its anterior surface, and the two bones, instead of being external and internal, are anterior and posterior. In many hoofed or Ungulate mammals, and in Bats, the ulna is reduced to little more than its upper articular extremity, and firmly ankylosed to the radius -stability of these parts being more essential than mobility.

Manus.-The terminal segment of the anterior limb is the hand or manus. Its skeleton consists of three divisions: (1) the carpus," a group of small, more or less rounded or angular bones with flattened surfaces applied to one another, and, though articulating by synovial joints, having scarcely any motion between them; (2) the "metacarpus," a series of elongated bones placed side by side, with their proximal ends articulating by almost immovable joints with the carpus; (3) the "phalanges" or bones of the digits, usually three in number to each, articulating to one another by freely movable hinge-joints, the first being connected in like manner to the distal end of the metacarpal bone to which it corresponds.

Carpus. To understand thoroughly the arrangement of the bones of the carpus in mammals, it is necessary to study their condition in some of the lower vertebrates. Fig. 17 represents the manus in one of its fullest and at the same time most

FIG. 17.-Dorsal surface of the

right manus of a Water Tortoise (Chelydra serpentina). After Gegenbaur. U, Ulna; R, radius; u, ulnare; i, intermedium; r, radiale;

c, centrale; 1-5, the five bones of the distal row of the carpus; ml

m3, the five metacarpals.

generalised forms, as seen in one of the Water Tortoises (Chelydra serpentina). The carpus consists of two principal rows of bones. The upper or proximal row contains three bones, to which Gegenbaur has applied the terms radiale (r), intermedium (i), and ulnare (u), the first being on the radial or preaxial side of the limb.' The lower or distal row contains five bones, called carpale 1, 2, 3, 4, and 5 respectively, commencing on the radial side. Between these two rows, in the middle of the carpus, is a single bone, the centrale (c). In this very symmetrical carpus it will be observed that the radiale supports on its distal side two bones, carpale 1 and 2; the intermedium is in a line with the centrale and carpale 3, which together form a median axis of the hand, while the ulnare has also two bones articulating with its distal end, viz. carpale 4 and 5. Each of the carpals of the distal row supports a metacarpal.

1 The opinion has recently been expressed by Baur that the bone termed radiale in Fig. 17 is really a second centrale, and that the radiale is represented by a minute bone generally known as the radial sesamoid. The mammalian

In the carpus of the Mammalia there are usually two additional bones developed in the tendons of the flexor muscles, one on each side of the carpus, which may be called the radial and ulnar sesamoid bones; the latter, which is the more constant and generally larger, is commonly known as the pisiform bone. The fourth and fifth carpals of the distal row are always united into a single bone, and the centrale is very often absent. As a general rule all the other bones are present and distinct, though it not unfrequently happens that two may have coalesced to form a single bone, or one or more may be altogether suppressed.

The following table shows the principal names in use for the various carpal bones,-those in the second column being the terms generally employed by English anatomists :

:

These

The radial and ulnar sesamoids are regarded by Bardeleben1 as the rudiments of a prepollex and a postminimus digit; the primitive number of digits being thus supposed to have been seven. bones have been observed in all orders of mammals having five complete digits. Occasionally, as in Pedetes caffer, the so-called prepollex consists of two bones, of which the distal one bears a distinct nail-like horny covering. In Bathyergus maritimus the pisiform, or postminimus, is likewise double; the two elements being regarded by their describer as representing the carpal and metacarpal of the presumed seventh digit.

Similarly in the posterior limb the tibial sesamoid, and a fibular ossification corresponding to the pisiform, are regarded as representing a prehallux and a postminimus.

Metacarpus and Phalanges. The metacarpal bones, with the digits which they support, are never more than five in number, and are described numerically-first, second, etc., counting from the radial towards the ulnar side. The digits are also sometimes named (1) the pollex, (2) index, (3) medius, (4) annularis, (5) minimus. scaphoid is accordingly also regarded as a second centrale. In the same communication, Dr. Baur expresses his disbelief in the existence of remnants of a prepollex and of a seventh digit in mammals and other vertebrates. (See Anat. Anzeiger, vol. iv. pp. 49-52, 1889.)

1 On the Præpollex and Præhallux, etc., Proc. Zool. Soc. 1889, pp. 259-262.

One or more may be in a rudimentary condition, or altogether suppressed. If one is absent, it is most commonly the first. Excepting the Cetacea, no mammals have more than three phalanges to each digit, but they may occasionally have fewer by suppression or ankylosis. The first or radial digit is an exception to the usual rule, one of its parts being constantly absent, since, while each of the other digits has commonly a metacarpal and three phalanges, it has only three bones altogether; whether the missing one is a metacarpal or one of the phalanges is a subject which has occasioned much discussion, and has not yet been satisfactorily decided. terminal phalanges of the digits are usually specially modified to support the nail, claw, or hoof, and are called "ungual phalanges." In walking, some mammals (as the Bears) apply the whole of the lower surface of the carpus, metacarpus, and phalanges to the ground; to these the term "plantigrade" is applied. Many others (as nearly all the existing Ungulata) only rest on the last one or two phalanges of the toes, the first phalanx and the metacarpals being vertical and in a line with the fore-arm. These are called "digitigrade." Intermediate conditions exist between these two forms, to which the terms "phalangigrade" (as the Camel) and "subplantigrade" (as in most Carnivora), are applied. When the weight is borne entirely on the distal surface of the ungual phalanx, and the horny structures growing around it, as in the Horse, the mode of progression is called "unguligrade."

In the Chiroptera the digits are enormously elongated, and support a cutaneous expansion constituting the organ of flight. In the Cetacea the manus is formed into a paddle, being covered by continuous integument, which conceals all trace of division into separate digits, and shows no sign of nails or claws. In the Sloths the manus is long and very narrow, habitually curved, and terminating in two or three pointed curved claws in close apposition with each other, and incapable, in fact, of being divaricated; so that it is reduced to the condition of a hook, by which the animal suspends itself to the boughs of the trees among which it lives. These are only examples of the endless modifications to which the distal extremity of the limb is subjected in adaptation to the various purposes to which it is applied.

Posterior Limb.-The posterior limb is constructed upon a plan very similar to that of the anterior extremity. It consists of a pelvic girdle and three segments belonging to the limb proper, viz. the thigh, the leg, and the foot or pes (Fig. 15).

Pelvic Girdle. The pelvic girdle is present in some form in all mammals, though in the Cetacea and the Sirenia it is in an exceedingly rudimentary condition. In all mammals except those belonging to the two orders just named, each lateral half of the pelvic girdle consists essentially, like the corresponding part of the anterior

limb, of a flattened rod of bone crossing the long axis of the trunk, having an upper or dorsal and a lower or ventral end. The upper end diverges from that of the opposite side, but the lower end approaches, and, in most cases, meets it, forming a symphysis, without the intervention of any bone corresponding to the sternum. The pelvic girdle differs from the shoulder girdle in being firmly articulated to the vertebral column, thus giving greater power to the hinder limb in its function of supporting and propelling the body. Like the shoulder girdle, it bears on its outer side, near the middle, a cup-shaped articular cavity ("acetabulum "), into which the proximal end of the first bone of the limb proper is received. Each lateral half of the girdle is called the OS innominatum," or innominate bone, and consists originally of three bones which unite at the acetabulum. The "ilium" or upper bone is that which articulates with the sacral vertebræ. Of the two lower bones the anterior or "pubis" unites with its fellow of the other side at the symphysis; the posterior is the "ischium." These lower elements form two bars of bone, united above and below, but leaving a space between them in the middle, filled only by membrane, and called the "thyroid" or "obturator" foramen. The whole circle of bone formed by the two innominate bones and the sacrum is called the pelvis. In the Monotremata and Marsupialia, a pair of thin, flat, elongated ossifications called epipubic or marsupial bones are attached to the fore part of the pubis, and project forward into the muscular wall of the abdomen.

Thigh and Leg.-The first segment of the limb proper has one bone, the femur, corresponding with the humerus of the anterior limb. The second segment has two bones, the tibia and fibula, corresponding with the radius and ulna. These bones always lie in their primitive unmodified position, parallel to each other, the tibia on the preaxial and the fibula on the postaxial side, and are never either permanently crossed or capable of any considerable amount of rotation, as in the corresponding bones of the fore limb. In the ordinary walking position the tibia is internal, and the fibula external. In many mammals the fibula is in a more or less rudimentary condition, and it often ankyloses with the tibia at one or both extremities. The patella or "knee-cap," which is found in an ossified condition in all mammals, with the exception of some of the Marsupialia, is a large sesamoid bone developed in the tendon of the extensor muscles of the thigh, where the tendon passes over the front of the knee-joint, to which it serves as a protection. There are frequently smaller ossicles, one or two in number, situated behind the femoral condyles, called "fabellæ." The processes for the attachment of muscles near the upper end of the femur are termed trochanters; and the third trochanter, found on the hinder

aspect of the shaft of this bone in many forms is of considerable taxonomic importance.

Pes. The terminal segment of the hind limb is the foot or pes. Its skeleton presents in many particulars a close resemblance to that of the manus, being divisible into three parts: (1) a group of short, more or less rounded or square bones, constituting the tarsus; (2) a series of long bones placed side by side, forming the metatarsus; and (3) the phalanges of the digits or toes.

The bones of the tarsus of many of the lower Vertebrata closely resemble both in number and arrangement those of the carpus, as shown in Fig. 17. They have been described in their most generalised condition by Gegenbaur under the names expressed in the first column of the following table. The names in the second column are those by which they are generally known to English anatomists, while in the third column some synonyms occasionally employed are added.

The bones of the tarsus of mammals present fewer diversities of number and arrangement than those of the carpus. The proximal row (see Fig. 18) always consists of two bones, namely the astragalus (a), which probably represents the coalesced scaphoid and lunar of the hand, and the calcaneum (c). The former is placed more to the dorsal side of the foot than the latter, and almost exclusively furnishes the tarsal part of the tibio-tarsal or ankle-joint. The calcaneum, placed more to the ventral or "plantar" side of the foot, is elongated backwards to form a more or less prominent tuberosity, the "tuber calcis," to which the tendon of the great extensor muscles of the foot is attached. The navicular bone (n) is interposed between the proximal and distal row on the inner or tibial side of the foot, but on the outer side the bones of the two rows come into contact. The distal row, when complete, consists of four bones, which, beginning on the inner side, are the three cuneiform bones, internal (c1), middle (c2), and external (c), articulated to the distal surface of the navicular, and the cuboid (cb), articulated with the calcaneum. Of these the middle cuneiform is usually the smallest in animals 1 Cope and Baur consider that the astragalus corresponds only with the intermedium, and that the tibiale may exist as a distinct element.