most of the species, particularly the specialized types, are more or less completely herbivorous.
The first family, Phascolomyidae, is typified by the wombats; but according to the view adopted by Mr H. Winge, and endorsed by Professor Max Weber, is also taken to include the koala. In this wider sense the family may be characterized as follows. The tympanic process of the alisphenoid bone of the skull is short, not covering the cavity of the tympanum, nor reaching the paroccipital process. The tail is rudimentary, the first hind-toe opposable, the first pair of upper incisors very large, but the second and third either absent or small and placed partially behind the larger pair ; and only five pairs of cheek-teeth in each jaw. The stomach has a cardiac gland, and the number of teats is two.
In the wombats (Phascolomys) the dentition is i. }, c. j, p. + m. |, total 24; all the teeth growing from persistent pulps, and the incisors large and chisel-like, with enamel only on the front surface. The cheek-teeth 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 (which appears to have no predecessor) 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 first toe, the second, third and fourth toes partially united by integument, of nearly equal length, the fifth distinct and rather shorter; all four with long and curved nails. In the skeleton the second and third toes are distinctly more slender than the fourth, showing a tendency towards the character so marked in the following families. Tail rudimentary. Caecum very short and wide, with a vermiform appendage (see WOMBAT).
In addition to remains referable to the existing genus, the Pleistocene deposits of Australia have yielded evidence of an extinct giant wombat constituting the genus Phascolonus (Sceparnodon).
The koala, or" native bear " (Phascolarctus cinereus), which differs widely from the wombats in its arboreal habits, is less specialized as regards its dentition, of which the formula is i. ?, c. J, p. + m. I, total 30. Upper incisors crowded together, cylindroidal, the first much larger than the others, with a bevelled cutting edge (fig. 9). Canine very small ; a considerable interval between it and the first premolar, which is as long from before backwards but not so broad as the molars, and has a cutting edge, with a smaller parallel inner ridge. The molar-like teeth slightly diminishing in size from the first to the fourth, with square crowns, each bearing four pyramidal cusps. The lower incisors are partially inclined forwards, compressed and tapering, bevelled at the ends. Cheek-teeth in continuous series, as in the upper jaw. 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-toot (fig. 10) with the first toe placed 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 hair. Caecum very long and dilated, with numerous folds. Vertebrae: C. 7, D. 1 1, L. 8, S. 2, Ca. 8. Ribs eleven pairs (see KOALA).
Here may be noticed three genera of large extinct marsupials from the Pleistocene of Australia whose affinities appear to ally them to the wombat-group on the one hand and to the phalangers on the other. The longest known is Diprotodon, an animal of the size of a rhinoceros, with a dental formula of i. f, c. $, p. |, m. f , total 28. The first upper FIG. 10. Skeleton of Right incisor very large and chisel-like, Hind-Foot of Koala (Phas- molars with prominent transverse colarctus cinereus), showing ridges, as in Macropus, but without stout opposable hallux, fol- the longitudinal connecting ridge, lowed by two slender toes, Complete skeletons disinterred by which in the living animal are Dr E. C. Stirling indicate that in the enclosed as far as the nails in structure of the feet this creature a common integument. presents resemblances both to the wombats and the phalangers, but is nearer to the former than to the latter. On the other hand, the considerably smaller Nototherium, characterized by its sharp and broad skull and smaller incisors, seems to have been much more wombat-like, and may perhaps have possessed similar burrowing habits.
The last of the three is Thylacoleo carnifex, so named on account of its supposed carnivorous habits. In the adult the dentition (fig. 1 1) is i. f , c. i, p.+m. j, total 24. The first upper incisor is much larger than the others; canine and first two premolars rudimentary. In the lower jaw there are also one or two small and early deciduous premolars; third premolars of both jaws formed on the same type as that of the rat- kangaroos, but relatively much larger; molars rudimentary, tubercular. The functional teeth are reduced to one From Flower, Quart. Journ. Geol. Soc. FIG. II. Front view of Skull of Thylacoleo carnifex, restored.
pair of large cutting incisors situated close to the middle line, and one great, cutting, compressed premolar, on each side above and below. As already mentioned, Thylacoleo was originally regarded as a carnivorous creature, but this view was subsequently disputed, and its diet supposed to consist of soft roots, bulbs and fruits, with an occasional small bird or mammal. Recently, however, the pendulum of opinion has swung back towards the original view: and Dr R. Broom believes Thylacoleo to have been " a purely carnivorous animal, and one which would be quite able to, and probably did, kill animals as large or larger than itself." The affinities of the creature are clearly with the phalangers.
By means of the little musk-kangaroo, the cuscuses and phalangers. constituting the family Phalangeridae, are so closely connected with the kangaroos, or Macropodidae, that in the opinion of some naturalists they ought all to be included in a single family, with three sub-families. Theoretically, no doubt, this is correct, but the typical members of the two groups are so different from one another that, as a matter of convenience, the retention of the two families seems advisable. From the Phascolomyidae, the two families, which may be collectively designated Phalangeroidea, differ by the circumstance that in the skull the tympanic process of the alisphenoid covers the tympanic cavity and reaches the paroccipital process. The tail is long and in some cases prehensile; the first hind-toe may be either large, small or absent; the dentition usually includes three pairs of upper and one of lower incisors, and six or seven pairs of cheekteeth in each jaw; the stomach is cither simple or sacculated-, without a cardiac gland ; and there are four teats.
With the exception of the aberrant long-snouted phalanger, the members of the family Phalangeridae have the normal number of functional incisors, in addition to which there may be one or two rudimentary pairs in the lower jaw. The first in the upper jaw is strong, curved and cutting, the other two generally somewhat smaller; the single lower functional incisor large, more or less inclined forwards; canines - , upper small or moderate, conical I oro and sharp-pointed; lower absent or rudimentary; premolars variable; molars j, or , with four obtuse tubercles, sometimes forming crescents. Limbs subequal. Fore-feet with five distinct subequal toes with claws. Hind-feet short and broad, with five welldeveloped toes; the first large, nailless and opposable; the second and third slender and united by a common integument as far as the claws. Caecum present (except in Tarsipes), and usually large.
From Gould.
FIG. 12. The Long-snouted Phalanger (Tarsipes rostratus).
The lower jaw has no pocket on the outer side. All are animals of small or moderate size and arboreal habits, feeding on a vegetable or mixed diet, and inhabiting Australia, Papua and the Moluccan Islands.
As the first example of the group may be taken the elegant little long-snouted phalanger (Tarsipes rostratus, fig. 12), a west Australian creature of the size of a mouse, which may be regarded as representing by itself a sub-family (Tarsipediinae), characterized by the rudimentary teeth, the long and extensile tongue, and absence of a caecum. The head is elongated, with a slender muzzle and the 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 jaw of the same individuals. The formula in one specimen was i-^-^i c. _ , p. +m.-S _ 4 ; total 20. The lower jaw is slender, nearly straight, and without a coronoid process or inflected angle. Fore-feet with five well-developed toes, carrying small, flat, scale-like nails, not reaching the extremity of the digits. Hind-feet rather long and slender, with a well-developed opposable and nailless first toe; second and third digits united, with sharp, compressed curved claws; the fourth and fifth free, with small flat nails. Ears of moderate size and rounded. Tail longer than the body and head, scantily clothed with short hairs, prehensile. Vertebrae: C. 7, D. 13, L. 5, S. 3, Ca. 24.
As indicated in the accompanying illustration, the long-snouted phalanger is arboreal in habits, extracting honey and probably small insects from long-tubed flowers by means of its extensile tongue.
The remaining members of the family may be included in the sub family Phalangerinae, characterized by the normal nature of the dentition (which shows rudimentary lower canines) and tongue. C'uscuscs and phalangers form a numerous group, all the members of which are arboreal, and some of which are provided with lateral expansions of skin enabling them to glide from tree to tree like flying-squirrels. The typical members of the group are the cuscuses (Phalanger), ranging from the Moluccas and Celebes to New Guinea, in which the males are often different in colour from the females. The true phalangers, or opossums of the colonists, constitute the genus Trichosurus, while the ring-tailed species are known as Pseudochirus; the latter ranging to New Guinea. Dactylopsila is easily recognized by its attenuated fourth finger and parti-coloured fur; the flying species are classed as Petauroides, Petaurus, Gymnobelideus and Acrobates, the last no larger than a mouse ; while Dromicia, Distaechurus and Acrobates are allied types without parachutes (see PHALANGER).
An equally brief notice must suffice of the kangaroo tribe or Macropodidae, since these receive a special notice elsewhere. The dentition is i. 3-c. or * p..2.m..2;theincisorsbeingsharpandcutting, and those of the lower jaw frequently having a scissor-like action against one another. The broad molars are either bluntly tuberculated or transversely ridged; the outer side of the hind part of the lower jaw has a deep pocket; and the hind-limbs are generally very long, with the structure of the foot similar to that of the bandicoots. The family is connected with the Phalangeridae by means of the musk-kangaroo (Hypsiprymnodon moschatus) ; forming the sub-family Hypsiprymnodontinae. Then come the rat-kangaroos, or kangaroo-rats, constituting the sub-family Potoroinae; while the tree-kangaroos (Dendrolagus) , rock-wallabies (Petrogale), and wallabies and kangaroos (Macropus) form the Macropodinae (see KANGAROO).
Extinct Marsupials Reference has been made to the Australasian Pleistocene genera Phascolonus, Diprotodon, Nototherium and Thylacoleo, whose affinities are with the wombats and phalangers. The same deposits have also yielded remains of extinct types of kangaroo, some of gigantic size, constituting the genera Sthenurus, Procoptodon and Palorchestes. Numerous types more or less nearly allied to the phalangers, such as Burramys and Triclis have also been described, as well as a flying form, Paiaeopelaurus. It is also interesting to note that fossil remains indicate the former occurrence of thylacines and Tasmanian devils on the Australian mainland. Of more interest is the imperfectly known Wynyardia, from older Tertiary beds in Tasmania, which apparently presents points of affinity both to phalangers and dasyures. From the Oligocene deposits of France and southern England have been obtained numerous remains of opossums referable to the American family Didelphyidae. These ancient opossums have been separated generically from Didelphys (in its widest sense) on account of certain differences in the relative sizes of the lower premolars, but as nearly the whole of the species have been formed on lower jaws, of which some hundreds have been found, it is impossible to judge how far these differences are correlated with other dental or osteological characters. In the opinion of Dr H. Filhol, the fossils themselves represent two genera, Peratherium, containing the greater part of the species, about twenty in number, and Amphiperatherium, with three species only. All are comparatively small animals, few of them exceeding the size of a rat.
Besides these interesting European fossils, a certain number of didelphian bones have been found in the caves of Brazil, but these are either closely allied to or identical with the species now living in the same region.
The occurrence in the Santa Cruz beds of Patagonia of fossil marsupials allied to the living Caenolestes has been mentioned above. The alleged occurrence in the same beds of marsupials allied to the thylacine is based on remains now more generally regarded as referable to the creodont carnivores (see CREODONTA).
Mesozoic Mammals. Under the heading of MULTITUBERCULATA will be found a brief account of certain extinct mammals from the Mesozoic formations of Europe and North America which have been regarded as more or less nearly related to the monotremes. The same deposits have yielded remains of small mammals whose dentition approximates more nearly to that of either polyprotodont marsupials or insectiyores; and! these may be conveniently noticed here without prejudice to their true affinities. Before proceeding further it may be mentioned that the remains of many of these mammals are very scarce, even in formations apparently in every way suitable to the preservation of such fossils, and it hence seems probable that these creatures are stragglers from a country where primitive small mammals were abundant. Not improbably this country was either " Gondwana-land," connecting Mesozoic India with Africa, or perhaps Africa itself. At any rate, there seems little doubt that it was the region where creodonts and other primitive mammals were first differentiated from their reptilian ancestors.
Of the Old World forms, the family Triconodontidae is typified by the genus Triconedon, from the English Purbeck, in which the cheek-teeth carry three cutting cusps arranged longitudinally. There seems to have been a replacement of some of these teeth; and it has been suggested that this was of the marsupial type. To the same family are referred Phascolotherium (fig. 14), of the Lower Jurassic Stonesfield slate of England, and Spalacotherium (fig. 15), of the Dorsetshire Purbeck; the latter having the three cusps of the cheek-teeth rotated so as to assume a tritubercular type. Other From Owen.
FIG. 13. Lower Jawof Triconodon mordax (nat. size).
From Owen.
FIG. 14. Lower Jaw and Teeth of Phascolotherium bucklandi (mt. size in outline).
genera are Menacodon and Priacodon, the former American, and the latter common to Europe and North America. By one authority Amphilestes (fig. 16), of the Stonesfield Slate, is included in the same group, while by a second it is regarded as representing a family by itself. Amphitherium, of the From Owen. Stonesfield Slate, typifies the FIG. 15. Spalacotherium tricuspidens family Amphitheriidae, which (twice nat. size), Purbeck beds. includes the American Dryo- lestes, and in which some would class the European Purbeck genus Amblotherium, although Professor H. F. Osborn has made the last the type of a d'stinct family. Yet another family, according to the palaeontologist last named, is typified by the genus Stylacodon, of the English Purbeck. To mention the other forms which have received names will be unnecessary on this occasion.
It will be observed from the figures of the lower jaws, which are in most cases the only parts known, that in many instances the number of cheek-teeth exceeds that found in modern marsupials except Myrmecobius. The latter has indeed been regarded as the direct descendant of these Mesozoic forms; but as already stated, in the opinion of Mr B. A. Bensley, this is incorrect. It may be added that the division of these teeth into premolars and molars in figs. 14 and 16 is based upon the view of Sir R. Owen, and is not altogether trustworthy, while the restoration of some of the missing Prorn Owen.
FIG. 16. Lower Jaw and Teeth of Amphilestes broderipi (twice nat. size).
teeth is more or less conjectural. As regards the affinities of the creatures to which these jaws belonged. Professor Osborn has referred the Triconodontidae and Amphitheriidae, together with the Curtodontidae (as represented by the English Purbeck Curtodon), to a primitive group of marsupials, while he has assigned the Amblotheriidae and Stylacodontidae to an ancestral assemblage of Insectivora. On the other hand, in the opinion of Professor H. Winge, a large number of these creatures are primitive monotremes. Besides the above, in the Trias of North America we have Dromotherium and Microconodon, extremely primitive forms, representing the family Marsupial Mole (Notoryctes typhlops).
Dromotheriidae, and apparently showing decided traces of reptilian affinity. It may be added that a few traces of mammals have been obtained from the English Wealden, among which an incisor tooth foreshadows the rodent type.
AUTHORITIES. The above article is partly based on that by Sir W. H. Flower in the gth edition of this work. See also O. Thomas, Catalogue of Monotremata and Marsupialia in the British Museum (1888); "On Caenolestes, a Survivor of the Epanorthidae," Proc. Zool. Soc. London (1895); J. D. Ogilby, Catalogue of Australian Mammals (Sydney, 1805); B. A. Bensley, " A Theory of the Origin and Evolution of the Australian Marsupialia," American Naturalist (1901) ; " On the Evolution of the Australian Marsupialia, etc.," Trans. Linn. 5oc.(vol. ix., 1903) ; L.Dollo," Arboreal Ancestry of Marsupials," Miscell. Biologiques (Paris, 1899); B. Spencer, " Mammalia of the Horn Expedition " (1896) ; "Wynyardia, a Fossil Marsupial from Tasmania," Proc. Zool. Soc. London (1900) ; J. P. Hill, " Contributions to the Morphology of the Female Urino-genital Organs in Marsupialia," Proc. Linn. Soc. N. S. Wales, vols. xxiv. and xxv. ; " Contributions to the Embryology of the Marsupialia," Quart. Journ. Micr. Science, vol. xliii. ; E. C. Stirling, " On Notoryctes typhlops," Proc. Zool. Soc. London (1891) ; " Fossil Remains of Lake Cadibona," Part I. Diprotodon, Mem. R. Soc. S. Australia (vol. i., 1889) ; R. Broom, " On the Affinities of Thylacoleo," Proc. Linn. Soc. N. S. Wales (1898) ; H. F. Osborn, " Mesozoic Mammalia," Journ. Acad. Nat. Sci. Philadelphia (vol. ix., 1888); E. S. Goodrich, " On the Fossil Mammalia from the Stonesfield Slate," Quart. Journ. Micr. Science (vol. xxxv., 1894).
(R. L.*)
Note - this article incorporates content from Encyclopaedia Britannica, Eleventh Edition, (1910-1911)
