The formation of the fœtal membrane, or amnion, which distinguishes reptiles, birds, and mammals from all other Vertebrata, is evidently a very important process in their ontogeny, and in the phylogeny which corresponds with it. It coincides with a series of other processes, which essentially determine the higher development of Amnionate animals. The first of these important processes is the total loss of gills, for which reason the Amniota, under the name of Gill-less animals (Ebranchiata), were formerly opposed to all other Vertebrate animals which breathed through gills (Branchiata). In all the Vertebrate already discussed, we found that they either always breathed through gills, or at least did so in early life, as in the case of Frogs and Salamanders. On the other hand, we never meet with a Reptile, Bird, or Mammal which at any period of its existence breathes through gills, and the gill-arches and openings which do exist in the embryos, are, during the course of the ontogeny, changed into entirely different structures, viz., into parts of the jaw-apparatus and the organ of hearing. (Compare above, vol. i. p. 307.) All Amnionate animals have a so-called cochlea in the organ of hearing, and a “round window” corresponding with it. These parts are wanting in the Amnion-less animals; moreover, their skull lies in a straight line with the axis of the vertebral column. In Amniotic animals the base of the skull appears bent in on the abdominal side, so that the head sinks upon the breast. (Plate III. Fig. C, D, G, H.) The organs of tears at the side of the eye also first develop in the Amniota.

The question now is, When did this important advance take place in the course of the organic history of the earth? When did the common ancestor of all Amniota develop out of a branch of the Non-amniota, to wit, out of the branch of the Amphibia?

To this question, the fossil remains of Vertebrata do not give us a very definite, but still they do give an approximate, answer. For with the exception of two lizard-like animals found in the Permian system (the Proterosaurus and Rhopalodon), all the fossil remains of Amniota, as yet known, belong to the secondary, tertiary, and quaternary epochs. With regard to the two Vertebrata just named, it is still doubtful whether they are genuine reptiles, or perhaps Amphibia of the salamander kind. Their skeleton alone is known to us, and even this not perfectly. Now as we know nothing of the characteristic features of their soft parts, it is quite possible that the Proterosaurus and Rhopalodon were non-amnionate animals more closely allied to Amphibia than to Reptiles; possibly they belonged to the transition form between the two classes. But, on the other hand, as undoubted fossil remains of Amniota have been found as early as the Trias, it is probable that the main class of Amniota first developed in the Trias, that is, in the beginning of the Mesolithic epoch. As we have already seen, this very period is evidently one of the most important turning points in the organic history of the earth. The palæolithic fern forests were then replaced by the pine forests of the Trias period; important transformations then took place in many of the classes of Invertebrata. Articulated marine lilies (Colocrina) developed out of the plated ones (Phatnocrina.) The Autechinidæ, or sea-urchins with only twenty rows of plates, took the place of the palæolithic Palechinidæ, the sea-urchins with more than twenty rows of plates. The Cystideæ, Blastoideæ, Trilobita, and other characteristic groups of Invertebrata of the primary period became extinct. It is no wonder that transforming conditions of adaptation powerfully influenced the Vertebrate tribes also in the beginning of the Trias period, and caused the origin of Amniotic animals.

If, however, the two Lizard and Salamander-like animals of the Permian system, the Proterosaurus and Rhopalodon, are considered genuine Reptiles, and consequently the most ancient Amniota, then the origin of this main class must necessarily have taken place in the preceding period, towards the end of the primary, namely, in the Permian period. However, all other remains of Reptiles, which were formerly believed to have been found in the Permian and the Coal system, or even in the Devonian system, have been proved to be either not remains of Reptiles at all, or to belong to a more recent date (for the most part to the Trias). (Compare Plate XIV.)

The common hypothetical primary form of all Amniotic animals, which we may call Protamnion, and which was possibly nearly related to the Proterosaurus, very probably stood upon the whole mid-way between salamanders and lizards, in regard to its bodily formation. Its descendants divided at an early period into two different lines, one of which became the common primary form of Reptiles and Birds, the other the primary form of Mammals.

Of all the three classes of Amniota, Reptiles (Reptilia, or Pholidota, also called Sauria in the widest sense), remain at the lowest stage of development, and differ least from their ancestors, the Amphibia. Hence they were formerly universally included among them, although their whole organization is much more like that of Birds than Amphibia. There now exist only four orders of Reptiles, namely,—Lizards, Serpents, Crocodiles, and Tortoises. They, however, form but a poor remnant of the exceedingly various and highly developed host of Reptiles which lived during the Mesolithic, or Secondary epoch, and predominated over all other Vertebrata. The immense development of Reptiles during the Secondary epoch is so characteristic that we could as well name it after those animals as after the Gymnosperms (p. 111). Twelve of the twenty-seven sub-orders, given on the accompanying table, and four of the eight orders, belong exclusively to the secondary period. These mesolithic groups are marked by an asterisk. All the orders, with the exception of Serpents, are found fossilized even in the Jura and Trias periods.

In the first order, that of Primary Reptiles, or Primary Creepers (Tocosauria), we class the extinct Thecodontia of the Trias, together with those Reptiles which we may look upon as the common primary form of the whole class. To the latter, which we may call Primæval Reptiles (Proreptilia), the Proterosaurus of the Permian system very probably belongs. The seven remaining orders must be considered as diverging branches, which have developed in different directions out of that common primary form. The Thecodontia of the Trias, the only positively known fossil forms of Tocosauria, were Lizards which seem to have been like the still living monitor lizards (Monitor, Varanus).

Of the four orders of reptiles now existing, and which, moreover, have alone represented the class since the beginning of the tertiary epoch, that of Lizards (Lacertilia) is probably most closely allied to the extinct Primary Reptiles, and especially through the monitors already named. The class of Serpents (Ophidia) developed out of a branch of the order of lizards, and this probably not until the beginning of the tertiary epoch. At least we at present only know of fossil remains of serpents from the tertiary strata. Crocodiles (Crocodilia) existed much earlier; the Teleosauria and Steneosauria belonging to the class are found fossil in large quantities even in the Jura; but the still living alligators are first met with in a fossil state in the chalk and tertiary strata. The most isolated of the four existing orders of reptiles consists of the remarkable group of Tortoises (Chelonia); fossils of these strange animals are first met with in the Jura. In some characteristics they are allied to Amphibia, in others, to Crocodiles, and by certain peculiarities even to Birds, so that their true position in the pedigree of Reptiles is probably far down at the root. The extraordinary resemblance of their embryos to Birds, manifested even at later stages of the ontogenesis, is exceedingly striking.

The four extinct orders of Reptiles show among one another, and, with the four existing orders just mentioned, such various and complicated relationships, that in the present state of our knowledge we are obliged to give up the attempt at establishing their pedigree. The most deviating and most curious forms are the Flying Reptiles (Pterosauria); flying lizards, in which the extremely elongated fifth finger of the hand served to support an enormous flying membrane. They probably flew about, in the secondary period, much in the same way as the bats of the present day. The smallest flying lizards were about the size of a sparrow; the largest, however, with a breadth of wing of more than sixteen feet, exceeded the largest of our living flying birds in stretch of wing (condor and albatross). Numerous fossil remains of them, of the long-tailed Rhamphorhynchia and of the short-tailed Pterodactylæ are found in all the strata of the Jura and Chalk periods, but in these only.

Not less remarkable and characteristic of the Mesolithic epoch was the group of Dragons (Dinosauria, or Pachypoda). These colossal reptiles, which attained a length of more than fifty feet, are the largest inhabitants of the land which have ever existed on our globe; they lived exclusively in the secondary epoch. Most of their remains are found in the lower cretaceous system, more especially in the Wealden formations of England. The majority of them were fearful beasts of prey (the Megalosaurus from twenty to thirty, the Pelorosaurus from forty to fifty feet in length). The Iguanodon, however, and some others lived on vegetable food, and probably played a part in the forests of the chalk period similar to that of the unwieldy but smaller elephants, hippopotami, and rhinoceroses of the present day.

The Beaked Reptiles (Anomodontia), likewise also long since extinct, but of which very many remarkable remains are found in the Trias and Jura, were perhaps closely related to the Dragons. Their jaws, like those of most Flying Reptiles and Tortoises, had become changed into a beak, which either possessed only degenerated rudimentary teeth, or no teeth at all. In this order, if not in the preceding one, we must look for the primary parents of the bird class, which we may call Bird Reptiles (Tocornithes). Probably very closely related to them was the curious, kangaroo-like Compsognathus from the Jura, which in very important characteristics already shows an approximation to the structure of birds.

The class of Birds (Aves), as already remarked, is so closely allied to Reptiles in internal structure and by embryonal development, that they undoubtedly originated out of a branch of this class. Even a glance at Plates II. and III. will show that the embryos of birds at a time when they already essentially differ from the embryos of Mammals, are still scarcely distinguishable from those of Tortoises and other Reptiles. The cleavage of the yolk is partial in the case of Birds and Reptiles, in Mammals it is total. The red blood-cells of the former possess a kernel, those of the latter do not. The hair of Mammals develops in closed follicles in the skin, but the feathers of birds and also the scales of reptiles develop in hillocks on the skin. The lower jaw of the latter is much more complicated than that of Mammals; the latter do not possess the quadrate bone of the former. Whereas in Mammals (as in the case of Amphibia) the connection between the skull and the first neck vertebra is formed by two knobbed joints, or condyles, in Birds and Reptiles those have become united into a single condyle. The two last classes may therefore justly be united into one group as Monocondylia, and contrasted to Mammals, or Dicondylia.

The deviation of Birds from Reptiles, in any case, first took place in the mesolithic epoch, and this moreover probably during the Trias. The oldest fossil remains of birds are found in the upper Jura (Archæopteryx). But there existed, even in the Trias period, different Saurians (Anomodonta) which in many respects seem to form the transition from the Tocosauria to the primary ancestors of Birds, the hypothetical Tocornithes. Probably these Tocornithes were scarcely distinguishable from other beaked lizards in the system, and were closely related to the kangaroo-like Compsognathus from the Jura of Solenhofen. Huxley classes the latter with the Dinosauria, and believes them to be the nearest relations to the Tocornithes.

The great majority of Birds—in spite of all the variety in the colouring of their beautiful feathery dress, and in the formation of their beaks and feet—are of an exceedingly uniform organization, in much the same way as are the class of insects. The bird form has adapted itself on all sides to the external conditions of existence, without having thereby in any way essentially deviated from the strict hereditary type of its characteristic structure. There are only two small groups, the feather-tailed birds (Saururæ) and those of the ostrich kind, which differ considerably from the usual type of bird, namely, from those with keel-shaped breasts (Carinatæ), and hence the whole class may be divided into three sub-classes.

The first sub-class, the Reptile-tailed, or Feather-tailed Birds (Saururæ), are as yet known only through a single, and that an imperfect, fossil impression, which, however, in being the oldest and also a very peculiar fossil bird, is of great importance. This fossil is the Primæval Griffin, or Archæopteryx lithographica, of which as yet only one specimen has been found in the lithographic slate at Solenhofen, in the Upper Jura system of Bavaria. This remarkable bird seems on the whole to have been of the size and form of a large raven, especially as regards the legs, which are in a good state of preservation; head and breast unfortunately are wanting. The formation of the wings deviates somewhat from that of other birds, but that of the tail still more so. In all other birds the tail is very short and composed of but few short vertebræ; the last of these have grown together into a thin, bony plate standing perpendicularly, upon which the rudder-feathers of the tail are attached in the form of a fan. The Archæopteryx, however, has a long tail like a lizard, composed of numerous (20) long thin vertebræ, and on every vertebra are attached the strong rudder-feathers in twos, so that the whole tail appears regularly feathered. This same formation of the tail part of the vertebral column occurs transiently in the embryos of other birds, so that the tail of the Archæopteryx evidently represents the original form of bird-tail inherited from reptiles. Large numbers of similar birds with lizard-tails probably lived during the middle of the secondary period; accident has as yet, however, only revealed this one fossil.

The Fan-tailed, or Keel-breasted birds (Carinatæ), which form the second sub-class, comprise all living Birds of the present day, with the exception of those of the ostrich kind, or Ratitæ. They probably developed out of Feather-tailed Birds during the first half of the secondary period, namely, in the Jura or Chalk period, by the hinder tail vertebræ growing together, and by the tail becoming shortened. Only very few remains of them are known from the secondary period, and these moreover only out of the last section of it, namely, from the Chalk. These remains belong to a swimming bird of the albatross species, and a wading bird like a snipe. All the other fossil remains of birds as yet known have been found in the tertiary strata.

The Bushy-tailed, or Ostrich-like Birds (Ratitæ), also called Running Birds (Cursores), the third and last sub-class, is now represented only by a few living species, by the African ostrich with two toes, the American and Australian ostrich with three toes, by the Indian cassowary and the four-toed kiwi, or Apteryx, in New Zealand. The extinct giant birds of Madagascar (Æpyornis) and the New Zealand Dinornis, which were much larger than the still living ostriches, also belong to this group. The Birds of the ostrich kind—by giving up the habit of flying, by the degeneration of the muscles for flying resulting from this, and of the breast bone which serves as their support, and by the corresponding stronger development of the hinder legs for running—have probably arisen out of a branch of the Keel-breasted birds. But possibly, as Huxley thinks, they may be the nearest relations of the Dinosauria and of the Reptiles akin to them, especially of the Compsognathus; at all events, the common primary form of all Birds must be looked for among the extinct Reptiles.

CHAPTER XXI.

PEDIGREE AND HISTORY OF THE ANIMAL KINGDOM.

IV. Mammals.

There are only a few points in the classification of organisms upon which naturalists have always agreed. One of these few undisputed points is the privileged position of the class of Mammals at the head of the animal kingdom. The reason of this privilege consists partly in the special interest, also in the various uses and the many pleasures, which Mammals, more than all other animals, offer to man, and partly in the circumstance that man himself is a member of this class. For however differently in other respects man’s position in nature and in the system of animals may have been regarded, yet no naturalist has ever doubted that man, at least from a purely morphological point of view, belongs to the class of Mammals. From this there directly follows the exceedingly important inference that man, by consanguinity also, is a member of this class of animals, and has historically developed out of long since extinct forms of Mammals. This circumstance alone justifies us here in turning our especial attention to the history and the pedigree of Mammals. Let us, therefore, for this purpose first examine the groups of this class of animals.

Older naturalists, especially considering the formation of the jaw and feet, divided the class of Mammals into a series of from eight to sixteen orders. The lowest stage of the series was occupied by the whales, which seemed to differ most from man, who stands at the highest stage, by their fish-like form of body. Thus Linnæus distinguished the following eight orders: (1) Cetæ (whales); (2) Belluæ (hippopotami and horses); (3) Pecora (ruminating animals); (4) Glires (gnawing animals and rhinoceroses); (5) Bestiæ (insectivora, marsupials, and various others); (6) Feræ (beasts of prey); (7) Bruta (toothless animals and elephants); (8) Primates (bats, semi-apes, apes, and men). Cuvier’s classification, which became the standard of most subsequent zoologists, did not rise much above that of Linnæus. Cuvier distinguished the following eight orders: (1) Cetacea (whales); (2) Ruminantia (ruminating animals); (3) Pachyderma (hoofed animals, with the exclusion of ruminating animals); (4) Edentata (animals poor in teeth); (5) Rodentia (gnawing animals); (6) Carnassia (marsupials, beasts of prey, insectivora, and bats); (7) Quadrumana (semi-apes and apes); (8) Bimana (man).

The most important advance in the classification of Mammals was made as early as 1816 by the eminent anatomist Blainville, who has already been mentioned, and who first clearly recognised the three natural main groups or sub-classes of Mammals, and distinguished them according to the formation of their generative organs as Ornithodelphia, Didelphia, and Monodelphia. As this division is now justly considered by all scientific zoologists to be the best, on account of solid foundation on the history of development, let us here keep to it also.

The first sub-class consists of the Cloacal Animals, or Breastless animals, also called Forked animals (Monotrema, or Ornithodelphia). This class is now represented only by two species of living mammals, both of which are confined to Australia and the neighbouring island of Van Diemen’s land, namely, the well-known Water Duck-bill (Ornithorhynchus paradoxus) with the beak of a bird, and the less known Beaked Mole (Echidna hystrix), resembling a hedgehog. Both of these curious animals, which are classed in the order of Beaked Animals (Ornithostoma), are evidently the last surviving remnants of an animal group formerly rich in forms, which alone represented the Mammalia in the secondary epoch, and out of which the second sub-class, the Didelphia, developed later, probably in the Jurassic period. Unfortunately, we as yet do not know with certainty of any fossil remains of this most ancient primary group of Mammals, which we will call Primary Mammals (Promammalia). Yet they possibly comprise the oldest of all the fossil Mammalia known, namely, the Microlestes antiquus, of which animals, however, we as yet only know some few small molar teeth. These have been found in the uppermost strata of the Trias, in the Keuper, first in Germany (at Degerloch, near Stuttgart, in 1847), later also in England (at Frome), in 1858. Similar teeth have lately been found also in the North American Trias, and have been described as Dromatherium sylvestre. These remarkable teeth, from the characteristic form of which we can conclude that they belonged to an insectivorous mammal, are the only remains of mammals as yet found in the older secondary strata, namely, in the Trias. It is possible, however, that besides these many of the other mammalian teeth found in the Jura and Chalk systems, which are still generally ascribed to Marsupials, in reality belong to Cloacal Animals. This cannot be decided with certainty owing to the absence of the characteristic soft parts. In any case, numerous Monotrema, with well-developed teeth and cloaca, must have preceded the advent of Marsupial animals.

The designation, “Cloacal animals” (Monotrema), has been given to the Ornithodelphia on account of the cloaca which distinguishes them from all other Mammals; but which on the other hand makes them agree with Birds, Reptiles, and Amphibia, in fact, with the lower Vertebrata. The formation of the cloaca consists in the last portion of the intestinal canal receiving the mouth of the urogenital apparatus, that is, the united urinary and genital organs, whereas in all other Mammals (Didelphia as well Monodelphia) these organs have an opening distinct from that of the rectum. However, in these latter also the cloaca formation exists during the first period of their embryonal life, and the separation of the two openings takes place only at a later date (in man about the twelfth week of development). The Cloacal animals have also been called “Forked animals,” because the collar-bones, by means of the breast bone, have become united into one piece, similar to the well-known fork-bone, or merry-thought, in birds. In all other Mammals the two collar-bones remain separated in front and do not fuse with the breast bone. Moreover, the coracoid bones are much more strongly developed in the Cloacal animals than in the other Mammalia, and are connected with the breast bone.

In many other characteristics also—especially in the formation of their internal genital organs, their auricular labyrinth, and their brain—Beaked animals are more closely allied to the other Vertebrata than to Mammals, so that some naturalists have been inclined to separate them from the latter as a special class. However, like all other Mammals, they bring forth living young ones, which for a time are nourished with milk from the mother. But whereas in all other Mammals the milk issues through nipples, or teats, from the mammary glands, teats are completely wanting in beaked animals, and the milk comes simply out of a flat, sieve-like, perforated patch of the skin. Hence they may also be called Breastless or Teatless animals (Amasta).

The curious formation of the beak in the two still living Beaked animals, which is connected with the suppression of the teeth, must evidently not be looked upon as an essential feature of the whole sub-class of Cloacal animals, but as an accidental character of adaptation distinguishing the last remnant of the class as much from the extinct main group, as the formation of a similar toothless snout distinguishes many toothless animals (for instance, the ant-eater) from the other placental animals. The unknown, extinct Primary Mammals, or Promammalia—which lived during the Trias period, and of which the two still living orders of Beaked animals represent but a single degenerated branch developed on one side—probably possessed a very highly developed jaw like the marsupial animals, which developed from them.

Marsupial, or Pouched Animals (Didelphia, or Marsupialia), the second of the three sub-classes of Mammals, form in every respect—both as regards their anatomy and embryology, as well as their genealogy and history—the transition between the other sub-classes—the Cloacal and Placental Animals. Numerous representatives of this group still exist, especially the well-known kangaroos, pouched rats, and pouched dogs; but on the whole this sub-class, like the preceding one, is evidently approaching its complete extinction, and the living members of the class are the last surviving remnants of a large group rich in forms, which represented the Mammalia during the more recent secondary and the earlier tertiary periods. The Marsupial Animals probably developed towards the middle of the Mesolithic epoch (during the Jura) out of a branch of the Cloacal Animals, and in the beginning of the Tertiary epoch again, the group of Placental Animals arose out of the Marsupials, and the latter then succumbed to the former in the struggle for life. All the fossil remains of Mammals known to us from the Secondary epoch, belong either exclusively to Marsupials, or partly perhaps to Cloacal animals. At that time Marsupials seem to have been distributed over the whole earth; even in Europe (France and England), well-preserved fossil remains of them have been found. On the other hand, the last off-shoots of the sub-class now living are confined to a very narrow tract of distribution, namely, to Australia, the Australasian, and a small part of the Asiatic Archipelago. There are also a few species still living in America, but at the present day not a single marsupial animal lives on the continent of Asia, Africa, or Europe.

The name of pouched animals is given to the class on account of the purse-shaped pouch (marsupium) existing in most instances on the abdominal side of the female animals, in which the mother carries about her young for a considerable time after their birth. This pouch is supported by two characteristic marsupial bones, also existing in Cloacal animals, but not in Placental animals. The young Marsupial animal is born in a much more imperfect form than the young Placental animal, and only attains the same degree of development which the latter possesses directly at its birth, after it has developed in the pouch for some time. In the case of the giant kangaroo, which attains the height of a man, the newly born young one, which has been carried in the maternal womb not much longer than five weeks, is not more than an inch in length, and only attains its essential development subsequently, in the pouch of the mother, where it remains about nine months attached to the nipple of the mammary gland.

The different divisions generally distinguished as families in the sub-class of Marsupial animals, deserve in reality the rank of independent orders, for they differ from one another in manifold differentiations of the jaw and limbs, in much the same manner, although not so sharply, as the various orders of Placental animals. In part they perfectly agree with the latter. It is evident that adaptation to similar conditions of life has effected entirely coincident or analogous transformations of the original fundamental form in the two sub-classes of Marsupials. According to this, about eight orders of Marsupial animals may be distinguished, the one half of the main group or legion of which are herbivorous, the other half carnivorous. The oldest fossil remains of the two legions (if the previously mentioned Microlestes and the Dromatherium are not included) occur in the Jurassic strata, namely, in the slates of Stonesfield, near Oxford. The slates belong to the Bath, or the Lower Oolite formation—strata which lie directly above the Lias, the oldest Jura formation. (Compare p. 15.) It is true that the remains of Marsupials found in the slates of Stonesfield, as well as those which were found later in the Purbeck strata, consist only of lower jaws. (Compare p. 29.) But fortunately the lower jaw is just one of the most characteristic parts of the skeleton of Marsupials. For it is distinguished by a hook-shaped process of the lower corner of the jaw turning downwards and backwards, which neither occurs in Placental nor in the (still living) Cloacal animals, and from the existence of this process on the lower jaws from Stonesfield, we may infer that they belonged to Marsupials.

PEDIGREE OF THE MAMMALIA

Of Herbivorous marsupials (Botanophaga), only two fossils are as yet known from the Jura, namely, the Stereognathus ooliticus, from the slates of Stonesfield (Lower Oolite), and the Plagiaulax Becklesii, from the middle Purbeck strata (Upper Oolite). But in Australia there are gigantic fossil remains of extinct herbivorous Marsupials from the diluvial period (Diprotodon and Nototherium) which were far larger than the largest of the still living Marsupials. The Diprotodon Australis, whose skull alone is three feet long, exceeded even the river-horse, or Hippopotamus, in size and upon the whole resembled it in the unwieldy and clumsy form of body. This extinct group, which probably corresponded with the gigantic placental hoofed animals of the present day—the hippopotami and rhinoceroses—may be called Hoofed Marsupials (Barypoda). Closely allied to them is the order of kangaroos, or Leaping Marsupials (Macropoda), which all have seen in zoological gardens. In their shortened fore legs, their very lengthened hind legs, and very strong tail, which serves as a jumping pole, they correspond with the leaping mice in the class of Rodents. Their jaw, however, resembles that of horses, and their complex stomach that of Ruminants. A third order of Herbivorous Marsupials corresponds in its jaws to Rodents, and in its subterranean mode of life, especially, to digging mice. Hence they may be termed Rodent Marsupials, or root-eating pouched animals (Rhizophaga). They are now represented only by the Australian wombat (Phascolomys). A fourth and last order of Herbivorous Marsupials is formed by the climbing or Fruit-eating Marsupials (Carpophaga), whose mode of life and structure resembles partly that of squirrels, partly that of apes (Phalangista, Phascolarctus).

The second legion of Marsupials, the Carnivorous Marsupials (Zoophaga), is likewise divided into four main groups or orders. The most ancient of these is that of the primæval, or Insectivorous Marsupials (Cantharophaga). It probably includes the primary forms of the whole legion, and possibly also those of the whole sub-class. At least, all the lower jaws from Stonesfield (with the exception of the Stereognathus) belong to Insectivorous Marsupials, and the still living Myrmecobius is their nearest relative. But some of those oolitic Primæval Marsupials possessed a larger number of teeth than all the other known mammals, for each half of the lower jaw of the Thylacotherium contained sixteen teeth (three incisors, one canine tooth, six pseudo, and six genuine molars). If the upper jaw, which is unknown, had as many teeth, then the Thylacotherium had no less than sixty-four teeth, just double the number possessed by man. The Primæval Marsupials correspond, on the whole, with the Insectivora among Placental animals, which order includes hedgehogs, moles, and shrew-mice. A second order, which has probably developed out of a branch of the last, consists of the Snouted, or Toothless Marsupials (Edentula), which resemble the Toothless animals, or Edentata, among the Placental animals by their tube-shaped snout, their degenerated jaws, and their corresponding mode of life. On the other hand, the mode of life and formation of the jaws of Rapacious marsupials (Creophaga) correspond with those of the genuine Beasts of Prey, or Carnivora, among Placental animals. This order includes the pouched marten (Dasyurus) and the pouched wolf (Thylacinus) in Australia. Although the latter attains to the size of a wolf, it is but a dwarf in comparison with the extinct Australian pouched lions (Thylacoleo) which were at least as large as a lion, and possessed huge canine teeth more than two inches in length. Finally, the eighth and last order is formed by the marsupials with hands, or the Ape-footed Pouched animals (Pedimana), which live both in Australia and America. They are frequently kept in zoological gardens, especially the different species of the genus Didelphys, and are known by the name of pouched rats, bush rats, or opossums. The thumb on their hinder feet is opposable to the four other toes, as in a hand, and by this they are directly allied to the Semi-apes, or Prosimia, among Placental animals. It is possible that these latter are really next akin to the marsupials with hands, and that they have developed out of their long since extinct ancestors.

It is very difficult to discover the genealogy of Marsupials, and this more especially because we are but very imperfectly acquainted with the whole sub-class; and the Marsupials of the present day are evidently only the last remnants of a group that was at one time rich in forms. It is possible that Marsupials with hands, those with snouts, as well as rapacious Marsupials, developed as three diverging branches out of the common primary group of Primæval Marsupials. In a similar manner, on the other hand, the rodent, leaping, and hoofed Marsupials have perhaps arisen as three diverging branches out of the common herbivorous primary group, that is, out of the Climbing Marsupials. Climbing and Primæval Marsupials might, however, be two diverging branches of the common primary forms of all Marsupials, that is, of the Primary Marsupials (Prodidelphia), which originated during the older secondary period out of Cloacal animals.

The third and last sub-class of mammals comprises the Placental animals, or Placentals (Monodelphia, or Placentalia). It is by far the most important, comprehensive, and most perfect of the three sub-classes; for the class includes all the known mammalia, with the exception of Marsupials and Beaked animals. Man also belongs to this sub-class, and has developed out of its lower members.

Placental animals, as their name indicates, are distinguished from all other mammals, more especially by the formation of a so called placenta. This is a very peculiar and remarkable organ, which plays an exceedingly important part in nourishing the young one developing in the maternal body. The placenta (also called after-birth) is a soft, spongy, red body, which differs very much in form and size, but which consists for the most part of an intricate network of veins and blood vessels. Its importance lies in the exchange of substance between the nutritive blood of the maternal womb, or uterus, and the body of the germ, or embryo. (See vol. i. p. 298.) This very important organ is developed neither in marsupials nor in beaked animals. But placental animals are also distinguished from these two sub-classes by many other peculiarities, thus more especially by the absence of marsupial bones, by the higher development of the internal sexual organs, and by the more perfect development of the brain, especially of the so-called callous body or beam (corpus callosum), which, as the intermediate commissure, or transverse bridge, connects the two hemispheres of the large brain with each other. Placental animals also do not possess the peculiar hooked process of the lower jaw which characterizes Marsupials. The following classification (p. 246) of the most important characteristics of the three sub-classes will best explain how Marsupials, in these anatomical respects, stand midway between Cloacal and Placental animals.

Placental animals are more variously differentiated and perfected, and this, moreover, in a far higher degree, than Marsupials, and they have, on this account, long since been arranged into a number of orders, differing principally in the formation of the jaws and feet. But what is even of more importance than these, is the different development of the placenta, and the manner of its connection with the maternal uterus. For in the three lower orders of Placental animals, in Hoofed animals, Whales, and Toothless animals, the peculiar spongy membrane, which is called the deciduous membrane, or decidua, and which connects the maternal and the fœtal portions of the placenta, does not become developed. This takes place exclusively in the seven higher orders of Placental animals, and we may, therefore, according to Huxley, class them in the main group of Deciduata, or animals with decidua. They are contrasted with the three first-mentioned legions of indeciduous animals, or Indeciduata.

But in the various orders of Placental animals the placenta differs not only in important internal differences of structure, which are connected with the absence or the presence of a decidua, but also in the external form of the placenta itself. In the Indeciduata it consists, in most cases, of numerous, single, scattered bunches or tufts of vessels, and hence this group may be called tufted placental animals (Villiplacentalia). In the Deciduata, however, the single tufts of vessels are united into a cake, which appears in two different forms. In the one case it surrounds the embryo in the form of a closed band or ring, so that only the two poles of the oval egg bladder are free of tufts; this is the case in animals of prey (Carnaria) and the pseudo-hoofed animals (Chelophora), which may consequently be comprised as girdled-placental animals (Zonoplacentalia). In the other Deciduata, to which man also belongs, the placenta is a simple round disc, and we therefore call them disc-placentals (Discoplacentalia). This group includes the five orders of Semi-apes, Gnawing animals, Insectivora, Bats, and Apes, from the latter of which, in the zoological system, man cannot be separated.