Читать книгу Mammalia - Frank E. Beddard - Страница 17

The Marsupials may be thus defined:—Terrestrial, arboreal, or burrowing (rarely aquatic) mammals, with furry integuments; palate generally somewhat imperfectly ossified; jugal bone reaching as far as the glenoid cavity; angle of lower jaw nearly always inflected. The clavicle is developed. Arising from the pubes are well developed and ossified epipubic bones. Fourth toe usually the most pronounced. Teeth often exceed the typical Eutherian number of forty-four; molars generally four on each side of each jaw. As a rule but one tooth of the milk set is functional, which is (according to many) the fourth premolar. Teats lying within a pouch, in which the young are placed. Young born in an imperfect condition, and showing certain larval characters. There is a shallow cloaca. The testes are extra-abdominal, but hang in front of the penis. In the brain the cerebellum is completely exposed; the hemispheres are furrowed, but the corpus callosum is rudimentary. An allantoic placenta is rarely present.

Structurally the Marsupials are somewhat intermediate between the Prototheria and the more typical Eutheria, with a greater resemblance to the latter.

Fig. 59.—Rock Wallaby (Petrogale xanthopus), with young in pouch. × 1⁄7. (After Vogt and Specht.)

The name Marsupial indicates what is perhaps the most salient character of this order. The pouch in which the young are carried is almost universally present. It is less developed on the whole in the Polyprotodont forms, such as the Thylacine, Dasyures, etc., but is found in so many of them that the two divisions of the Marsupials, the Diprotodonts and the Polyprotodonts, cannot be raised to distinct orders on this and other grounds. The marsupial pouch of the Marsupials must not, as has been already pointed out, be confounded with the pouch of the Monotreme mammals. Distinct teats are found in the marsupium of the Marsupials, while there are none in the mammary pouch of the Monotreme, the pouch itself indeed representing an undifferentiated teat, of which the walls have not closed up. The pouch opens forward in the Kangaroos, and backwards in the Phalangers and in the Polyprotodonts. Its walls are supported by a pair of bones diverging from each other in a -shaped manner; these are cartilaginous and vestigial in the Thylacine. They are the precise equivalents of similar bones in the Monotremata. It has been held, but apparently erroneously, that these bones are mere ossifications in the tendons of the external oblique muscle of the abdomen, or of the pyramidalis of the same region; and vestiges have been asserted to exist in the Dog. Such bonelets are undoubtedly present in the Dog; but it seems clear from their development in Marsupials, as structures actually continuous with the median unossified portion of the symphysis pubis, that the "marsupial bones" belong to that part of the skeleton, and that they correspond with the epipubis of certain amphibians and reptiles. The pouch, it may be remarked, exists in a rudimentary form in the males of many Marsupials.

Fig. 60.—Ventral surface of innominate bone of Kangaroo (Macropus major). × ⅓. a, Acetabulum; ab, acetabular border of ilium; is, iliac surface; m, "marsupial" bone; pb, pubic border; pt, pectineal tubercle; s, symphysis; si, supra-iliac border; ss, sacral surface; thf, thyroid foramen; ti, tuberosity of ischium. (From Flower's Osteology.)

Fig. 61.—Mammary foetus of Kangaroo attached to the teat. (Nat. size.) (From Parker and Haswell's Zoology.)

The most salient feature in the life-history of the Marsupials is the imperfect condition in which the young are born. The egg is no longer laid, as in the Monotremes; but curiously enough the ovum, which has the small size of that of the Eutheria, divides incompletely at the first division (as Mr. Caldwell has shown), and this developmental feature may perhaps be looked upon as a reminiscence of a former large-yolked condition. The young when born are small and nude; the newly born young of a large Kangaroo is perhaps as large as the little finger. The young are transferred by the lips of the mother to the pouch, where they are placed upon a teat. It is an interesting fact that they are not merely imperfect foetuses, but that they are actual larvae. They possess in fact at any rate one larval organ in the shape of a special sucking mouth. This sucking mouth is an extra-uterine production, and is of course an adaptation to the particular needs of the young, just as are other larval organs, such as the chin-suckers of the tadpole, or the regular ciliated bands of the larvae of various marine invertebrate organisms.

There are a number of other features which distinguish the Marsupials from other mammals.

The cloaca of the Marsupials is somewhat reduced, but is still recognisable. Its margins in Tarsipes are even raised into a wall, which projects from the body.

The tooth series of the Marsupials was once held to consist of one dentition only, with the exception of the last premolar, which has a forerunner. The interpretation of the teeth of Marsupials are various. Perhaps most authorities regard the teeth as being of the milk dentition, with the exception of course of the single tooth that has an obvious forerunner. But there are some who hold that the teeth are of the permanent dentition. In any case it is proved that a set of rudimentary teeth are developed before those which persist. Those who believe in the persisting milk dentition describe these as prelacteal. Another matter of importance about the teeth of this order of mammals is that their numbers are sometimes in excess of the typical Eutherian 44. This, however, holds good of the Polyprotodonts only.

It was for a long time held that the Marsupials differed from all other mammals in having no allantoic placenta. But quite recently this supposed difference has been proved to be not universal by the discovery in Perameles of a true allantoic placenta. The Marsupials have been sometimes called the Didelphia. This is on account of the fact that the uterus and the vagina are double. Very frequently the two uteri fuse above, and from the point of junction an unpaired descending passage is formed (see Fig. 48 on p. 74).

A character of the brain of Marsupials has been the subject of some controversy. Sir Richard Owen stated many years ago that they were to be distinguished from the higher mammals by the absence of the corpus callosum. Later still it was urged that a true corpus callosum, though a small one, was present; while, finally, Professor Symington^[68] seems to have shown that 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.