VI. †Miacidæ.

†Didymictis, Paleoc. and low. Eoc. †Viverravus, mid. Eoc. †Miacis, low. Eoc. †Uintacyon, low. to up. Eoc. †Oödectes, mid. Eoc. †Vulpavus, do. †Palæarctonyx, do.

The †Creodonta were an extremely varied assemblage, of carnivorous, omnivorous and presumably insectivorous habits, so that few statements, not subject to exceptions, can be made of them all. Only seven genera are known from skeletons, and several more from skulls, but most are represented only by jaws and teeth; limb- and foot-bones, however, give us a conception of the general structure of a considerable number. As a rule, the dentition was complete, according to the formula, i 3/3, c 1/1, p 4/4, m 3/3, × 2 = 44, but the first premolar or the last molar may be lost. The canines were always large, as was befitting for beasts of prey. In only one family, the Miacidæ, were the carnassial teeth confined to a single pair and those the same as in the Fissipedia, the fourth upper premolar and first lower molar; in all the other families there were either no sectorial teeth, or else there was more than one pair. In the Fissipedia the first is the largest of the lower molars, while in the †Creodonta (except the †Miacidæ) it was usually the smallest. The premolars were generally simple, compressed-conical teeth and the molars, with all their great variety, may be reduced to a common plan; those of the upper jaw were primitively tritubercular, with a triangle of two external and one internal cusps, and those of the lower jaw were in two distinct parts, an anterior, elevated triangle of three cusps and a low heel of two.

The skull was almost always very large in proportion to the size of the animal; the cranium, though long, was of small capacity and the face varied much in length in the different families. Primitively, the face and jaws were short in correlation with the small size of the teeth, and this primitive condition was modified in two opposite directions; in one the face and jaws were elongated, as the teeth enlarged, and in the other they were shortened still further. The zygomatic arches were stout and curved out strongly from the sides of the skull, making very wide openings, and, in almost all cases, the sagittal and occipital crests were very high, as would be necessary from the combination of powerful jaws and small brain-case (see [p. 63]). The tympanic bullæ were not ossified. The brain was extremely small, especially in the more ancient genera, and the convolutions were almost always few and simple, which indicates a low grade of intelligence and very marked inferiority to the Fissipedia.

In all the genera of which sufficient material has been obtained the body was long and had 19 or 20 trunk-vertebræ: in the lumbar and posterior part of the dorsal regions the processes by which the successive vertebræ were articulated together (zygapophyses) were cylindrical and interlocking, as in the artiodactyl ungulates ([p. 360]). To this general statement, the †Miacidæ formed a partial exception. The tail was very long and heavy in all the forms of which the caudal vertebræ are known, and this was probably true of all. The limbs were short and generally heavy; the femur had the third trochanter and the humerus, save in a few of the later genera, the epicondylar foramen, and the manus could, in nearly all, be freely rotated. Except in the most advanced forms of one family, the †Mesonychidæ, the feet were five-toed and plantigrade, or semi-plantigrade, and of decidedly primitive structure. The scapho-lunar bone of the Fissipedia (see [p. 519]) was not formed, its three elements, with very few exceptions, remaining separate. The astragalus nearly always had a shallow groove, or none at all. The claws were thick and blunt and the ungual phalanges cleft at the end, except in the †Arctocyonidæ and †Miacidæ, which had sharp claws and uncleft phalanges.

From this brief description, it is obvious that the †Miacidæ occupied a very isolated position among the †creodonts and, in my judgment, it would be better to transfer that family to the Fissipedia and include the others in a separate order.

Throughout the Paleocene and Eocene epochs the †Creodonta were numerous and varied, the first of the Fissipedia appearing in the upper Eocene. Till then the †creodonts were the only predaceous mammals in North America and Europe, and they were especially abundant in the former. Most members of the suborder and all the Paleocene forms were of small or moderate size, but some of the Eocene species were very large. In the Uinta the †creodonts were greatly decreased in numbers and in the White River there were only two genera of one family, the †Hyænodontidæ, and since the Oligocene the suborder has been extinct.

1. †Miacidæ. Fissipede-like †Creodonts

It is unfortunate that no member of this family is known from a complete skeleton, but the material collected is sufficient to give a fairly adequate conception of these most interesting animals. These were the only †creodonts with a single pair of carnassials, the fourth upper premolar and first lower molar, but in some of the genera the carnassials did not differ greatly from the other teeth. In the various genera the skull differed considerably in length and in the proportions of cranium and face; the brain-case was larger than in most other †creodonts and the brain more advanced, though smaller than in the fissipedes, and the sagittal and occipital crests were very prominent; the tympanic bullæ were not ossified. The humerus had the epicondylar foramen and the femur the third trochanter; in the wrist the scaphoid, lunar and central were separate, almost the only important difference from the Fissipedia and merely the primitive stage of the latter. The feet were pentadactyl and the digits were arranged in spreading fashion; the claws were small, sharp and partially retractile and the ungual phalanges not cleft at the tip.

Within the family several different phyla may be distinguished, one of which (†Miacis†Uintacyon) led to the dogs, another to the †bear-dogs, or †amphicyons. A third phylum (†Didymictis†Viverravus) is by several authorities regarded as ancestral to the civet family, or viverrines, of the Old World, and a fourth (†Oödectes, †Vulpavus) as the forerunner of the kinkajous (Potos). Except for the connection with the dogs, the hiatus in time between the supposed ancestors and descendants is too great to permit any confident statements. It seems very probable, however, that the †Miacidæ represented the common stock, from which the fissipede families were all derived, directly or indirectly, though for most of them the details of the connection remain to be learned.