the original statement of Owen was correct, at least in part. It is at most feebly developed (see Fig. 58, p. [118]).
As to skeletal characters, the Marsupial skull has on the whole a tendency towards a permanent separation of bones usually firmly ankylosed. Thus the orbitosphenoids remain distinct from the presphenoid. The palate is largely fenestrated, a return as it were—says Professor Parker—to the Schizognathous palate of the bird. The mandible is inflected; this familiar character of the Marsupials goes back to the earliest representatives of the order in Mesozoic times (see p. [96]); but it is not absolutely universal, being absent from the much weakened skull of Tarsipes. On the other hand, the inflection is nearly as great in certain Insectivores, in Otocyon, etc. The malar always extends back to form part of the glenoid cavity. The shoulder girdle has lost the large coracoid of Monotremes; this bone has the vestigial character that it possesses in other Eutheria. The clavicle is present except in the Peramelidae. A third trochanter upon the femur seems to be never present.
Fig. 62.—Skull of Rock Wallaby (Petrogale penicillata). (Ventral view.) ali, Alisphenoid; bas.oc, basi-occipital; bas.sph, basi-sphenoid; ex.oc, ex-occipital; ju, jugal; max, maxilla; pal, palatine; par.oc, paroccipital; p.max, premaxilla; pr.sph, presphenoid; pt, pterygoid; sq, squamosal; ty, tympanic. (From Parker and Haswell's Zoology.)
The Marsupials cannot be regarded as an intermediate stage in the origin of the Eutheria for a number of reasons. In the first place, the nature of their teeth shows them to be degenerate animals; one set, whether we regard it as the milk or permanent dentition, has become vestigial. The recent discovery of a true allantoic placenta in Perameles removes one reason for regarding
the Marsupials as primitive creatures. It implies on the whole that the Marsupials have sprung from a stock with an allantoic placenta. The alternative is to assume the independent development of an allantoic placenta in both groups of the Mammalia; unless indeed the genus Perameles is to be held to be the most primitive race of Marsupials living, a hypothesis which does not appear on the face of it likely. So long as it was believed that the mammary pouch of the Monotremes was the equivalent of the marsupium of the Marsupials, the persistence of this structure seemed to be a bond of union between the groups. But it is now known that the marsupium is a special organ confined to the Marsupials, an argument which is rather in favour of their being a lateral development of the mammalian stem. It is to be remarked also that the marsupium is feeblest in the Polyprotodonts, which may perhaps be looked upon as the most primitive of the Marsupials, owing to their more numerous teeth and other points to be referred to immediately.
Not only are the Marsupials interesting from the point of view of their structure; their present and past distribution is of equal interest. During the Mesozoic epoch they occurred in Europe and North America; but not, so far as negative evidence means anything, in Australia, which is now their headquarters. In Europe Marsupials lingered on into the Tertiary period, when they finally became extinct. In America, of course, the group has persisted to the present day. Now it is important to notice that the two main subdivisions of the Marsupials, the Polyprotodontia and the Diprotodontia, exist to-day in both Australia and South America. These two divisions, it should be explained, differ principally in that one has numerous, the other rarely more than two,[[69]] incisors in the lower jaw. It is perhaps the more widely distributed opinion that the Polyprotodontia are the more archaic group; this opinion rests upon one or two facts in addition to the absence of specialisation in the incisor teeth. Among the Polyprotodontia the total number of teeth is greater—a clearly primitive character; secondly, the general form of the body of these animals, with four subequal limbs and carnivorous or omnivorous diet, contrasts with the purely vegetarian and much specialised Kangaroos at any rate. Finally—and sufficient stress
has perhaps not been laid upon this matter—the brain among the Polyprotodonts is less convoluted than among the genera of the other division. This statement is of course made with due regard to parallelism in size (see p. [77]). It is well known that the complexity of a brain bears a distinct relation to the size of its possessor within the group. Now the most ancient Marsupials are decidedly more Polyprotodont-like. No European form from the earlier periods is distinctly to be referred to the Diprotodonts. But both divisions now exist in America and Australia.
We must assume, therefore, one of three hypotheses. Either the differentiation into the two great divisions occurred in Jurassic or Cretaceous times before the migration of the order southwards; or the Diprotodont type is only a type, and not a natural group, i.e. it has been separately evolved in America and Australia; or, finally, there was formerly a land-connexion in the Antarctic hemisphere, along which the Diprotodonts of Australia wandered into South America. The middle hypothesis has this to commend it, that syndactylism occurs in both divisions, and that in some Diprotodonts the pouch opens backwards as it does in the Polyprotodonts. So great are the resemblances that but little difference is really left—of great importance that is to say. Hence it is not difficult to imagine the reduction of the incisors having taken place twice. In favour of the first hypothesis there are no positive facts. Finally, in favour of the last, which is so strongly supported by the facts of distribution derived from the study of other groups of animals,[[70]] there is at least this striking fact or rather series of facts: that some of the South American fossil Polyprotodonts have a "strictly Dasyurine relationship."[[71]] If there has not been a direct migration, then the Dasyurine type has been twice evolved, an improbability that few will attempt to explain away. In any case we shall adopt here the usual division of the Marsupials into Diprotodontia and Polyprotodontia.