The skull (cf. Fig. 54, H, I, K, p. [280]) agrees fundamentally with that of Sphenodon and of the Crocodilia, but it is characterised by several special features. There are no ectopterygoids or ossa transversa; no lacrymal bones, no interparietal or pineal foramen; the vomer is unpaired and the nasal bones are mostly absent, unless they are fused with the prefrontals. The premaxillae are very small. The single vomer forms a septum between the choanae; and these are, except in Sphargis, ventrally roofed over by wings sent out by the palatines. The latter form a continuous bony roof to the mouth with the pterygoids, and these diverge posteriorly, being connected suturally with the quadrates, lateral and basi-occipital bones, and with the unpaired basi-sphenoid, which appears between the basi-occipital and the diverging pterygoids, but is in most cases to a great extent overlapped by the latter. The occipital condyle is distinctly triple; the basi-occipital sometimes helps to border the foramen magnum. The supra-occipital sends out a long vertical blade, directed backwards and generally projecting far over the neck, for the attachment of the powerful cranio-cervical muscles. The quadrate is very peculiar. Firmly attached, and hemmed in on nearly all sides by the neighbouring bones, it stands nearly vertically and forms a broad articulating surface for the mandible. Its posterior side shows either a transverse, horizontal groove, in which lies the columella auris, or the groove is transformed into a more or less closed canal. Moreover, the hinder lateral margin of the quadrate forms most of the tympanic frame; its margins being curved backwards, leaving in the Cryptodira, however, a wide notch behind; in the Pleurodira this part of the quadrate is transformed into a trumpet, the wide rim of which, forming a complete ring, carries the tympanic membrane. The tympanic cavity thus formed often leads into a deep recess which extends beneath the squamosal towards the opisthotic and bears some resemblance to the intricate tympanic recesses which pervade that region of the Crocodilian skull.
Dorsally the quadrate is broadly overlaid by the squamosal, which frequently forms an arch with the parietal. Anteriorly the quadrate is connected through a variably sized quadrato-jugal with the jugal; and this, by joining the maxilla and postfrontal, helps normally to form the posterior rim of the orbit. All the bones which border the temporal fossa vary much in extent in the different groups of Chelonia. The extremes are represented by Cistudo and Geoemyda, in which the bony infratemporal arch is absent, owing to the loss of the quadrato-jugal; and on the other hand by the Chelonidae and by Sphargis, in which the whole temporal region is covered over by an additional "false cranial" roof. This roof is produced chiefly by lateral wing-like expansions of the parietal and postfrontal bones, which meet the likewise much expanded jugal, quadrato-jugal, and squamosal bones. In the lower diagram of Fig. 63 (Chelone mydas) the squamosal has been removed, and the other bones have been reduced to their normal, or rather primitive condition, for comparison with the external view of the complete skull of the same animal. The lower diagram shows also the connexion of the pterygoid with a descending process of the parietal; this column, paired of course, usually contains a separate bone, the epipterygoid, the portion between Ptg and Par.
The hyoidean apparatus is well developed, and sometimes assumes large dimensions, especially in Chelys. The two pairs of "horns" are the first and second branchial arches, whilst the hyoid arches are reduced to a pair of small, frequently only cartilaginous, nodules attached near the anterior corners of the basis linguae, which generally fuses with the os entoglossum in the tip of the tongue.
The pectoral arch consists of a pair of long coracoids sloping obliquely backwards, the distal cartilages of which scarcely touch each other in the middle line, and the scapulae. The upper end of the scapula frequently touches the inside of the first costal plate, protected by a cartilaginous pad. Near the glenoid cavity arises a long process (PC in Fig. 65), placed transversely and approaching its fellow. The distal end is connected with that of the coracoid by a fibro-cartilaginous band. The homology of this scapular process is not quite clear. The band just mentioned favours the idea that the process represents the precoracoid, but its being an outgrowth from the scapula suggests that it is merely the much enlarged acromion. It certainly does not represent the clavicle, which forms part of the plastron: and this is not in contact with the shoulder-girdle at all.
Fig. 64.–Diagram of the skeleton of Testudo elephantopus, after removal of the left half of the carapace. The plastron is roughly indicated by a section through the middle line. Fe, Femur, foreshortened; Fi, fibula; H, humerus; Il, ilium; Is, ischium; P.P., pubis; R, radius; Scap, scapula; Tb, tibia; u, ulna; 3, third cervical vertebra; 1, 3, 5, first, third, and fifth fingers; XIII, thirteenth (fifth thoracic) vertebra.
The pelvis is strong. Ilium, pubis, and ischium meet at the acetabulum. The dorsal end of the ilium is generally broadened, and is attached to one or both sacral vertebrae, but it is also in contact with the superimposed last costal plate. This additional connexion often becomes predominant and the sacral vertebrae are partly or completely relieved of the iliac support, fusing in this case more or less with the costal plates. The pubes have strong lateral processes, directed obliquely forwards and downwards. The pubes and the ischia, which latter are much smaller, form broad symphyses, and these are connected with each other by a longitudinal cartilaginous band (Chelone, Trionyx); or the connecting bridge is broad and quite ossified (Testudo), forming in the latter case two roundish obturator-foramina. Cartilage frequently remains at the anterior end of the pubic symphysis, and a smaller, longer, and narrow piece of cartilage extends sometimes backwards from the ischiadic symphysis, as the so-called hypo-ischium. In the Pleurodira the ends of the ilia, and those of the lateral processes of the pubes, are much broadened and firmly ankylosed with the posterior costal plates and with the xiphiplastron respectively.
Fig. 65.–Ventral view of the bony shell of Chelone mydas, the Green Turtle, after removal of the plastron (Fig. 66). The costal plates are marked by cross lines to distinguish them from the ribs. C, coracoid; Fe, femur; Fi, fibula; H, humerus; Ma.1-Ma.12, marginal plates, some of which are fused together; Nu, nuchal plate; PC, "precoracoid"; R, radius; Sc, scapula; I, V, first and fifth digits; IX, Ninth vertebra or first thoracic.
The limbs are typically pentadactyle and complete, and are most primitive in water-tortoises, e.g. Chelydra and Emys, in which the carpus consists of the typical ten separate elements, including the pisiform. In Testudo the centrale is fused with the intermedium, and the first three distal carpals are also fused together. In the marine turtles the limbs are transformed into paddles, but all the bones retain their independence; the pisiform and the first metacarpal are enlarged and flattened, thereby giving additional width to the paddle. The tarsus remains less primitive; the centrale and the proximal elements have a tendency to fuse together, most completely in land-tortoises; the fifth distal carpal is enlarged, and stands out hook-like from the rest. The number of the phalanges of the fingers and toes varies slightly. It is noteworthy that none of the Chelonia possess more than three phalanges. The three middle fingers and toes have mostly three phalanges; the pollex and hallux have always two; the number of phalanges of the fifth finger varies from three to one, of the fifth toe from two to none. The greatest reduction occurs in Testudo and its allied genera of typical land-tortoises, Homopus, Pyxis, and Cinixys, the formula for the fingers being 2, 2, 2, 2, 2 or 1, and 2, 2, 2, 2, 0 for the toes. In Pelomedusa all the fingers possess two phalanges only, owing to fusion of the first and second phalanges with each other.