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 stock. 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.
1. Polyprotodonts.—The Polyprotodonts 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 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.
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Fig. 2.—Front View of Skull of the Tasmanian Devil (Sarcophilus ursinus) to exhibit polyprotodont type of dentition. |
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 sub-equal 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, &c. 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. 43, c. 11, p. 44, m. 33 = 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 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-vark, and may be directly inherited from reptilian ancestors (see Thylacine).
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Fig. 3.—The Tasmanian Wolf, or Thylacine (Thylacinus cynocephalus). |
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. 43, c. 11, p. 33, m. 33: 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. 43, c. 11, p. 44, m. 33: 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 almost 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 Phascologale, 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.
Sminthopsis 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)
