1911 Encyclopædia Britannica/Artiodactyla

by Richard Lydekker

ARTIODACTYLA (from Gr. ἄρτιος, even, and δάκτυλος, a finger or toe, “even-toed”), the suborder of ungulate mammals in which the central (and in some cases the only) pair of toes in each foot are arranged symmetrically on each side of a vertical line running through the axes of the limbs. As contrasted with the Perissodactyla living, and in a great degree extinct, Artiodactyla are characterized by the following structural features. The upper premolar and molar teeth are not alike, the former being single and the latter two-lobed; and the last lower molar of both first and second dentition is almost invariably three-lobed. Nasal bones not expanded posteriorly. No alisphenoid canal. Dorsal and lumbar vertebrae together always nineteen, though the former may vary from twelve to fifteen. Femur without third trochanter. Third and fourth digits of both feet almost equally developed, and their terminal phalanges flattened on their inner or contiguous surfaces, so that each is not symmetrical in itself, but when the two are placed together they form a figure symmetrically disposed to a line drawn between them. Or, in other words, the axis or median line of the whole foot is a line drawn between the third and fourth digits (fig. 1). Lower articular surface of the astragalus divided into two nearly equal facets, one for the navicular and a second for the cuboid bone. The calcaneum with an articular facet for the lower end of the fibula. Stomach almost always more or less complex. Colon convoluted. Caecum small. Placenta diffused or cotyledonary. Teats either few and inguinal, or numerous and abdominal.

Artiodactyla date from the Eocene period, when they appear to have been less numerous than the Perissodactyla, although at the present day they are immeasurably ahead of that group, and form indeed the dominant ungulates. As regards the gradual specialization and development of the modern types, the following features are noteworthy.

1. As regards the teeth, we have the passage of a simply tubercular, or bunodont (βουνός, a hillock) type of molar into one in which the four main tubercles, or columns, have assumed a crescentic form, whence this type is termed selenodont (σελήνη, the new moon). Further, there is the modification of the latter from a short-crowned, or brachyodont type, to one in which the columns are tall, constituting the hypsodont, or hypsiselenodont, type. It is noteworthy, however, that in some instances there appears to have been a retrograde modification from the selenodont towards the bunodont type, the hippopotamus being a case in point. Other modifications are the loss of the upper incisors; the development of the canines into projecting tusks; and the loss of the anterior premolars.

2. As regards the limbs. Reduction of the ulna from a complete and distinct bone to a comparatively rudimentary state in which it coalesces more or less firmly with the radius. Reduction of the fibula till nothing but its lower extremity remains. Reduction and final loss of outer pair of digits (second and fifth), with coalescence of the metacarpal and metatarsal bones of the two middle digits to form a cannon-bone. Union of the navicular and cuboid, and sometimes the ectocuneiform bone, of the tarsus.

3. Change of form of the odontoid process of the second or axis vertebrae from a cone to a hollow half-cylinder.

4. Development of horns or antlers on the frontal bones, and gradual complication of form of antlers.

5. By inference only, increasing complication of stomach with ruminating function superadded. Modification of placenta from simple diffused to cotyledonary form.

Fig. 1.—Bones of Right Fore Feet of existing Artiodactyla.
A,	Pig (Sus scrofa).	l,	Lunar.
B,	Red deer (Cervus elaphus).	s,	Scaphoid.
C,	Camel (Camelus bactrianus).	u,	Unciform.
U,	Ulna.	m,	Magnum.
R,	Radius.	td,	Trapezoid.
c,	Cuneiform.
In the Sheep and the Camel the long compound bone, supporting the two main (or only) toes is the cannon-bone.

The primitive Artiodactyla thus probably had the typical number (44) of incisor, canine and molar teeth, brachyodont molars, conical odontoid process, four distinct toes on each foot, with metacarpal, metatarsal and all the tarsal bones distinct, and no frontal appendages.

As regards classification, the first group is that of the Pecora, or Cotylophora, in which the cheek-teeth are selenodont, but there are no upper incisors or canine-like premolars, while upper canines are generally absent, though sometimes largely developed. Inferior incisors, three on each side Pecora.with an incisiform canine in contact with them. Cheek-teeth consisting of p.3/3, m.3/3, in continuous series. Auditory bulla simple and hollow within. Odontoid process of second vertebra in the form of a crescent, hollow above. Lower extremity of the fibula represented by a distinct malleolar bone articulating with the outer surface of the lower end of the tibia. Third and fourth metacarpals and metatarsals confluent into cannon-bones (fig. 1 B), and the toes enclosed in hoofs. Outer toes small and rudimentary, or in some cases entirely suppressed; their metacarpal or metatarsal bones never complete. Navicular and cuboid bones of tarsus united. The skull generally lacks a sagittal crest; and the condyle of the lower jaw is transversely elongated. Horns or antlers usually present, at least in the male sex. Left brachial artery arising from a common innominate trunk, instead of coming off separately from the aortic arch. Stomach with four complete cavities. Placenta cotyledonous. Teats 2 or 4.

The group at the present day is divided into Giraffidae (giraffe and okapi), Cervidae (deer), Antilocapridae (prongbuck), and Bovidae (oxen, sheep, goats, antelopes, &c.). (See Pecora.)

The second group is represented at the present day by the camels (Camelus) of the Old, and the llamas (Lama) of the New World, collectively constituting the family Camelidae. They derive their name of Tylopoda (“boss-footed”) from the circumstance that the feet form large cushion-like pads, Tylopoda.supporting the weight of the body, while the toes have broad nails on their upper surface only, instead of being encased in hoofs. The cheek-teeth are selenodont, and one pair of upper incisors is retained, while some of the anterior premolars assume a canine-like shape, and are separated from the rest of the cheek-series. Auditory bulla filled with honeycombed bony tissue. Odontoid process of second vertebra semi-cylindrical; skull with a sagittal crest; and the condyle of the lower jaw rounded. Third and fourth metacarpals and metatarsals (which are alone present) fused into cannon-bones for the greater part of their length, but diverging inferiorly (fig. 1, C) and with their articular surfaces for the toes smooth, instead of ridged as in the Pecora. Navicular and cuboid bones of tarsus distinct. No horns or antlers. Stomach, although complex, differing essentially from that of the Pecora. Placenta diffuse, without cotyledons. Teats few. (See Tylopoda.)

In the same sectional group is included the North American family of oreodonts (Oreodontidae), which are much more primitive ruminants, with shorter necks and limbs, the full series of 44 teeth, all in apposition, and the metacarpal and metatarsal bones separate, and the toes generally of more normal type, although sometimes claw-like. (See Oreodon.) The Eocene American genus Homacodon is regarded as representing a third family group, the Homacodontidae (= Pantolestidae), in which the molars were of a bunodont type, and approximate to those of the Condylarthra from which this family appears to have sprung, and to have given origin on the one hand to the Oreodontidae, and on the other to the Camelidae. The family is represented in the Lower, or Wasatch, Eocene by Trigonolestes, in the Middle (Bridger) Eocene by Homacodon (Pantolestes), and in the Upper (Uinta) Eocene by Bunomeryx.

The third group is that represented by the chevrotains or mouse-deer, forming the family Tragulidae, with Tragulus in south-eastern Asia and Dorcatherium (or Hyomoschus) in equatorial Africa. The cheek-teeth are selenodont, as in the two preceding groups; there are no upper incisors, but Tragulina.there are long, narrow and pointed upper canines, which attain a large size in the males; the lower canines are incisor-like, as in the Pecora, and there are no caniniform premolars in either jaw. Cheek-teeth in a continuous series consisting of p.3/3, m.3/3. Odontoid process of axis conical. Fibula complete. Four complete toes on each foot. The middle metacarpals and metatarsals generally confluent, the outer ones (second and fifth) slender but complete, i.e. extending from the carpus or tarsus to the digit. Navicular, cuboid and ectocuneiform bones of tarsus united. Auditory bulla of skull filled with cancellar tissue. No frontal appendages. Ruminating, but the stomach with only three distinct compartments, the maniplies or third cavity of the stomach of the Pecora being rudimentary. Placenta diffused. (See Chevrotain.)

In this place must be mentioned the extinct Oligocene European group typified by the well-known genus Anoplotherium of the Paris gypsum-quarries, and hence termed Anoplotherina, although the alternative title Dichobunoidea has been suggested. It includes the two Anoplotherina.families Anoplotheriidae and Dichobunidae, of which the first died out with the Oligocene, while the second may have given origin to the Tragulina and perhaps the Pecora. There is the full series of 44 teeth, generally without any gaps, and most of the bones of the skeleton are separate and complete; while, in many instances at any rate, the tail was much longer than in any existing ungulates, and the whole bodily form approximated to that of a carnivore. The upper molars, which may be either selenodont or buno-selenodont, carry five cusps each, instead of the four characteristic of all the preceding groups; and they are all very low-crowned, so as to expose the whole of the valleys between the cusps. In Anoplotherium, some of the species of which were larger than tapirs, there were either two or three toes, the latter number being almost unique among the Artiodactyla. Allied genera are Diplobune and Dacrytherium.

The Dichobunidae include the genus Dichobune, of which the species were small animals with buno-selenodont molars. Xiphodon and Dichodon represent another type with cutting premolars and selenodont molars; while Caenotherium and Plesiomeryx form yet another branch, with resemblances to the ruminants. The most interesting genera are however, the Upper Oligocene and Lower Miocene Gelocus and Prodremotherium, which have perfectly selenodont teeth, and the third and fourth metacarpal and metatarsal bones respectively fused into an imperfect cannon-bone, with the reduction of the lateral metacarpals and metatarsals to mere remnants of their upper and lower extremities. While Gelocus exhibits a marked approximation to the Tragulidae, Prodremotherium comes nearer to the Cervidae, of which it not improbably indicates the ancestral type. The Dichobunidae may be regarded as occupying a position analogous to that of the Homacodontidae in the Tylopoda, and like the latter, are probably the direct descendants of Condylarthra.

Fig. 2.—Restoration of Anoplotherium commune.

The last section of the Artiodactyla is that of the Suina, represented at the present day by the pigs (Suidae), and the hippopotamuses (Hippopotamidae), and in past times by the Anthracotheriidae, in which may probably be included the Elotheriidae. In the existing members of the group Suina.the cheek-teeth approximate to the bunodont type, although showing signs of being degenerate modifications of the selenodont modification. There is at least one pair of upper incisors, while the full series of 44 teeth may be present. The metacarpals and metatarsals are generally distinct (fig. 1 A), and never fuse into a complete cannon-bone; and the navicular and cuboid bones of the tarsus are separate. The odontoid process of the second vertebra is pig-like: and the tibia and fibula and radius and ulna are severally distinct. The stomach is simple or somewhat complex, and the placenta diffused. The Suidae include the Old World pigs (Suinae) and the American peccaries (Dicotylinae), and are characterized by the snout terminating in a fleshy disk-like expansion, in the midst of which are perforated the nostrils; while the toes are enclosed in sharp hoofs, of which the lateral ones do not touch the ground. There is a caecum. The Dicotylinae differ from the Suinae in that the upper canines are directed downwards (instead of curving upwards) and have sharp cutting-edges, while the toes are four in front and three behind (instead of four on each foot), and the stomach is complex instead of simple. In the Old World a large number of fossil forms are known, of which the earliest is the Egyptian Eocene Geniohyus. Originally the family was an Old World type, but in the Miocene it gained access into North America, where the earliest form is Bothriolabis, an ancestral peccary showing signs of affinity with the European Miocene genus Palaeochoerus. (See Swine and Peccary.)

The Hippopotamidae are an exclusively Old World group, in which the muzzle is broad and rounded and quite unlike that of the Suidae, while the crowns of the cheek-teeth form a distinctly trefoil pattern, when partially worn, which is only foreshadowed in those of the latter. The short and broad teeth terminate in four subequal toes, protected by short rounded hoofs, and all reaching the ground. The hinder end of the lower jaw is provided with a deep descending flange. Both incisors and canines are devoid of roots and grow throughout life, the canines, and in the typical species one pair of lower incisors, growing to an immense size. The stomach is complex; but there is no caecum. Although now exclusively African, the family (of which all the representatives may be included in the single genus Hippopotamus, with several subgeneric groups) is represented in the Pliocene of Europe and the Lower Pliocene of northern India. Its place of origin cannot yet be determined.

The extinct Anthracotheriidae were evidently nearly allied to the Hippopotamidae, of which they are in all probability the ancestral stock. They agree, for instance, with that family in the presence of a descending flange at the hinder end of each side of the lower jaw; but their dentition is of a more generalized type, comprising the full series of 44 teeth, among which the incisors and canines are of normal form, but specially enlarged, and developing roots in the usual manner. The molars are partially selenodont in the typical genus Anthracotherium, with five cusps, or columns, on the crowns of those of the upper jaw, which are nearly square. The genus has a very wide distribution, extending from Europe through Asia to North America, and occurring in strata which are of Oligocene and Miocene age. In Ancodon (Hyopotamus) the cusps on the molars are taller, so that the dentition is more decidedly selenodont; the distribution of this genus includes not only Europe, Asia and North Africa, but also Egypt where it occurs in Upper Eocene beds in company with the European genus Rhagatherium, which is nearer Anthracotherium. On the other hand, in Merycopotamus, of the Lower Pliocene of India and Burma, the upper molars have lost the fifth intermediate cusp of Ancodon; and thus, although highly selenodont, might be easily modified, by a kind of retrograde development, into the trefoil-columned molars of Hippopotamus. In the above genera, so far as is known, the feet were four-toed, although with the lateral digits relatively small; but in Elotherium (or Entelodon), from the Lower Miocene of Europe and the Oligocene of North America, the two lateral digits in each foot had disappeared. This is the more remarkable seeing that Elotherium may be regarded as a kind of bunodont Anthracotherium. It shows the characteristic hippopotamus-flange to the lower jaw, but has also a large descending process from the jugal bone of the zygomatic arch of the skull. Finally, we have in the Pliocene of India the genus Tetraconodon, remarkable for the enormous size attained by the bluntly conical premolars; as the molars are purely bunodont, this genus seems to be a late and specialized survivor of a primitive type.  (R. L.*)