Cambridge Natural History Mammalia/Chapter VI
INTRODUCTION TO THE SUB-CLASS EUTHERIA
Definition.—Mammalia with teats. Mammary glands of sebaceous type. Heart with entirely membranous and complete right auriculo-ventricular valve. Brain generally with a corpus callosum. Coracoid much reduced and not reaching sternum. No interclavicle. Vertebrae with epiphyses. Ribs double-headed. Viviparous, with a small ovum.
In this group are included not only the Eutheria in the sense of Huxley, but also his Metatheria. Though the Metatheria, or Marsupials as we shall term them, undoubtedly form a most distinct order of mammals, perhaps even a trifle more distinct than most others, their differences from the remaining tribes are not by any means so great as those which separate Ornithorhynchus and Echidna from all other mammals. In his well-known memoir upon the arrangement of the Mammalia, Professor Huxley enumerated eleven characters as distinguishing the Metatheria either from the Prototheria or from the Eutheria. Of these only three were characters in which they approach the lower mammals. According to his showing, therefore, the preponderance of marsupial features are Eutherian. The three characters of Prototherian type are (1) the presence of epipubes; (2) the small corpus callosum; (3) the absence of an allantoic placenta.
The last of these can be dismissed, in consequence of the recent discovery of an allantoic placenta in Perameles. The first character is apparently a valid distinction between the Marsupials and their mammalian relatives higher in the series; but it is not a character that should have been made use of by Huxley, since he believed in the existence of a corresponding element in the Dog. As to the corpus callosum (Fig. 50, p. 77) being small, that seems to be not more than a slight difference of degree. A number of other characters of secondary importance were added by Huxley to the weight of evidence which led him to form a group Metatheria for the Marsupials. Some of these, however, are now known to be not evidence in that direction. For instance he observed that no Marsupial had more than a single successional tooth. It seems at the present moment to be fairly clear that Marsupials have a milk dentition like other Eutherians, but that only one of these teeth, the fourth premolar, comes to functional maturity. That it is really one of a complete milk series is evidenced by the fact that this tooth is differentiated contemporaneously with another series formerly held to belong to the so-called prelacteal dentition. There still remains, of course, the actual fact that the milk dentition is not for the most part functional, but its significance breaks down with these fresh discoveries. Of this Professor Osborn has remarked: "The discovery of the complete double series seems to have removed the last straw from the theory of the marsupial ancestry of the Placentals." But Huxley did not lay much stress upon this matter of the teeth, since he observed that similar suppressions of the milk dentition were to be found in many other mammals admittedly Eutherian.
Fig. 57.—Brain of Echidna aculeata; sagittal section. ant.com, Anterior commissure; cbl, cerebellum; c.mam, corpus mammillare; col.forn, column of the fornix; c.qu, corpora quadrigemina; gang.hab, ganglion habenulare; hip.com, hippocampal commissure; med, medulla oblongata; mid.com, middle commissure; olf, olfactory lobe; opt, optic chiasma; tub.olf, tuberculum olfactorium; vent. 3, third ventricle. (From Parker and Haswell's Zoology.)
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. 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.
The remaining Eutheria are obviously all to be referred to one great division with the possible exception of the Whales, whose affinities form one of the principal difficulties to the student of this group. A short résumé of what is at present thought of the systematic position of this anomalous order is appropriate here. Albrecht went so far as to regard the Cetacea as the nearest group of animals to the hypothetical Promammalia. But discounting his arguments by the removal of such of them as relate to structure plainly altered by the singular mode of life of these creatures, there is really a great deal to be said in favour of his view.
The chief facts which argue a primitive position among mammals for the Cetacea are perhaps: (1) the slight union of the rami of the lower jaw; (2) the occasionally rather marked traces of the double constitution of the sternum; (3) the long and simple lungs; (4) the retention of the testes within the body-cavity; (5) the occasional presence (in Balaenoptera) of a separate supra-angular bone. These points, however, are but few, and are not of such great weight as those which ought to be present to establish a claim to separate treatment for the Cetacea as opposed to the Eutheria. If this group of mammals can be tacked on anywhere, it appears to us that the nearest relatives are not, as is sometimes put forward, the Ungulata or the Carnivora, but the Edentata. There are quite a number of rather striking features in which a likeness is shown between these apparently diverse orders of mammals. The chief ones are these: (1) the existence of traces of a hard exoskeleton, of which vestiges remain in the Porpoise; (2) the double articulation of the rib of the Balaenopterids to the sternum, with which compare the conditions obtaining in the Great Anteater; (3) the concrescence of some of the cervical vertebrae; (4) the share which the pterygoids may take in the formation of the hard palate; (5) the fact that in the Porpoise, at any rate, as in many Edentates, the vena cava, instead of increasing in size as it approaches the liver, diminishes.
Another group which is perfectly isolated is that of the Sirenia. The alliance advocated by some with the Cetacea, and quite recently renewed by Professor Haeckel, is contradicted by so many important features that it seems necessary to abandon it. The recent discovery of a fossil Sirenian jaw by Dr. Lydekker with teeth highly suggestive of those of Artiodactyla, may prove a clue. A third group which is so isolated as to have been placed in a primary division, proposed to be called Paratheria, is that of the Edentates. Probably the group so called should really be divided into the Edentata and the Effodientia, the latter containing the Old World forms. Whether or not it be ultimately shown that the Ganodonta are ancestral Edentates (sensu strictiori), the connexion of the group with others is not at present plain. The same is the case with the extensive order of Rodents. It is true that the extinct order of the Tillodontia shows certain Rodent-like characters on the one hand, and likenesses to Ungulates on the other. Certain likenesses shown by such apparently diverse animals as the Rabbit and the Elephant used to be insisted upon by Professor Huxley. For the present, however, the Rodents must remain as an isolated group with only very dubious affinities to others. The remaining groups of existing mammals are easier to connect. At first the differences between a Cat and a Horse seem to be quite as wide as those which separate any two of the higher Eutherian orders. But it seems to become clearer and clearer, as palaeontological investigation proceeds, that the bulk of the Ungulate and the Carnivorous, Insectivorous, and perhaps Lemuroid stocks converge into the early Eocene Creodonta. From the Lemuroid branch the higher Primates can be derived. The only "Ungulates" which cannot be fitted in with some reasonable probability is the group of the Proboscidea. But of the early forms of this division we have at present no knowledge.
- 62 ^ Proc. Zool. Soc. 1880, p. 649.
- 63 ^ Moreover, the "corpus callosum and the anterior commissure ... in ... Erinaceus and Dasypus are almost Monotreme-like."
- 64 ^ See Wilson and Hill, Quart. J. Micr. Sci. xxxix. 1899, p. 427.
- 65 ^ In Dendrolagus at any rate. See Proc. Zool. Soc. 1895, p. 132.
- 66 ^ Anat. Anz. i. 1886, p. 338; and see Weber, ibid. ii. 1887, p. 42.