Huxley regarded the peculiarities in the reproductive organs
of the Marsupials as "singularly specialised characters," in no way intermediate in character. This view applies also to the pouch, which, as already stated, distinguishes the adults of that group. But the impossibility of using this last character as one of any importance has been shown by the discovery of rudiments of it in embryos of undoubtedly Eutherian mammals (see p. [18]).
Fig. 58.—Sagittal section of brain of Rock Wallaby (Petrogale penicillata). ant.com, Anterior commissure; cbl, cerebellum; c.mam, corpus mammillare; c.qu, corpora quadrigemina; crur, crura cerebri; epi, epiphysis, with the posterior commissure immediately behind it; f.mon, position of foramen of Monro; hip.com, hippocampal commissure, consisting here of two layers continuous behind at the spleneium, somewhat divergent in front where the septum lucidum extends between them; hypo, hypophysis; med, medulla oblongata; mid.com, middle commissure; olf, olfactory lobe; opt, optic chiasma; vent. 3, third ventricle. (From Parker and Haswell's Zoology.)
Less stress is laid now upon the existence of four molars in the Marsupials as dividing them from the higher mammals than was formerly the case. The total dentition of the group is on the whole composed of more numerous individual teeth than in the typical Eutheria; but we have exceptions like the Whales, the Armadillo Priodontes, and the Manatee; or better, because free from the suspicion of secondary multiplication, Otocyon and occasionally (according to Mr. Thomas) Centetes. In the last two there are at least sometimes four molars.
On the other hand, a few archaic characters of some importance crop up here and there among the Marsupials, which are sometimes held to point to a primitive ancestry. It has been remarked that in Marsupials it is the fourth toe which is dominant in size, whereas in Ungulates it is the third. An attempt has been made to explain this on the view (reasonable enough in itself) of a tree-living ancestry for the group. A greater development of the fourth toe is, however, by no means a necessary character of arboreal creatures; the Primates themselves are an exception. Nor is this prevalence universal among the Marsupials;
in Myrmecobius (alone) is the third toe the longest; and no great difference can be detected between the third and fourth toes in the case of the genera Phascologale, Didelphys, and some others. Professor Leche compares the predominance of the fourth toe with the hyperphalangeal condition in the fourth toe of the embryo Crocodile, and considers it an archaic feature, not surpassed by the ancient characteristics of the Monotremata. Again it has been pointed out that in Phascologale and Perameles, the epistropheus (axis vertebra) has a separate rib as in Ornithorhynchus. In the third place, the likeness of the teeth of Myrmecobius to those of Ornithorhynchus is an argument in the same direction, which is furthermore supported by the great age (Mesozoic) of the Metatherian group, if we are right in regarding those extinct creatures as Marsupials.
We may now mention certain facts which are not so generally used. The partly primitive structure of the right auriculo-ventricular valve in the Monotremata has no counterpart in any Marsupial which has been dissected; but there are traces in the latter of the characteristic "ventral mesentery" of Ornithorhynchus and Echidna.[[65]] Mr. Caldwell's interesting observation upon the segmenting egg of the Marsupial, the incompleteness of the first segmentation furrow (reminding us of the meroblastic ovum of the Monotreme), may possibly not turn out to be so exclusively Marsupial a feature as has been thought.
The balance of evidence thus points to the nearer relationship of the Marsupials to the Eutherian mammals; and their great specialisation combined with certain evidences of degeneration (disappearance in part of the milk dentition), and their age, point to the fact that they are, at any rate, the descendants of an early form of Eutherian. But they must have separated from the Eutherian stock after it had acquired a definite diphyodonty and the allantoic placenta, the two principal features of the Eutherian as opposed to the Prototherian mammals.
Nevertheless it seems probable that the Marsupial tribe is derived from some of the earliest Eutherians. And on this view may be explained the retention of Prototherian characters.