98

JURASSIC MAMMALS

CHAP.

the family, is Plagiaulax. As it is not Triassic, the consideration of its characters will be deferred until later. Microlestes is a Rhaetic genus, known from rocks in Germany and England; but it is entirely based upon molar teeth. M. antiquus has a tworooted molar of an elongated form with a row of tubercles on either side of a median groove, which traverses the long axis of the tooth. To some extent the teeth of the ancient form resemble those of Ornithorhynchus. Microlestes has been sometimes spoken of as a Marsupial, but Mr. Tomes has found that it does not show one very universal character of the Marsupial teeth: it has not those continuations of the dentinal tubes which traverse the enamel in all Marsupials that have been examined with the sole exception of the Wombat.

1

The rarity of the remains of mammals in these earliest rocks of the Secondary epoch has been accounted for in another way from that which has been suggested above. It may be that the group Mammalia was not evolved in Europe at all, and that the stray remains which have been found in that continent represent the fragmentary remnants of a few scattered immigrants which heralded the later invasion of more numerous genera during the Jurassic period.

The Mammals of the Jurassic Period.-Some of the Allotheria or Multituberculata described in the last section occur in the rocks of this early part of the Secondary epoch. They are doubtful in position, as already stated; some of them indeed, as for instance Tritylodon and Dromatherium, are possibly not mammals at all, while the remainder probably belong to a existent order of mammals. Along with these dubious creatures are the fragmentary remains of small animals which are not merely mammals, but in all probability definitely Marsupials. It is true that here again we have little beyond lower jaws and teeth to deal with; so that there may be less certainty in referring them to the Marsupials than appears to be the opinion of the majority of Palaeontologists.

Professor Osborn in fact considers that the Mesozoic mammals consist of three groups: (1) The Multituberculata, including the Bolodontidae, Stereognathidae, Plagiaulacidae, Polymastodontidae, and possibly the Tritylodontidae (which, however, are regarded by him and by others as more probably reptiles of the 1 Dental Anatomy, 5th ed. 1898, p. 304.

IV

PLAGIAULAX

99

were

Theromorphous group). (2) The Triconodonta, which Marsupials, though in all probability with a complete succession of teeth and with an allantoic placentation. This group will include the genera Phascolotherium and Amphilestes, as well as Triconodon and Spalacotherium. Finally we have (3) the Trituberculata (or Insectivora Primitiva) with the genera Amphitherium, Peramus, Amblotherium, Stylacodon, and Dryolestes.

We shall take these three groups in order. The Multituberculata have already been to some extent defined, if such a word can be used to express the summation of the very scanty information at our disposal. Of this group, Plagiaulax is a genus which occurs in the Purbeck beds; it is only known by lower jaws implying an animal of the size of a Rat or rather smaller. The jaws have in

front a large incisor which looks Rodent-like, and also like those of the Diprotodont Marsupials; but it is held that these teeth did not. grow from persistent pulps, and there is in any case no anterior thickened coating of enamel. Canines are absent; the diastema is followed by four premolars increasing progressively in size and possessing somewhat complicated grinding surfaces. These surfaces

are formed by several obliquely-set ridges. The succeeding teeth are termed molars on account of their difference in structure, and there are but two of them on each side. The molars are of a pattern common in the Multituberculata; the centre is hollowed, and the raised rim is beset by tubercles. Other Jurassic genera of Multituberculates are Bolodon, Allodon, and Stereognathus. All of these possess the same multituberculate molars.

Of the Triconodonta the type-genus is Triconodon. This genus is better known than most Jurassic mammals, since both the upper and the lower dentition have been described. It appears to have possessed the typical Eutherian dentition of forty-four teeth, to which a fourth molar is added in some species. The great difference between the molars and premolars argues a complete toothchange. The genus is American as well as European.

Spalacotherium has more molars, five or six.

Phascolotherium bucklandi, on the other hand, is a much older type in the form of its teeth. There are, however, not so many of them as in Amphitherium; Phascolotherium has but two premolars and five molars, making a total of forty-eight teeth. The teeth are of the triconodont form, the three cusps being in line, and the middle one the largest.

100

STONESFIELD MAMMALS

CHAP.

Amphilestes has teeth of the same pattern but has more of them, the premolars and molars being respectively four and five. All these animals had the lower jaw inflected. Whether they are all Marsupials or not, it is clear that Phascolotherium and Amphilestes should be united and placed away from Amphitherium on account of the more primitive form of their teeth.

We next come to the Trituberculata.

Among the most celebrated of these remains are a few jaws discovered in the Stonesfield slates near Oxford, and examined by Buckland, Cuvier, and some of the most eminent naturalists of the beginning of the last century. These jaws have been lately submitted to a careful re-examination by Mr. Goodrich,' who has increased our knowledge of the subject by exposing from the rocky matrix in which the jaws lie fresh details of their structure; it is probable therefore that now all that there is to be learnt from these specimens has been recorded.

The

Amphitherium prevostii was a creature about the size of a Rat. Its jaw was first brought to Dean Buckland about the year 1814, and described six years later. Buckland thought the jaw to be that of an Opossum, an opinion in which Cuvier concurred. jaw, however, is marked by a groove running along its length, and this groove was regarded by de Blainville as evidence of the composition of the jaw out of more than one element, which would naturally lead to its being regarded as the jaw of a reptile. This species and another named after Sir Richard Owen have a dental formula which, like that of the Marsupials, is large as compared with that of the Placental mammals; it runs: I 4, C 1, Pm 5, M 6—i.e. 64 teeth altogether. This is a larger number than we find in any existing Marsupial. But as in Marsupials, and in certain Insectivora also, the angle of the jaw is inflected. These teeth are of the tritubercular pattern with a "heel." They are in fact closely like those of the living Myrmecobius; but not, it should be remarked, unlike those of certain Insectivora.

The Mammals of the Cretaceous Period. At one time. there was a totally inexplicable gap between the Jurassic and the basal Eocene, a series of strata which occupy an enormous expanse of time in the history of the earth having appeared to

1 "On the Fossil Mammalia from the Stonesfield Slate," Quart. Journ. Mier. Sei. xxxv. 1894, p. 407.

2 This groove has been found in the existing Myrmecobius, see p. 154.

IV

EARLY TERTIARY MAMMALS

ΙΟΙ

be devoid of mammalian remains. This gap, however, has been filled up by the discovery of mammalian remains in the North American Laramic formation, which seems to be clearly of Cretaceous age. Furthermore, it is held by some that the Purbeck beds are more properly to be placed with the Cretaceous, which would then necessitate the consideration under the present heading of some of the types already dealt with; and if, as is suggested in the following section, the lowest so-called Eocene beds are really referable to the Cretaceous, there is no lack of mammalian remains in that period. And, moreover, it was in that case the Cretaceous period which witnessed the evolution of the existing orders of Placental mammals. Otherwise the mammalian remains of the Cretaceous agree with those of the Jurassic. We find remains

of the Multituberculata in fragments of Plagiaulacidae and Polymastodontidae. Ptilodus is a genus which has two premolars; and Meniscoessus is another multituberculate from the same Laramic formation. The other detached fragments of mammals are thought by Osborn to represent both Placentals and Marsupials.

The Mammals of the Tertiary Period.-Unless the lowest beds of the earliest Tertiary period, the Eocene, such as the Torrejon of North America, should be in reality referred to the Cretaceous, there is no evidence that the modern groups of Mammalia existed before the present epoch of the earth's history. It is probable, however, that the Eutheria as a group were Mesozoic. The fossil jaws that have been considered in the last chapter may quite probably be primitive Eutherians, or even divisible, as believed by Professor Osborn, into Marsupials and Insectivores. In the Tertiary, however, apart from the question as to the nature of the Puerco and Torrejon formations, and as to certain South American strata whose fossil contents have been investigated by Professor Ameghino, we find the first traces of mammals definitely referable to existing orders, or to be distinctly compared with existing orders. Since, however, representatives of types which have obvious relationships to modern types appear in considerable profusion in the very earliest strata of the Eocene, it seems clear that much remains to be discovered in beds earlier than these. Confining ourselves, however, to facts and to comparisons which can be made on more than a few lower jaws and scattered teeth, which is practically all that we

102

PRIMITIVE UNGULATA

CHAP.

possess of earlier mammals, we must arrive at the general conclusion that two of the existing larger groups of the Eutherian, non-Marsupial, mammals were differentiated at quite the beginning of the Eocene, and were represented by forms from which it is possible to derive at least the existing Carnivora, Insectivora, Artiodactyla, and Perissodactyla. These were the Creodonta and the Ungulate Condylarthra. In addition to these we may enumerate as very early types the Lemuroidea, represented by such forms as Indrodon in the New World, and (though later) by Necrolemur, etc., in the Old World, and the Edentata, if we are to allow as their ancestors the Ganodonta.

The early Eocene strata also contain representatives of at least one order, the Amblypoda, which increased subsequently, but has died out without descendants, unless we are to believe with some that the Elephants are to be derived from these Eocene "pachyderms." In later Eocene times the great majority of the existing orders, and even subdivisions of orders, are to be met with; and there are in addition such totally extinct orders as the Typotheria, Ancylopoda, and Tillodontia. Coupled with this gradual specialisation in the orders of Eutherian mammals, there is naturally a vast increase in the number of generic and family types. This culminates perhaps in the Miocene, from which time there has been a gradual decline in mammalian variety, so that it is justly said that we live now in an epoch which is impoverished of mammals. This gradual decay has persisted until to-day, as is witnessed by the extinction of the Rhytina and the Quagga, and the growing rarity of the White Rhinoceros and the American Bison.

The early Eutherian stock consisted of small mammals with small heads and slender, long tails. The limbs were pentadactyle, ensheathed in claws or broader hoofs. The forelimbs may have been partly prehensile. The teeth were fortyfour, completely differentiated into incisors, canines, molars, and premolars; and there appears to have been a complete diphyodontism. The canines were not greatly enlarged, and no diastema separated any of the teeth. The molars were bunodont or of a more cutting pattern, with some five or six tubercles. These animals were, moreover, very small-brained. This early stock is represented by Creodont and Condylarthrous animals, the exact boundaries between which are hardly marked in the