Page:EB1911 - Volume 17.djvu/542

Rh the horse and the dog may be cited as instances where the single caecum is of large size, this being especially the case of the former, where it is of enormous dimensions; in human beings, on the other hand, the caecum is rudimentary, and best known in connexion with “appendicitis.” The existence of paired caeca was previously known in a few armadillos and anteaters, but Dr Mitchell has shown that they are common in these groups, while he has also recorded their occurrence in the hyrax and the manati. With the aid of these instances of paired caeca, coupled with the frequent existence of a rudiment of its missing fellow when only one is functional, the author has been enabled to demonstrate conclusively that these double organs in birds correspond in relations with their normally single representative in mammals.

In mammals both caecum and colon are often sacculated, a disposition caused by the arrangement of the longitudinal bands of muscular tissue in their walls; but the small intestine is always smooth and simple-walled externally, though its lining membrane often exhibits contrivances for increasing the absorbing surface without adding to the general bulk of the organ, such as the numerous small tags, or “villi,” by which it is everywhere beset, and the more obvious transverse, longitudinal, or reticulating folds projecting into the interior, met with in many animals, of which the “valvulae conniventes” of man form well-known examples. Besides the crypts of Lieberkühn found throughout the intestinal canal, and the glands of Brunner confined to the duodenum, there are other structures in the mucous membrane, about the nature of which there is still much uncertainty, called “solitary” and “agminated” glands, the latter more commonly known by the name of “Peyer’s patches.” Of the liver little need be said, except that in all living mammals it has been divided into a number of distinct lobes, which have received separate names. It has, indeed, been suggested that in the earlier mammals the liver was a simple undivided organ. This, however, is denied by G. Ruge (vol. xxix. of Gegenbaur’s Morphologisches Jahrbuch).

Origin of Mammals.—That mammals have become differentiated from a lower type of vertebrates at least as early as the commencement of the Jurassic period is abundantly testified by the occurrence of the remains of small species in strata of that epoch, some of which are mentioned in the articles and (q.v.). Possibly mammalian remains also occur in the antecedent Triassic epoch, some palaeontologists regarding the South African Tritylodon as a mammal, while others consider that it was probably a reptile. Whatever may be the true state of the case with regard to that animal probably also holds good in the case of the approximately contemporaneous European Microlestes. Of the European Jurassic (or Oolitic) mammals our knowledge is unfortunately very imperfect; and from the scarcity of their remains it is quite probable that they are merely stragglers from the region (possibly Africa) where the class was first differentiated. It is not till the early Eocene that mammals become a dominant type in the northern hemisphere.

It is now practically certain that mammals are descended from reptiles. Dr H. Gadow, in a paper on the origin of mammals contributed to the Zeitschrift für Morphologie, sums up as follows: “Mammals are descendants of reptiles as surely as they [the latter] have been evolved from Amphibia. This does not mean that any of the living groups of reptiles can claim their honour of ancestry, but it means that the mammals have branched where the principal reptilian groups meet, and that is a long way back. The Theromorpha, especially small Theriodontia, alone show us what these creatures were like.” It may be explained that the Theromorpha, or Anomodontia, are those extinct reptiles so common in the early Secondary (Triassic) deposits of South Africa, some of which present a remarkable resemblance in their dentition and skeleton to mammals, while others come equally near amphibians. A difficulty naturally arises with regard to the fact that in reptiles the occipital condyle by which the skull articulates with the vertebral column is single, although composed of three elements, whereas in amphibians and mammals the articulation is formed by a pair of condyles. Nevertheless, according to Professor H. F. Osborn, the tripartite reptilian condyle, by the loss of its median element, has given rise to the paired mammalian condyles; so that this difficulty disappears. The fate of the reptilian quadrate bone (which is reduced to very small dimensions in the Anomodontia) has been referred to in an earlier section of the present article, where some mention has also been made of the disappearance in mammals of the hinder elements of the reptilian lower jaw, so as to leave the single bone (dentary) of each half of this part of the skeleton in mammals.

Most of the earliest known mammals appear to be related to the Marsupialia and Insectivora. Others however (inclusive of Tritylodon and Microlestes, if they be really mammals), seem nearer to the Monotremata; and the question has yet to be decided whether placentals and marsupials on the one hand, and monotremes on the other are not independently derived from reptilian ancestors.

With regard to the evolution of marsupials and placentals, it has been pointed out that the majority of modern marsupials exhibit in the structure of their feet traces of the former opposability of the thumb and great toe to the other digits; and it has accordingly been argued that all marsupials are descended from arboreal ancestors. This doctrine is now receiving widespread acceptation among anatomical naturalists; and in the American Naturalist for 1904, Dr W. D. Matthew, an American palaeontologist, considers himself provisionally justified in so extending it as to include all mammals. That is to say, he believes that, with the exception of the duckbill and the echidna, the mammalian class as a whole can lay claim to descent from small arboreal forms. This view is, of course, almost entirely based upon palaeontological considerations; and these, in the author’s opinion, admit of the conclusion that all modern placental and marsupial mammals are descended from a common ancestral stock, of which the members were small in bodily size. These ancestral mammals, in addition to their small size, were characterized by the presence of five toes to each foot, of which the first was more or less completely opposable to the other four. The evidence in favour of this primitive opposability is considerable. In all the groups which are at present arboreal, the palaeontological evidence goes to show that their ancestors were likewise so; while since, in the case of modern terrestrial forms, the structure of the wrist and ankle joints tends to approximate to the arboreal type, as we recede in time, the available evidence, so far as it goes, is in favour of Dr Matthew’s contention.

The same author also discusses the proposition from another standpoint, namely, the condition of the earth’s surface in Cretaceous times. His theory is that in the early Cretaceous epoch the animals of the world were mostly aerial, amphibious, aquatic or arboreal; the flora of the land being undeveloped as compared with its present state. On the other hand, towards the close of the Cretaceous epoch (when the Chalk was in course of deposition), the spread of a great upland flora vastly extended the territory available for mammalian life. Accordingly, it was at this epoch that the small ancestral insectivorous mammals first forsook their arboreal habitat to try a life on the open plains, where their descendants developed on the one hand into the carnivorous and other groups, in which the toes are armed with nails or claws, and on the other into the hoofed group, inclusive of such monsters as the elephant and the giraffe. The hypothesis is not free from certain difficulties, one of which will be noticed later.

Classification.—Existing mammals may be primarily divided into three main groups, or subclasses, of which the second and third are much more closely related to one another than is either of them to the first. These three classes are the Monotremata (or Prototheria), the Marsupialia (Didelphia, or Metatheria), and the Placentalia (Monodelphia, or Eutheria); the distinctive characters of each being given in separate articles (see, and .)

The existing monotremes and marsupials are each represented only by a single order; but the placentals are divided into the following ordinal and subordinal groups, those which are extinct being marked with an asterisk (*):—