Handbook to the Primates/Extinct Anthropoidea

[ 209 ]


As we have seen above (vol. i., p. 110) the earliest Lemuroids appeared in the Lower Eocene division of the Tertiary period in the New World, and in the Old World in its upper strata; they continued during the whole of the Eocene in the Western Hemisphere, and are last seen in the Lower Miocene of North America.

Fossil Apes, on the other hand, appear first in South America, in the Santa Cruz beds of Patagonia, in strata of Upper Eocene or Oligocene age. In the Old World they come on the scene only during the tropical ages of the Miocene epoch. When the middle and upper strata of the latter period were being deposited in Europe, Anthropoid Apes ranged from the Mediterranean shores to further north than the present northern limit of the Old World Apes.

In the Pliocene age Anthropoidea were living in Southern Asia, around where the Sivalik hills now stand, and in Southern Europe, as at Pikermi and Samos, being represented almost entirely by species of still existing genera, and one living species—the Orang. Chimpanzees had already then become differentiated, and perhaps Man had even appeared, though the evidence is not sufficiently conclusive.

In the Pleistocene, remains of many still living species have been brought to light both in the New and the Old Worlds, and unmistakable osseous remains, as well as abundant evidences [ 210 ]of his handiwork, prove the existence of Man at that remote epoch.

FAMILY HAPALIDÆ (Vol. I., p. 129).

GENUS HAPALE (op. cit., p. 131).

Of this genus abundant remains of two species have been found in many of the Brazilian caverns of Pleistocene or recent age. These have been referred to two species: Hapale grandis (Lund), and the still-living H. jacchus (Linn.; cf. Vol. I., p. 132).

FAMILY CEBIDÆ (Vol. I., p. 150).


Protopithecus, Lund, Ann. Sc. Nat. (2), xi., p. 230 (1839); Zittel, Handb. Palæont., iv., p. 705 (1893).

This genus is founded on a very large leg-bone from the Pleistocene bone-caves of Brazil. The species has been described as Protopithecus brasiliensis, Lund.

GENUS CALLITHRIX (Vol. I., p. 158).

Two species have been described from the Pleistocene bone-caves of Brazil: Callithrix chlorocnomys, Lund, and C. primæva, Lund (= C. antiqua, Lund).

GENUS ALOUATTA (Vol. I., p. 192).

Remains of one species, Alouatta ursina (p. 149), has been discovered in the Pleistocene bone-caves of Brazil.

GENUS CEBUS (Vol. I., p. 204).

The Pleistocene bone-caverns of Brazil have preserved three species: one extinct, Cebus macrognathus, Lund, and two still living, C. fatuellus, Linn., and C. cirrifer, Geoffr.

[ 211 ]


Homunculus, Ameghino, Rev. Argent. Hist. Nat., i., pp. 290, 384 (1891).

Ecphantodon, Mercenat, Rev. Mus. La Plata, ii., p. 74, pl. ii.; Zittel, Handb. Palæont., iv., p. 704 (1893).

The dental formula of this genus is I 2/2, C 1/1, P 3/3, M 3/3. The diastema, or break, in the dental series is very small; the incisors are chisel-shaped, the outer pair smaller than the inner pair. The canines, which have a small basal cusp behind, are only slightly prominent; the pre-molars have one root, and one low outer cusp, and two higher inner cusps. The molars are quadrangular, with two pairs of cusps, each united obliquely by a ridge; the anterior molar is smaller than the two hinder. The arm-bone (humerus) has an ent-epi-condylar foramen. (Zittel.) The front surface of the line of union of the two halves of the lower jaw is vertical. The terminal joints of the digits have nails. The thumb and the great-toe are opposable. Homunculus patagonicus, Ameghino (= Ecphantodon ceboides, Mercenat), the only known species, is found in the Upper Eocene or Oligocene of Santa Cruz, Patagonia.


Anthropops, Ameghino, Rev. Arg. Nat. Hist., i., p. 387 (1891); Zittel, Handb. Palæont., iv., p. 704 (1893).

This genus is known from only a fragment of a lower jaw containing four small incisors, two strong canines, and anterior and median pre-molars, both one-rooted. One species, Anthropops perfectus, Ameghino, from the older Tertiary (Upper Eocene or Oligocene) beds of Santa Cruz, Patagonia, is known.

Two genera, Homocentrus (H. argentinus, Amegh.) and [ 212 ]Eudiastus (E. lingulatus, Amegh.), described by Ameghino, from the Santa Cruz beds in Patagonia, are not yet sufficiently characterised.


GENUS PAPIO (suprà, p. 253).

Several species of this still living genus have been recovered from strata of the Tertiary epoch: Papio sub-himalayanus (Meyer), from the Sivalik hills, of Lower Pliocene age; P. falconeri (Lydekker), from the Pleistocene bone-caves of Madras, India, and in the superficial deposits of Algeria, North Africa; and P. atlanticus (Thomas).

The Sivalik species was closely related to the existing North-African Baboons.


Oreopithecus, Gervais, C. R., p. 1223, lxxiv. (1872); Ristori, Boll. Com. Geol. (3), i., pp. 178, 226, pls. vii., viii. (1890); Zittel, Handb. Palæont, iv., p. 705 (1893).

The characters which distinguish this genus are the incisors, which are chisel-shaped above and scoop-shaped below; the large upper and lower canine teeth; the upper pre-molars, which approach in shape to the molars, with the outer cusps higher than the inner, and the inner one strong; the upper molars with two pairs of opposite conical cusps, separated by a longitudinal furrow, and with a strong cingulum; the posterior upper molar smaller than the median; the lower molars smaller than the upper, with two pairs of cusps, and a fifth on their hind border, which in the hindmost tooth is developed into a strong talon. The face is short, and the chin rounded. Oreopithecus bambolii, Gervais, is the best known species, and was obtained from the Mid-Miocene lignites of Monte Bamboli, [ 213 ]Casteani, and Monte Massi, in Tuscany. It has been placed by some Palæontologists among the Simiidæ, and by others in the Cercopithecidæ. According to Ristori, the under jaw shows its alliance with Papio and Cercopithecus; while the upper jaw more resembles the Anthropoid Apes. It is the largest known fossil Ape, and is excelled in strength only by Dryopithecus, Zittel.

GENUS MACACUS (suprà, p. 1).

Species belonging to this still living genus, occurred in Asia and in Europe in the age—the Pliocene—which immediately preceded the Great Ice age, as well as in the Pleistocene epoch itself. Macacus sivalensis is the oldest fossil of the genus, and was described by Mr. Lydekker from the Sivalik beds of the Punjaub. M. priscus is known from the Pliocene of Montpellier, in France; M. florentinus, Cocchi (the same as Aulaxinuus florentinus of Cocchi, and M. ausonianus of Forsyth Major), from the Upper Pliocene beds in the valley of the Arno. M. suevicus (Hedinger), which has been described from a well-preserved palate-bone, having all the molar, and two of the pre-molar teeth present, was found at Heppenlochs, in Würtemberg. M. trarensis (Pomel) is found in Algeria, in beds of the Ice age; while, in holes on the rock of Gibraltar, remains of the same species as is now living there—M. inuus—were discovered by Mr. Calderon in 1879. From another crevasse at Monstaines, in the Haute Garonne, M. Harlé obtained a fragment of a lower jaw of a species of Macacus, associated with the bones of Mammals of the Ice age. (Zittel.) Of the same antiquity is a jaw found, according to Mr. Lydekker, near the village of Grays, in Essex, a fact which indicates a very great difference in the climate of that part of England from that of the present day.

[ 214 ]


Dolichopithecus, Depéret, Mem. Soc. Geol. Fr., Palæont., i., p. 11 (1890); Zittel, Handb. Palæont., iv., p. 707 (1893).

Allied to Semnopithecus, but having the muzzle longer and the limbs shorter and stouter. The genus has been based on three crania, several teeth, and a number of the bones of the skeleton, belonging to the species Dolichopithecus ruscinensis, Depéret, from the Pliocene strata of Perpignan, in France. (Zittel.)


Mesopithecus, Wagner, Abh. K. Bayer, Ak. (1) iii., p. 154; vii., abth., ii., p. 9; Zittel, Handb. Palæont., iv., p. 706 (1893).

This genus is based on a skull and teeth, which indicate an alliance with Semnopithecus, while the skeleton more resembles that of Macacus Inuus (the Barbary Ape). The male had much longer and more powerful canines than the female. Mesopithecus pentelici, Wagner, the typical species, was founded on a fragment originally brought by a soldier in 1838 from Pikermi to Munich. Since then the whole skeleton has been recovered, and this is now one of the best-known species of the fossil Anthropoidea. It lived in Pliocene times, apparently in troops in the forests of the Pikermi plains, which at that date extended far into what is now the Mediterranean Sea. Remains of the same species have been discovered near Baltavar, in Hungary.

GENUS COLOBUS (suprà, p. 85).

In the Mid-Miocene forests of Europe this genus was represented by a species described by Professor Fraas as Colobus grandævus, from Steinheim, in Würtemburg.

[ 215 ]

GENUS SEMNOPITHECUS (suprà, p. 100).

Among the forests in which bamboos, liquidambars, tulip-trees, magnolias, laurels, and pomegranates flourished in Upper Pliocene days, in the middle of Europe, there lived troops of Langurs, closely allied to those of our own time. Semnopithecus monspessulanus, Gervais, has been recovered from the strata of that age, at Montpellier, and near Casino in Tuscany. S. palæindicus (Lydekker) inhabited the forests in the region where the Sivalik hills now rise at the foot of the Himalayas, while S. entellus roamed over that region in the Pleistocene age, as its actual descendants do to-day.

FAMILY SIMIIDÆ (suprà, p. 143).


Pliopithecus, Gervais, C. R., xliii., p. 221 (1856); id., Zool. et Pal. Franc., p. 8 (1859); Forsyth Major, Atti. Soc. Ital. Sc. Nat., xv., p. 82 (1872); Zittel, Handb. Palæont., p. 708 (1893).

Protopithecus, Ed. Lartet (nec Lund), Ann. Dep. Gers., 1851, p. 11.

This genus is very nearly allied to Hylobates, but differs from it in the form and proportions of its teeth. The genus is based on a lower jaw found in the Mid-Miocene of Central Europe. The incisors are small and long; the canines strong and but little taller than the incisors; the pre-molars are low, the anterior having one cusp, and the next two cusps; the molars have two pairs of opposite short, thick, conical cusps, with an additional one on the hind border, which enlarges into a talon in the hindmost of the set. The type species, Pliopithecus antiquus, which very closely resembles the Gibbons, lived in the luxuriant forests of Sansan (Gers), and a variety of [ 216 ]it, described as P. chantrei, Depéret, inhabited the woods round Mont Ceindre. Remains of the same animals have been obtained in the Brown-coal beds of Elgg, in Switzerland and Göriach, in Steyermark.

GENUS HYLOBATES (suprà, p. 148).

True Gibbons, indistinguishable from those now living in the island, have been found in the caves of Borneo.

A finely preserved limb-bone, from the Eppelsheim beds of the Pliocene age, has also been ascribed to a species of this genus.


Dryopithecus, Lartet, C. R., xliii., p. 221 (1856); id., Mem. Soc. Geol., Palæon., i., p. 1, pl. 1 (1890); Gaudrey, C. R. Cx., p. 373 (1890); Zittel, Handb. Palæont., iv., p. 709 (1893).

This genus is based on remains from the Mid-Miocene of St. Gaudens (Haute Garonne), which indicate the former existence of an Ape more Man-like than any other. In size it approached the dimensions of the Chimpanzee; the incisors are smaller—an elevated character—and shorter than those in the Gorilla or the Chimpanzee. The canines are, as in the Gorilla, thick, sharp behind, and taller than the cheek-teeth; the anterior pre-molar is large, as in the Gorilla, has one root, and a strong cingulum on the inner side; the posterior pre-molar is longer than broad, is two-cusped, and has a flattened talon. The molar teeth have two pairs of opposite cusps, and a fifth on the hind border, which develops, on the hindmost tooth, into a two-cusped talon. The line of union of the lower jaw is high, [ 217 ]projects obliquely forward, and is longer and narrower than in Man. The late appearance of the last molar in the upper jaw was supposed to be a character which was alone common to Dryopithecus and Man; but Dr. Forsyth Major has observed that in Macacus the same late in-coming of the "wisdom tooth" occurs. The type species, Dryopithecus fontani, Lartet, which lived in the Mid-Miocene forests of St. Gaudens, though the most Man-like of all the Tertiary Apes, was nevertheless further distant from Man than the Chimpanzees (Anthropopithecus). The form of the symphysis of its lower jaw indicates that its snout was considerably lengthened. Certain molar teeth found in the Bohnerz strata from Melchingen and Salmendingen, in Würtemberg, and at one time considered to be human, have now been ascribed to D. fontani.

GENUS SIMIA (suprà, p. 170).

To this genus has been referred a molar tooth found in the Pliocene Strata of the Sivalik hills in India. It is considered to belong to an Orang-Utan, Simia satyrus.


A fragmentary jaw, also from the Pliocene beds in the Sivalik hills, has been described as Anthropopithecus sivalensis by Lydekker, who at first placed it in a new genus, Palæopithecus, but has more recently determined it to belong really to this now exclusively African genus. The relative smallness of the premolars distinguish it from the Orang. Should this determination be confirmed, the presence of a true Chimpanzee in Asia will be a fact of the highest interest in the geographical distribution of the Simiidæ.

[ 218 ]

FAMILY HOMINIDÆ (suprà, p. 203).

GENUS HOMO (suprà, p. 203).

Although, as has been stated above, the Primates, represented by lowly Lemuroids evincing relationship with the ancestors of the hoofed animals (Ungulata), first appeared in Eocene times, it would be a hopeless quest, as Professor Boyd-Dawkins points out, to seek for a highly specialised Man in a fauna where no living genus of Mammals was present.

The earliest appearance of Man on the globe has been considered by Dr. Hamy and M. de Mortillet to be in France in the middle of the Miocene age. They base their belief on flint fragments supposed to be artificially made, and on a cut upon the bone of an extinct Manatee considered to be of human handiwork. The evidence is, however, doubtful and unsatisfactory. In this age appeared such Anthropoids as Pliopithecus and the highly-developed Dryopithecus (p. 216), when the climate was tropical in mid-Europe, and warm and genial even within 8° 15′ of the North Pole. Professor Boyd-Dawkins believes that notwithstanding the favourable climate and the existence of so highly-developed an Ape as Dryopithecus, "were any Man-like animal living in the Miocene age, he might reasonably be expected to be not Man, but intermediate between Man and something else."

The Pliocene, i.e., that portion of the Tertiary period in which the genera of mammals are mostly the same as those now living—only one species is known to be identical,—is the next horizon in which human remains have been asserted to have been found. The evidence is based on a skull found in a railway cutting in France after a landslip, and on a supposed artificially incised bone; but both these data require confirmation. Senhor Ribeiro has, however, obtained in Portugal implements [ 219 ]said to be of undoubted human manufacture in strata of this age, 1,200 feet below the surface; and it has been claimed by Professor Whitney that, in California, a skull, as well as a mortar and pestle, have been recovered from Pliocene beds. The latter evidence has also been called in question.

The discovery at Crayford and in Kent's Hole in England, and in the Grotte d'Église in France, of flint implements of human manufacture, demonstrates without doubt that Man was living in Europe in the Pleistocene age—at which time most of the species of Mammals were identical with those now living—before the climate (which had been cooling since the Miocene) had become so cold as to cause the Arctic Mammals to swarm down in front of the approaching glaciation of the Northern Hemisphere. At that epoch the River-drift Men, as they are called, would have had to contend with Wolves, Bears, and Lions; while Elephants and Rhinoceroses, Horses, Oxen, and Bison roamed wild around them. The implements of this "long-headed" race were stones, conveniently picked up and rough-hewn into rude choppers and scrapers, pointed borers, and cutting chips. There is evidence that their makers ranged across a more extended Europe than now, into Africa and continental India. After the River-drift Men, who disappeared with the Ice age, there came on the scene a race known as the Palæolithic "Cave Men." Associated with their bones there have been found, in numerous caverns, remains of the Reindeer (Cervus tarandus), the Woolly Rhinoceros (R. tichorhinus), and the Mammoth (Elephas primigenius). They were an artistic people, who have left drawings of extraordinary fidelity of the animals with which they were familiar, scratched on bones and horns of the animals themselves. Their implements were better chipped and shaped than were those of the [ 220 ]River-drift Men. They appear to have been ignorant of the potter's art; but they clothed themselves in skins, wore teeth-ornaments, and hunted the Reindeer and other animals—they were men, as Sir A. Geikie remarks, who must have had much similarity with the Esquimo—an identification, however, which has lately been strongly contested. Many fragments of their skeletons have been found in caverns in various parts of Europe: a lower jaw and an ulna at Naulette, a skull at Cro-Magnon, a lower jaw in the Grotte des Fées at Arcy-sur-Cure (Yonne), another from the rock shelter of La Madelaine in the Dordogne; portions of skulls from Neanderthal, Cannstatt, and Gibraltar, and as far north as Derbyshire, in England. The remains are, unfortunately, all very fragmentary, and afford little more information as to the physical characters of the Palæolithic races, than that they were "long-headed." In 1886, however, in the Grotto of Spy, in the Belgian Province of Namur, were discovered two nearly complete skeletons, which showed that the Neanderthal skull, the lower jaw from Naulette, and the skulls from Cannstatt and Gibraltar all belonged to the same race. This race, which was widely spread over Europe in the Palæolithic age, presents more Simian characters than any yet unearthed. MM. Lohest and Fraipont, of Liege, who discovered and described the remains from Spy, have given in detail the following Simian characteristics which they present: The superciliary crests are far greater, and the forehead more retreating, than in any other known race—characters which closely resemble those in female and young male Orangs and Chimpanzees; and the occipital region of the skull shows a transverse crest as in some African tribes and in the above-named Anthropoid Apes. The lower jaw presents little or none of that markedly [ 221 ]human character—the chin; and the slope of the interior (or posterior) surface of its symphysis is intermediate between that of Man and the higher Apes. The bones of the fore-arm (the ulna and radius) are curved so as to produce a space between them, wider than in any human subject, and resembling what is seen in Apes. The thigh-bone (femur) is so shaped and articulated to the leg-bone (tibia) "that in order to maintain equilibrium the head and body must have been thrown forward." This relation of the femur and tibia is found in the Apes, and it is highly probable that the Man of Spy presented a somewhat similar figure when walking; that is to say, the knees were bent and the body thrown forward. The crowns of the molar teeth of this race have, as in the lowest races of Man, four cusps, but with distinct and divergent roots, as among the Chimpanzees, but they increase in size from in front to behind, as they do in Apes. "The other and much more numerous characters of this long-headed skull, of the trunk and of the limbs, seem to be all human." (Fraipont.) "Under whatever aspect we view this [the Neanderthal] cranium ... whether we regard its vertical depression, the enormous thickness of its supra-ciliary ridges, its sloping occiput, or its long and straight squamosal suture—we meet with Ape-like characters, stamping it as the most pithecoid of human crania yet discovered." The cranial capacity being, however, about seventy-five cubic inches, "so large a mass of brain as this would alone suggest that the pithecoid tendencies indicated by the skull did not extend deep into the organisation.... In no sense, then, can the Neanderthal bones be regarded as the remains of a human being intermediate between Man and Apes." (Huxley, 1867.) "The distance which separates the Man of Spy from the [ 222 ]modern Anthropoid Ape is undoubtedly enormous; between the Man of Spy and the Dryopithecus it is a little less. But we must be permitted to point out that if the Man of the later Quaternary age is the stock whence existing races have sprung, he has travelled a great way. From the data now obtained, it is permissible to believe that we shall be able to pursue the ancestral type of Man and the Anthropoid Apes still further, perhaps as far as the Eocene, and even beyond." (Fraipont.) As these fossil human remains are now admitted to be of the Palæolithic age of the Pleistocene period, they give some idea of "the rate of evolution of the human species, and indicate that it has not taken place at a much faster or slower pace than that of other Mammalia. And if that is so, we are warranted in the supposition that the genus Homo, if not the species which the courtesy or the irony of naturalists has dubbed sapiens, was represented in Pliocene or even Miocene times.... There is no reason to suppose that the genus Homo was confined to Europe in the Pleistocene age; it is much more probable that this, like other Mammalian genera of that period, was spread over a large extent of the surface of the globe. At that time, in fact, the climate of regions nearer the equator must have been far more favourable to the human species, and it is possible that under such conditions it may have attained a higher development than in the north." (Huxley.) Professor Huxley points out also, in the interesting article "The Aryan Question," in The Contemporary Review for November, 1890, from which we have taken the above extracts, that the Irish river-bed skulls, belonging to a dark-haired, long-headed race, and those of the Frisians, the blond, long-headed race, now living on the North German coast, unmistakably approach the Neanderthal and Spy type in many of their distinctive [ 223 ]characters, "a sure indication" of the physiological continuity with the Pleistocene Neanderthaloid Men. The skulls of some of the Australian aboriginals and of the broad-headed people of Borreby, in Denmark, also present a remarkable similarity to the Neanderthal skull—perhaps an indication that those are characters of a stage in the pedigree of the human species before it differentiated into any of the existing races. (Huxley.)

The next palæontological evidence of Man is found in the Neolithic cavern deposits, alluvial accumulations, peat mosses, lake bottoms, pile dwellings, and shell-mounds in various parts of Europe. Between the time that Palæolithic Man left the caves he occupied, and the date when the earlier Neolithic people began to deposit fragments of the records of their history in the kitchen-midden, which they piled in front of their shelters, a long period appears to have elapsed in many districts. The objects found in these refuse-heaps are not associated with the remains of the Mammoth, the Woolly Rhinoceros, or the Elephant, but with those of animals still living, or such as have lived down to within historical times. The remains of his skeleton indicate that Neolithic Man varied very much in stature. Some were tall, some short; some had long and others broad skulls. The long-skulled people had the same tall stature and cranial peculiarities as the blue-eyed, light haired, and long-headed Xanthochroi living at the present day in Eastern Prussia, North Belgium, Northern France, and Britain, though their bony fabric "bears marks of somewhat greater ruggedness and savagery." The broad-skulled Men were short, and agreed in physical characters with the majority of the people now inhabiting the Mediterranean sea-board—the Melanochroi—with black hair and black eyes. Many Neolithic graves have [ 224 ]given up also the remains of a tall, broad-skulled, and a short, long-skulled race.

Such are the only recovered links in the pedigree of our race, and extremely unsatisfactory they are; indeed, beyond these few spots in Western Europe, in California, and the Mississippi valley in North America, Palæontology is silent as to the history of Man, and sheds no light upon his origin, or his last pithecoid parents; for, in Professor Huxley's impressive words, "so far as that light is bright it shows him substantially as he is now, and when it grows dim it permits us to see no sign that he was other than he is now."