Chapter VII: The Successive Mammalian Faunas of North and South America (2)

_North America._—Upper Miocene beds cover extensive areas of the Great Plains region and are scattered from Montana far into Mexico. The rich fauna is an outgrowth and development of that of the middle Miocene, with but few immigrant additions and, on the other hand, passes so gradually into that of the lower Pliocene, that any line of separation between them is very difficult to draw. The rodents, numerous as they are among the fossils, are almost certainly very incompletely represented in the collections; the families are almost all still in existence, but nearly every genus is extinct, and thus the vernacular names used to designate them must be understood in a broad sense. Hares, mice, pocket-gophers, squirrels, marmots, beavers and the extraordinary †mylagaulids were all abundant.

In even more strongly marked sense must the broad meaning for the vernacular names of the other mammals be emphasized, for we have to deal almost exclusively with extinct genera, which differed much from their modern descendants. Many of the Carnivora have been obtained; there were numerous dogs, some rivalling the largest of existing bears in size, true felines and †sabre-tooth tigers, which were smaller and lighter animals than the great beasts of the Pleistocene; weasels, martens, otters and raccoons, but no bears. The bears, a family of Old World origin, are not certainly known in America before the Pleistocene, but had probably reached this continent in the Pliocene.

As is so very generally true, the commonest and best-preserved of the fossils are those of the hoofed animals. The †mastodons were of the four-tusked kind (_†Gomphotherium_ or _†Trilophodon_), the skull and teeth of which differed so markedly from those of the true elephants. The relatively small, low-crowned and simple grinding teeth were common to all the †mastodons, but the tusks were different from those of the larger members of the group. The upper tusks were comparatively short and nearly straight and retained a band of enamel, while the lower tusks were still shorter, chisel-shaped and so worn as to prove that they were regularly used, no doubt in cropping leaves; the shortness of these lower tusks was compensated for by the great elongation of the lower jaw. The head was proportionately broad and low and, for Proboscidea, these were small animals, not more than five or six feet high at the shoulder. The body, limbs and feet had already attained substantially their modern grade of structure, advance among the Proboscidea being chiefly restricted to the teeth and skull.

Four families of Perissodactyla were represented in the upper Miocene. The rhinoceroses, which were very abundant, were present in considerable variety; some were hornless, others had a single small horn on the end of the nose. Among these rhinoceroses there was much difference in bodily proportions, some being extremely heavy, with very short legs and feet, and these were the commonest, while others had longer legs and less massive bodies. Tapirs, on the contrary, would seem to have been scantily represented; at least, they are rare among the fossils. The extraordinary, aberrant †chalicotheres, perissodactyls with claws instead of hoofs, still persisted, but are far better known from the lower Miocene, in connection with which they will be described. The dominant perissodactyl family was that of the horses, of which no less than five genera are already known. There were some with very low-crowned teeth, which must have fed principally by browsing upon leaves and such soft diet; but the grazing kinds, which had high-crowned, cement-covered and very complex grinding teeth, had come to the fore. Still retaining three toes in each foot, with the middle toe so enlarged as to bear nearly the entire weight, save in snow or soft ground, these eminently cursorial animals, which had the slender limbs of a deer, must have roamed the plains in great herds.

Still commoner were the Artiodactyla. Many species of grazing camels, which were the predominant artiodactyl family in North America during upper Miocene times, were the ancestors both of the true camels of the Old World and the South American llamas. †Giraffe-camels have not yet been found and no doubt they were much less abundant than in the middle Miocene, but that they had not completely disappeared is shown by their recurrence in the Pliocene. As compared with earlier ages, the †oreodonts had begun a rapid decline and had lost notably both in numbers and variety, but one most curious beast (_†Pronomotherium_, Fig. 197, p. 375) marked the final step in the development of the short-faced, proboscis-bearing series, which may be traced back to its beginnings in the Oligocene. In this wonderful creature the skull was so short and deep as to suggest that of a gorilla or some other great ape. No other artiodactyls even approximate these later proboscis-bearing †oreodonts in the altogether exceptional form of the skull. Grazing †oreodonts (_†Merychyus_), of moderate and small size with high-crowned teeth, were evidently quite common on the upper Miocene plains. The †hornless deer and “†deer-antelopes” differed but little from those of the lower Pliocene. Peccaries were fairly abundant.

The upper Miocene fauna was especially characterized by the large number of mammals, belonging to several different orders, which had acquired the high-crowned, persistently growing pattern of grinding teeth. Many of the horses, camels, ruminants and rodents displayed this structure, and, as was first pointed out by Kowalevsky, the explanation is probably to be found in the spread of grassy plains at the expense of the forests. On account of the silica which they contain, the grasses are very abrasive and rapidly wear the teeth down. In adaptation to this new source of abundant and nutritious food, many kinds of mammals developed a form of tooth which was fitted to compensate by growth for the loss through abrasion.

The middle Miocene, small areas of which occur in Montana, eastern Oregon and northeastern Colorado, has received various local names, the typical one being the Deep River of Montana. Very probably, these scattered areas are not strictly contemporaneous, but form a closely connected series. That a land-connection with the eastern hemisphere existed, is made clear by the appearance of several unmistakably Old World types of animals and the beginnings of migration from South America are perhaps also to be noted, though this cannot be positively stated. The evidence for the South American connection is the finding in the middle Miocene of Oregon of what are believed to be the earliest remains of †ground-sloths yet discovered in North America, but the material is too scanty for altogether certain determination.

The smaller animals are not very well represented in the middle Miocene faunas, as conditions appear to have been unfavourable to their preservation; something is known of them, nevertheless. The very curious extinct family of rodents known as the †Mylagaulidæ, the presence of which was noted in the upper Miocene and lower Pliocene, first appeared here. These †mylagaulids, which were distantly related to the modern Sewellel (_Aplodontia rufa_), were characterized by the great enlargement and complication of one of the grinding teeth in each jaw and the consequent reduction of the others. One genus of this family, as in the Pliocene, had the peculiarity, unique among rodents, of developing a large horn upon the nose, like a miniature rhinoceros. Among the Carnivora, we find a great variety of dogs, large and small, all belonging to extinct genera, as indeed is true of the other carnivores also. True felines have been found, but as yet, none of the †sabre-tooth series; the abundance of the latter, however, in both preceding and succeeding formations, is sufficient proof that the discovery of them in the middle Miocene is merely a question of time. Mustelines were present, and especially noteworthy is the appearance of the first American otters, immigrants from the Old World.

Of the hoofed animals, the most interesting are the Proboscidea, the most ancient of which that are definitely determinable in America occur in this horizon. The place of origin and ancestry of these animals were long exasperating puzzles. Appearing suddenly in the Miocene of Europe and North America, in which regions nothing was known that could, with any plausibility, be regarded as ancestral to them, they might as well have dropped from the moon, for all that could be told concerning their history. The exploration of the Eocene and Oligocene beds of Egypt has dispelled the mystery and shown that Africa was the original home of the group, whence they gradually spread to every continent except Australia. Little is known of these earliest American proboscideans, but they were doubtless small †mastodons of the four-tusked type.

Among the Perissodactyla, the rhinoceroses were perhaps the most conspicuous; the native American stocks of this family appear to have mostly died out and to have been replaced by two or more phyla of immigrants from the Old World, some of which were hornless, others had a small horn on the tip of the nose and others again had a second and smaller horn on the forehead. Tapirs, though unquestionably present, are rare as fossils and not well known. Several distinct phyla of horses may be distinguished, which were like small ponies in size, but of more slender form; they were all three-toed, but there were marked differences among them with regard to the degree to which the middle toe (the third of the original five) had been enlarged to carry the whole weight and the lateral toes (second and fourth) reduced to mere “dew-claws.” While browsing horses, with low-crowned teeth, still persisted in large numbers, we find also the extremely interesting beginnings of the highly complex, cement-covered and high-crowned teeth of the grazing kinds. The clawed †chalicotheres were present, though very little is known about them because of the fragmentary character of the remains.

The Artiodactyla were much more varied and abundant, though they did not rival the great assemblage of these animals found in the European Miocene. Of the peccaries little more can be said than that they were present in these faunas. The †oreodonts were very numerous, both individually and generically; two stages of the proboscis-bearing kind are found here together, the older, long-faced genus (_†Promerycochœrus_) surviving from the Oligocene, while the newer Miocene type was short-faced and had a moderate proboscis (see Fig. 196, p. 373). Others had more the proportions of peccaries and still others were very small and presumably aquatic in habits. Camels abounded, both the grazing kinds which were ancestral to the modern forms of South America and Asia, and the great, browsing †giraffe-camels. The †hornless deer and the antlered †deer-antelopes were much like those of the Upper Miocene, slender and graceful little creatures, and there were also considerably larger ruminants (_†Dromomeryx_) with straight, simple and non-deciduous horns, which may be called antelopes.

The line of division between the lower Miocene and the uppermost Oligocene is a very obscure and difficult one to draw. Personally, I prefer to begin the Miocene with the widespread formation of the Great Plains, which has been variously named Arikaree, Harrison, Rosebud, etc., but this is a moot point. Concerning the lower part of these beds Osborn says: “They may be either: (1) Upper Oligocene or (2) transitional from Oligocene to Miocene, or (3) of pure Lower Miocene age.” The upper division is referred to the Miocene without question by any one, but for the purposes of this rapid sketch it will be best to treat the two faunas together. This many-named formation, for which the term _Arikaree_ is here employed, as having priority, is found over extensive areas of South Dakota, northern Nebraska and central Wyoming. The fauna was almost entirely a development from that of the North American Oligocene, with very little admixture of foreign elements, so that the land communication with the eastern hemisphere must have been difficult. In this, as in most of the Miocene formations, the smaller mammals are not fairly represented, and it is evident that much remains to be learned with regard to them; this is especially true of the upper division of this stage.

The rodents, which were fairly numerous, were directly continuous with those of the upper Oligocene and included forms which were more or less distantly connected with the modern hares, squirrels, beavers, sewellels, pocket-gophers and kangaroo-rats. A few Insectivora of doubtful reference have been found. Among the Carnivora there was also considerable variety: dogs, large and small, were abundant, but all of them were decidedly primitive from the modern standpoint; the cats were represented both by the true felines, which were probably immigrants, and by the †sabre-tooth series. There were several large and powerful mustelines, or members of the weasel family, which were likewise immigrants, one of which resembles in many ways the modern Wolverene (_Gulo_). Very interesting is the beginning of the raccoon family (Procyonidæ) or, at least, what is believed to be such, which arose from a branch of the dogs; this most ancient of the raccoons was _†Phlaocyon_, a small and slender animal.

The earliest traces of the Proboscidea in America have been reported from this formation, but the fragmentary specimens are inconclusive. The Perissodactyla are among the commonest fossils. The rhinoceroses belonged to native stocks, including both the horned and hornless forms. The horned genus (_†Diceratherium_) differed from all other rhinoceroses in having a transverse _pair_ of horns on the nose, and the species of the lower Miocene were quite small and light; the hornless genus (_†Cænopus_) was a larger and heavier animal. Tapirs are rare as fossils and consequently not well known. While there were several kinds of horses, they all agreed in having short-crowned and relatively simple grinding teeth and three-toed feet; they were smaller and of lighter, more slender build than those of the middle Miocene. The wonderful aberrant perissodactyls with clawed feet, the †chalicotheres (suborder †Ancylopoda), appear to have been more abundant in the Arikaree than at any other time in North America, though their history in this continent extends from the middle Eocene to the lower Pliocene. _†Moropus_, the lower Miocene genus, was as grotesque a creature as could well be imagined and, in advance of experience, no one ever did imagine such a beast. With rather small and somewhat horse-like head, long neck, long fore limbs and shorter hind limbs, these extraordinary animals united short, three-toed feet, which were armed with enormous claws. The long persistence (to the Pleistocene of Asia) and wide geographical range of the †chalicotheres are sufficient evidence that their very unusual structure must have been advantageous to them, but the problem of their habits and mode of life is still unsolved. From the character of the teeth, the long neck and fore limbs, it may, however, be inferred that they fed chiefly upon the leaves of trees.

Even more numerous and varied were the Artiodactyla. Peccaries of a primitive sort were common, and we find the last of the series of “†giant pigs,” which had been a very conspicuous group throughout the Oligocene. The lower Miocene genus, _†Dinohyus_, was a monstrous beast, six feet or more in height, with formidable canine tusks and a very long head made grotesque by bony excrescences upon the skull and jaws. For a pig, the legs were very long and the feet slender, having but two toes. The †oreodonts were present in great numbers, both small and large forms; except for bodily stature and modifications of the head, they all looked very much alike; _†Merycochœrus_, with its incipient proboscis, here made its first appearance. The last representatives of a family (†Hypertragulidæ) of small and graceful artiodactyls are found in this formation. One of these (_†Syndyoceras_, _see_ Fig. 215, p. 403), an animal considerably larger than the existing Musk-Deer, was in its way even more bizarre-looking than the †chalicotheres; with an antelope-like head, it had four horns, one pair over the eyes, curving inward, and a shorter pair, with outward curvature, on the muzzle. Another genus (_†Hypertragulus_) was very much smaller and very slender.

The camels were beginning to diversify and give rise to several phyla. One of the genera (_†Protomeryx_), which did not much exceed a sheep in size, probably represented the main stock, which led to the camels and llamas of to-day. A second (_†Stenomylus_) was a still smaller animal, with remarkably long and slender legs, and might be called a “gazelle-camel,” while a third (_†Oxydactylus_, _see_ Fig. 209, p. 392), which was larger and apparently the beginning of the †giraffe-camels, was noteworthy for its long neck. All of these lower Miocene camels had deer-like hoofs, the characteristic pad or cushion which gives such an exceptional appearance to the feet of modern llamas and camels not being fully developed till a later period. A very important new element in the North American fauna was the appearance of the first deer (_†Blastomeryx_), which came in the latter part of the Arikaree stage and were the forerunners of a renewed immigration from the Old World, which had been broken off during the upper Oligocene. This, however, is a disputed point; Professor Osborn and Dr. Matthew believe that these animals were truly indigenous and derived from a long line of American ancestry. The same genus continued through the middle Miocene, as we have already seen, and therefore no further description of it is called for.

The limits of the _South American_ Miocene are very doubtful. The Paraná formation, here regarded as lower Pliocene, may prove to be more properly referable to the upper Miocene. No other upper Miocene is known.

To the earlier, probably middle, Miocene may be referred the wonderful Santa Cruz fauna of Patagonia. It is extremely difficult to convey to the reader any adequate conception of this great assemblage of mammals, because most of them belonged to orders which have altogether vanished from the earth and are only remotely like the forms with which we are familiar in the northern hemisphere. To one who knows only these northern animals, it seems like entering another world when he begins the study of the Santa Cruz fossils. If any North American mammals had then entered South America, which is not probable, they had not extended their range as far as Patagonia. Marvellously rich and varied as the Santa Cruz fauna was, it did not contain everything that we should expect to find in it; several recent families of undoubtedly indigenous South American origin have left no ancestors in the early Miocene formations. For this, there are several obvious reasons. In part, these gaps in the history are merely due to the accidents of collecting and some of them will almost certainly be filled by future exploration. Other absentees will probably never be found, because the Santa Cruz beds are known only in the very far south, and the Miocene climate of the region, though much milder and more genial than the present one, must have been unsuitable for many tropical animals. Again, the Patagonia of that time appears to have been a country of open plains, with few trees, and hence arboreal forms were rare.

While great numbers of large, flightless birds, some of them of enormous size, were entombed in the volcanic ash and dust which were spread over such wide areas and to such great depths, the extreme scarcity of reptiles is surprising; a few remains of lizards have been found, but no snakes, crocodiles, or tortoises, and we have no information as to the plant-life of the region at that time. The mammals were almost all of small or moderate size; only one or two species were really large.

One very striking and characteristic feature of the Santa Cruz fauna is the great abundance of marsupials which it contained and which resembled more or less those of modern Australia. There were no true Carnivora and their places were taken by a variety of carnivorous marsupials, some of which (_e.g._ _†Prothylacynus_) were as large as wolves and were closely similar to the so-called Tasmanian Wolf (_Thylacynus_). Another genus (_†Borhyæna_) had a short, bullet head, not unlike a small Puma in appearance and, besides, there were many smaller beasts of prey, in size like badgers and minks. Opossums were common and there were many very small herbivorous marsupials, which resembled, though perhaps but superficially, the Australian phalangers. At the present day South America contains no Insectivora, but in the Santa Cruz there was one family (†Necrolestidæ) of this order which bore considerable resemblance to the “golden moles” of South Africa. An extraordinary variety of rodents inhabited Patagonia in Santa Cruz times, all of them belonging to the Hystricomorpha, or porcupine suborder, and all referable to existing South American families. There were none of the northern forms of rodents, neither rats, mice, squirrels, marmots, hares, nor rabbits, but a very numerous assembly of tree-porcupines, cavies, chinchillas, coypus and the like. The genera, though closely allied to existing ones, are all extinct, and the animals were very generally smaller than their modern descendants. A few small monkeys of unmistakably Neotropical type have been found, but like other arboreal and forest-living animals, they are very rare among the fossils.

The Edentata were more abundant and diversified than at any other time in South American history of which the record is preserved. Two of the modern subdivisions of this order have not been certainly identified in the Santa Cruz collections, the arboreal sloths and the anteaters, and though they may be found there at any time, it will only be as stragglers from the warmer forested regions to the north, where these forms had doubtless long been present. Unfortunately, however, nothing is directly known concerning the life of those regions in Miocene times. On the other hand, three groups of edentates, two of them now extinct, were very copiously represented in the Santa Cruz formation, the armadillos, †glyptodonts and †ground-sloths. Of the many armadillos, some quite large, others very small, only a few can be regarded as directly ancestral to those now in existence; the truly ancestral forms were probably then living in the forests of Brazil and northern Argentina, in the same areas as the ancestral tree-sloths and anteaters. In comparison with the giants of the Pliocene and Pleistocene, the Santa Cruz †glyptodonts were all small, the carapace rarely exceeding two feet in length, and, what it is particularly interesting to note, they departed much less widely from the armadillo type than did their gigantic successors. The †ground-sloths were present in actually bewildering variety and they also were very small as compared with the huge animals of the Pleistocene, none of them exceeding the Black Bear in height or length, though proportionally much more massive, and many were no bigger than foxes. They had small heads, long bodies, heavy tails and short, thick legs; their teeth show that they were plant-feeders, but their feet were armed with long, sharp and formidable claws. Among this great host of Santa Cruz †ground-sloths may readily be noted the probable ancestors of the gigantic creatures which were such characteristic elements of the Pliocene and Pleistocene faunas.

There was an extraordinarily rich and varied assemblage of hoofed animals, all utterly different from those of the northern hemisphere and belonging to groups which have never been found outside of South and Central America. Of these groups there were five, which by different writers are variously regarded as orders or suborders, a matter of very secondary importance. Individually, the commonest of the hoofed mammals were the †Toxodonta, which ranged in size from a sheep to a tapir, heavily built and clumsy creatures, with absurdly small, three-toed feet; in some of the species there was a small median horn on the forehead. As with the †glyptodonts and †ground-sloths, the contrast in size between the Santa Cruz ancestors and the Pleistocene descendants was very striking. A very numerous and varied group was that of the †Typotheria, all small animals, some no larger than rabbits, others the size of small foxes. It requires a decided effort to think of these †typotheres as being really hoofed animals at all, as their whole appearance must have been much more like that of rodents, yet their structure clearly demonstrates their near relationship to the †toxodonts. Still a third group of the same series, the †Entelonychia, is of great interest, for, as in the †chalicotheres of the northern hemisphere, the hoofs had been transformed into claws and their five-toed feet had a truly grotesque appearance, not diminished by the long and powerful limbs and relatively small head.

This is the third example of that paradoxical creature, a “hoofed animal” with claws instead of hoofs, and in each of the three instances, there is every reason to believe, the transformation proceeded independently. Among the perissodactyls the †chalicotheres (p. 238) underwent this change; in North America the †Agriochœridæ, a family of artiodactyls, had a very similar history, while in South America the †Entelonychia arose from the same stock as the †toxodonts, with which they were nearly allied. They were among the largest animals of Santa Cruz times and ranged in size from an ox to a rhinoceros.

There was a fourth group, the †Astrapotheria, concerning which our knowledge is tantalizingly incomplete, some species of which were the largest of known Santa Cruz mammals, while others were much smaller. They had short, domed heads, with a considerable proboscis, and were armed with formidable tusks, which were the enlarged canine teeth, the only known instance of large canine tusks among the indigenous South American hoofed animals. The limbs were long and not very massive, the feet short, five-toed and somewhat elephantine in appearance. These bizarre animals would seem to have held a rather isolated position among the South American ungulates, and though they may be traced back to the most ancient mammal-bearing beds of that continent, their relationships are still obscure; much more complete material must be obtained before this problem can be definitely solved. Both the †Astrapotheria and the †Entelonychia died out shortly after the end of the Santa Cruz.

From many points of view the most interesting members of the Santa Cruz fauna were the †Litopterna, an order which also went back to the earliest South American Tertiary. In the Miocene and Pliocene the order was represented by two very distinct families, the †Macrauchenidæ and †Proterotheriidæ, which were superficially very unlike. In the Santa Cruz beds is found a genus (_†Theosodon_) which was apparently the direct ancestor of the Pampean _†Macrauchenia_. The Miocene genus was a much smaller animal and had hardly more than an incipient proboscis, but otherwise was very like its Pampean successor; it was somewhat larger and heavier than a Llama and probably bore some resemblance to that animal in appearance. The long, narrow head, with its prehensile upper lip, must have had an almost reptilian likeness from the numerous uniform and sharp-pointed teeth with which the front of the jaws was supplied; the neck was elongate, the body short and rather slender and the legs long, ending in three nearly equal toes.

The †proterotheres, on the other hand, were almost the only Santa Cruz ungulates which had nothing _outré_ or grotesque about them to the eye of one habituated to the faunas of the northern hemisphere. They were small, graceful animals, very like the Miocene horses of the north in their proportions, though having much shorter necks and shorter, heavier heads. In some genera of this family (_e.g._ _†Diadiaphorus_, _†Proterotherium_) the feet were three-toed and most surprisingly horse-like in shape, but one genus (_†Thoatherium_) was absolutely single-toed, more completely monodactyl than any horse. The horse-likenesses ran all through the skeleton and are so numerous and so striking that several writers have not hesitated to incorporate the †Litopterna with the Perissodactyla, but this I believe to be an error. If the †proterotheres were not perissodactyls, as I am convinced they were not, they afford one of the most remarkable examples of convergent evolution among mammals yet made known.

3. _Oligocene_

_North America._—The John Day formation of eastern Oregon represents the upper Oligocene and has yielded a very extensive series of mammals, though with some obvious gaps that remain to be filled by future work. The land-connection with the Old World which had existed in the lower Oligocene and was restored in the lower, or at latest in the middle, Miocene, was interrupted in John Day times, and so the mammals assumed a purely indigenous character.

No opossums or other marsupials have been found, and nothing is known of the Insectivora. Of the Carnivora, there were but three families, and one of these, the mustelines, was represented but scantily by a few small species. Cats of the †sabre-tooth subfamily were common and one species was quite large, almost equalling the Jaguar in length; but most of the species were small, much smaller than the Pleistocene members of the group. True cats are not definitely known to have been present, but there were two genera (_†Nimravus_ and _†Archælurus_) which have been called the “false †sabre-tooths,” which may prove to be referable to that series. The dogs, on the other hand, were remarkably numerous and diversified, more so than ever before or since; none of them was very large, the largest but little exceeding the Timber Wolf in size, and some were extremely small; but the number of distinct genera and species and the differences among them are quite remarkable. Both long and short-faced forms and early stages of the “†bear-dogs,” and “†hyena-dogs,” and ancestral forms of the wolves and dholes may be distinguished, a truly wonderful assemblage. The rodents also were numerous and varied, including ancient and extinct genera of the beavers, squirrels, mice, pocket-gophers and hares and the earliest distinguishable ancestors of the sewellels (Aplodontiidæ).

The remainder of the known John Day fauna was composed of artiodactyls and perissodactyls. The latter had suffered serious losses as compared with the preceding or White River stage. Up to and through White River times the perissodactyls had held their own in actual diversity, though the rise of the artiodactyls had put an end to the dominant position which they had maintained in the Eocene. With the John Day the actual decline may be said to have begun. The rhinoceroses were represented chiefly by the †diceratheres, with a transverse pair of horns, some species of which were much larger than those of the lower Miocene. Hornless rhinoceroses have not yet been certainly found, though there is every reason to believe that they then existed, as they unquestionably did both before and after. Tapirs occurred but rarely and the horses were individually abundant, though in no great diversity; they were smaller and lighter than the horses of the lower Miocene. Enough has been found to demonstrate the presence of the clawed †chalicotheres, but not to show how they differed from their immediate successors.

In the number of individuals, species, genera and families, the artiodactyls of the John Day much exceeded the perissodactyls. The peccaries were numerous, but smaller and more primitive than those of the succeeding age, as were also the †giant pigs, or †entelodonts, but the latter were very large. The peculiarly North American family of the †oreodonts was very numerously represented, and one genus (_†Promerycochœrus_), comprising animals not unlike the Wild Boar in size and shape, was the probable beginning of the series of proboscis-bearing †oreodonts, which led to such grotesque forms in the middle and upper Miocene. A family closely allied to the †oreodonts, and by many writers included in the latter, is the very remarkable group of the †Agriochœridæ, which was distinguished by the long, stout and cat-like tail and by the possession of claws instead of hoofs. The family is not known to have existed later than the John Day and no trace of it has been found in the succeeding formations. The camels seem to be all comprised in a single genus (_†Protomeryx_) which was the same as that found in the lower Miocene. A very small and dainty little creature (_†Hypertragulus_) belonged to another family, the relationships of which are not clear.

To the middle and lower Oligocene is referred the great White River formation of South Dakota, Nebraska, Wyoming, etc., which is divisible into three clearly marked substages. The White River contains the best-known fauna of all of the North American Tertiaries, for collecting in these beds has been carried on for more than sixty years, and a greater number of complete and nearly complete skeletons has been secured than from any of the other formations. It is plainly evident that a land-connection existed with the Old World, which was interrupted in the John Day, as is shown by the intermigration of characteristic forms; but some barrier, presumably climatic, prevented any complete interchange of mammals, and very many genera and even families remained confined to one continent or the other.

The aspect of the White River fauna changes in accordance with the direction from which it is approached. If one comes to the study of it from the Eocene, it displays a very modern aspect, given by the almost complete disappearance of the archaic groups of mammals and by the great multiplication of genera and species belonging to the progressive orders. These genera, it is true, are all extinct, but many of them stood in an ancestral relationship to modern forms. On the other hand, if approached from the Miocene side, the White River mammals seem to be very ancient and primitive and very different from anything that now lives. We speak of horses and rhinoceroses, dogs and cats, in this fauna, but those terms can be employed only in a very wide and elastic sense to designate animals more or less distantly allied to those of the present day.

Several species of opossums, some of them very small, were the only marsupials in North America then, as they are now. There was quite a variety of Insectivora; some were survivals of a family that was abundant in the Eocene, others, like the hedgehogs, moles and shrews, were probably immigrants. Here we find the last of a group (order or suborder) of ancient and primitive flesh-eaters, the †Creodonta, that had played a great rôle in the Eocene and Paleocene of North America and Europe. In White River times but a single family (†Hyænodontidæ), with two genera, remained of the Eocene host. One of these genera (_†Hemipsalodon_), a very large beast of prey, which was almost identical with the Old World genus _†Pterodon_, was confined to the lower substage of the White River beds in the Northwest Territory of Canada; the other, _†Hyænodon_, which was also an Old World form, was represented abundantly in the United States by many species. In size, these species ranged from a small fox to a large wolf, but they all had disproportionately large heads, and small, weak feet, with blunt claws, so that they must have been very curious-looking creatures and were probably carrion-feeders rather than active catchers of prey. The White River members of the family were migrants from the eastern hemisphere, for, though small and primitive representatives of it occurred in the North American Eocene, as well as in the corresponding formations of Europe, the family appears to have died out in America and to have been renewed by the Oligocene migration.

Coincident with this decline of the †creodonts and, no doubt, causally connected with it, was the rise of the true Carnivora, which for the first time were numerous and were divisible into three distinct families. Small and primitive representatives of the wolves (_†Daphœnus_) and possibly also of the foxes (_†Cynodictis_) were quite common, and there were a few species of the musteline family, evidently immigrants and the most ancient yet found in America. There were several species of the †sabre-tooth cats (_†Dinictis_ and _†Hoplophoneus_) all of which, except in the uppermost substage, were quite small, few of them exceeding the Canada Lynx in size. A much larger animal (_†Eusmilus_, also European) appeared in the latter part of the stage. None of the true cats, or feline subfamily, has been obtained. Nothing is yet known of the time and place of origin of the †sabre-tooth series, for they appeared at approximately the same date in Europe and America, and in neither continent have any possible ancestors been found in preceding formations. The problem is like that of the Proboscidea (_see_ p. 234), but Egypt has given no help in the case of the †sabre-tooths, and, by a process of elimination, we reach the conclusion that these strange creatures probably arose somewhere in Asia and sent out migrants eastward and westward.

The Rodentia were fairly abundant and present a strange mixture of ancient and comparatively modern types. One very common genus (_†Ischyromys_), which was the last remnant of a family almost limited to the North American Eocene, was associated with the earliest American mice, arboreal and ground squirrels, beavers and rabbits; some, if not all, of these were immigrants.

The hoofed mammals were present in fairly bewildering variety, but were restricted to the two orders of the Perissodactyla and Artiodactyla. The Perissodactyla, while they no longer had the relatively dominant position which they held in the middle Eocene (_see_ p. 270), had suffered no actual loss; and no less than seven families of them, or six by another scheme of classification, had members in the North America of White River times, a very notable difference from the present order of things, when there are but three families in the entire world, none of which enters North America. The Eocene family of the †titanotheres became extinct at the end of the lower substage of the White River, but in that substage there was a marvellous abundance of these huge beasts, some of which were of almost elephantine stature and bulk. The pair of great bony, horn-like protuberances on the nose varied much in size and form in the different species, short to very long, triangular, cylindrical, flattened and shovel-shaped, and gave these ungainly creatures somewhat the appearance of strange and very large rhinoceroses. The †titanotheres were a typically North American family, but sent migrants to the Old World, at least two species reaching southeastern Europe. Rhinoceroses too were extremely numerous and diversified throughout the stage and are very plainly divisible into three strongly contrasted series, which are sometimes regarded as three subdivisions of the same family and sometimes put into two separate families. One of these series, the †hyracodonts (_†Hyracodon_), was composed of small, long-necked and long-legged, slender and lightly built, cursorial animals, but with short, heavy heads, which gave them a somewhat clumsy look; having neither horns nor tusks, they were entirely defenceless and depended for their safety upon speed alone. The second series, or †amynodonts (_†Metamynodon_), formed the very antithesis of the first,—large, heavy, short-necked, and short-legged and probably amphibious in manner of life, they were armed with formidable tusks; and their skulls were so curiously modified as to bear a distinct resemblance to the skull of a huge carnivore. The †amynodonts migrated to the Old World and occur in the Oligocene of France, but the †hyracodonts would seem never to have left North America. The third series, that of the true rhinoceroses, comprised several genera at different levels in the White River beds (_†Trigonias_, _†Cænopus_, etc.); they were of uncertain origin and it has not yet been determined whether they were immigrants or of native stock. Many species have been found, varying much in size, up to that of a modern tapir, and not unlike one in proportions, for they were of lighter build and had relatively longer legs than any existing rhinoceros. The species of the lower and middle substages were all hornless, but in the uppermost substage we find skulls with a pair of nasal horns in an incipient stage of development. This was the beginning of the †paired-horned rhinoceroses (_†Diceratherium_) which so flourished in the John Day and the lower Miocene.

Of the horses there was no great variety and all the species so far discovered are included in a single genus (_†Mesohippus_), though there was a decided increment in the size of the successive species from the earlier to the later portion of the stage. Looked at superficially, it seems absurd to call these little creatures “horses” at all and the term can be justified only as implying that they were ancestral members of the family. The largest of the White River species hardly exceeded a sheep in size and all of them had comparatively short necks, long and slender legs and three-toed feet. The low-crowned grinding teeth show that they were browsers, not grazers. The abundant Eocene family of the †Lophiodontidæ made its last appearance in the White River, where it was scantily represented by slender, long-legged animals (_†Colodon_), with feet singularly like those of the contemporary horses, except that there were four toes in the front foot. Tapirs (_†Protapirus_) were very much less common than rhinoceroses or horses and were hardly half as large as the existing species of the family and of relatively far more slender form; the development of the proboscis had already begun. Lastly, the presence of the clawed †chalicotheres has been reported from the lower Oligocene of Canada, but the material is too fragmentary for generic reference.

Though the number of artiodactyl families yet identified among the White River fossils is no larger than that of the perissodactyl families, the artiodactyls greatly preponderated in individual abundance. The peccaries, which were fairly common, resembled those of the John Day, but were considerably smaller. Of the camels, there were two series, one of which (_†Eotylopus_), lately described by Dr. Matthew, is of yet unknown significance, while the other (_†Poëbrotherium_) was apparently the ancestor common to all the subsequent phyla of camels and llamas. This extremely interesting genus had species which ranged in size from a gazelle to a sheep, had two toes in each foot, a moderately elongate neck and teeth which were beginning to assume the high-crowned character. From this it may be inferred that those animals were, partly at least, of grazing habit, which was rare among White River ungulates, most of which fed upon leaves and soft and succulent plants. An extinct family, the †Hypertragulidæ, were a greatly diversified group of dainty little creatures, one of which (_†Hypisodus_) was no larger than a rabbit and had high-crowned teeth. The other genera (_†Leptomeryx_, _†Hypertragulus_) must have resembled in form and proportions the tiny little chevrotains or “mouse-deer” of the East Indian islands. Late in the age arose a larger form of this family, nearly equalling the Musk-Deer in size, the extraordinary genus _†Protoceras_, which was, especially the males, a grotesque object. The males had a pair of upper canine tusks and two pairs of prominent long protuberances on the skull. This, or some similar form, must have been the ancestor of the still more bizarre _†Syndyoceras_ of the lower Miocene.

The †oreodonts were by far the commonest of White River mammals, and evidently they roamed the woods and plains in great herds. There were several species, larger and smaller, of the abundant genus (_†Merycoidodon_) but the largest did not surpass a modern peccary in size and was somewhat like that animal in appearance, but had a shorter head and much longer tail. In the upper substage appeared a very peculiar genus of this family (_†Leptauchenia_), animals with short, deep, almost monkey-like heads, and presumably aquatic in habits. The _†agriochœrids_ were very much less common; they may be described roughly as †oreodonts with very long, cat-like tails and clawed feet.

All of the foregoing artiodactyl families were exclusively North American in Oligocene distribution; even the camels did not reach Asia till the Pliocene, and the other families never invaded the Old World at all. There were, however, two additional families, which also occurred in the eastern hemisphere, whence one of them, and possibly the other, was derived. The unquestionably Old World family, that of the †anthracotheres, was represented in the White River by two genera (_†Bothriodon_ and _†Anthracotherium_), which were short-legged, long-snouted, swine-like animals, which have no near relations in the modern world. The other family, the †giant pigs, which we have already met with in the lower Miocene and upper Oligocene, is of doubtful origin, and nothing has yet been found in the preceding formations of either North America or Europe which can be regarded as ancestral to them. The White River genus (_†Archæotherium_) was very like the John Day and Arikaree genera, but most of the species were much smaller and some were not so large as a domestic pig. In the uppermost beds, however, are found huge species, which rivalled those of the subsequent formations. That these strange animals were rooters and diggers and therefore pig-like in habits is indicated by the manner in which the teeth are worn.

_South America._—The older continental Tertiary formations of South America cannot be correlated with those of North America or Europe, because they have nothing in common. Difficult as it is to give a correct and adequate conception of the Tertiary mammalian life of the northern hemisphere to one who has not made a study of it, it is far more difficult in the case of South America. The stock of adjectives, such as “peculiar,” “bizarre,” “grotesque” and the like, already overworked in dealing with northern forms, is quite hopelessly inadequate where everything is strange. In addition to this, we are seriously handicapped in treating of the Oligocene and Eocene of South America by very incomplete knowledge. Many fossils have been collected and named, but the great majority of these are known only from teeth; a few skulls and limb-bones have been described, but no skeletons, and therefore much is very uncertain regarding these faunas.

The Deseado formation (Pyrotherium Beds) has been variously referred by different writers from the upper Cretaceous to the lower Miocene, but its most probable correlation is with the Oligocene. Though most of the mammalian groups are the same as those of the Santa Cruz, the proportions of the various orders in the two faunas are very different, but, to some extent, the difference is probably illusory and due to the conditions of fossilization, for, as a rule, the small mammals are much less frequent and well preserved in the older beds. As in the Santa Cruz, the marsupials were the only predaceous mammals, and some of them attained gigantic size; but no such variety of these beasts of prey has been found in these beds as occurred in the middle Miocene. In addition, there were numerous small herbivorous marsupials. One of the most striking differences from the Santa Cruz fauna was in the very much smaller number of Edentata, which, instead of being extremely common, are quite rare among the fossils. No doubt there was a real and substantial difference in this respect, but it was probably not so great as it seems, and the same three suborders are found in both formations. One of the few †ground-sloths that have been obtained was very large (_†Octodontherium crassidens_), a much larger animal than any species of the suborder that is known from the Santa Cruz. The †glyptodonts were also rare, and only two genera and species have been described from very scanty remains. Armadillos, on the other hand, were much more common, and no less than eleven genera have been named, three of which occurred also in the Santa Cruz. Among these was the remarkable genus _†Peltephilus_, in which the anterior two pairs of plates of the head shield were modified into horn-like spines.

Equally striking was the remarkable diminution of the Rodentia, as compared with those of the Santa Cruz, though, of course, this is an inaccurate mode of stating the truth, occasioned by the fact that we are following the history in reverse order. It would be preferable to say that the rodents underwent a remarkable expansion in the Santa Cruz. These rodents of the Deseado stage are the most ancient yet discovered in South America and represent only two families, both belonging to the Hystricomorpha, or porcupine group. If, as Dr. Schlosser and other European palæontologists maintain, the Hystricomorpha were all derived from a family of the European Eocene, this would necessitate a land-connection between South America and the Old World independent of North America, for the latter continent had no hystricomorph rodents until the connection between the two Americas was established.

The great bulk of the Deseado fauna is made up, so far as individual abundance is concerned, of hoofed animals belonging to the typically South American groups. The †Toxodonta were represented partly by genera which were the direct ancestors of the common Santa Cruz genera (_†Pronesodon_, _†Proadinotherium_), and, more numerously, by a very peculiar family, the †Notohippidæ, which had highly complex, cement-covered grinding teeth. Still a third family of this suborder, the †Leontiniidæ, was highly characteristic of the Deseado fauna and is not known from the Santa Cruz. These were large animals, with a small horn on the tip of the nose and low-crowned, comparatively simple grinding teeth. Even more abundant were the †Typotheria, small forms which were ancestral to the Santa Cruz genera, larger ones which died out without leaving successors and one quite large animal (_†Eutrachytherus_) which seems to have been the ancestor of the Pliocene and Pleistocene _†Typotherium_. This series is not known to have been represented in the Santa Cruz and may have withdrawn from Patagonia at the end of the Deseado stage.

The †Entelonychia, those strange toxodont-like animals with claws instead of hoofs, were much more numerous and varied than they were afterward in the Santa Cruz, when they were on the verge of extinction, and included both very small and very large species. The †Pyrotheria, a suborder which is not met with in the Santa Cruz or later formations, likewise included some very large forms. The typical genus, _†Pyrotherium_, included large, relatively short-legged and very massive animals; the upper incisors formed two pairs of short, downwardly directed tusks, and in the lower jaw was a single pair of horizontally directed tusks; the grinding teeth were low-crowned and had each two simple, transverse crests. These grinding teeth and the lower tusks so resemble those of the ancestral Proboscidea in the Oligocene of Egypt, that the †pyrotheres have actually been regarded as the beginnings of the †mastodons and elephants, but this is undoubtedly an error. The †Astrapotheria, another group which became extinct at or soon after the end of the Santa Cruz, were relatively abundant in the Deseado and counted some very large species. Finally, the †Litopterna were represented by the same two families as continued through the Pliocene and one of them far into the Pleistocene. The horse-like †proterotheres were present, but not enough of them has been obtained to show whether or not they were in a notably less advanced stage of development than those of the Santa Cruz. The †macrauchenids were quite similar to those of the latter formation, though considerably smaller. In addition, there were a few genera, survivals from earlier times, which were not referable to either of these families.

The large number of genera, especially among the †toxodonts and †typotheres, which had high-crowned, cement-covered teeth, may be taken as an indication that grazing habits had already begun to be prevalent.

Of this wonderful assemblage of hoofed animals, divisible into six separate groups, whether of ordinal or subordinal rank, not a trace remains to-day. Not only are all the species, genera and families extinct, but the suborders and orders also. Further, this was a very strictly autochthonous fauna, so far as the hoofed animals were concerned, and no member of any of the six groups has ever been found outside of the Neotropical region.

4. _Eocene_

_North America._—In the western interior of North America the Oligocene followed so gradually upon the Eocene, that there is great difficulty in demarcating them and much difference of opinion and practice obtains as to where the boundary line should be drawn. Not to depart too widely from the scheme used by Professor Osborn, the Uinta stage is here treated as uppermost Eocene, though this is a debatable procedure. For several reasons, the extraordinarily interesting and significant Uinta fauna is far less completely known than that of the preceding Bridger and succeeding White River stages. For one thing, it has been much less thoroughly explored, and it may be confidently expected that future exploration will greatly enlarge our knowledge.

The smaller mammals of the Uinta are particularly ill-known. No Insectivora have yet been found, though this gap will assuredly be filled; rodents are scanty in the collections and include only two families, one the †ischyromyids, which were still common in the White River, the other of doubtful position, but not improbably to be considered as the beginning of the pocket-gophers (Geomyidæ). The archaic flesh-eaters, or †Creodonta, were represented by two families, one comprising smaller animals with somewhat cat-like, shearing teeth (†Oxyænidæ), the other, very large beasts with crushing teeth (†Mesonychidæ), neither of which continued into the White River. As compared with the middle and lower Eocene, the †creodonts had greatly diminished and, to replace them, the true Carnivora were beginning to come in. As yet, however, only small and very primitive dog-like forms are known and no trace of †sabre-tooths or mustelines has been found. Indeed, it is very doubtful whether members of these families ever will be found in the Uinta, for their presence in the succeeding White River was probably due to immigration.