nts of the first gill-clefts) by bone.

In the earlier members they were still wide

slits or open grooves on the ventral side of the basi-occipital bone. Since the
Cretaceous epoch they have been transformed into bony canals and open through one
median hole, situated between the basi-occipital and the {435}basisphenoid,
immediately behind the posterior symphysis of the dorsal portion of the pterygoids,
which latter almost completely cover the basisphenoid. The vomer is not visible
(except in Caiman niger), being covered by the ventral junction of the palatines
and maxillaries. The broad, lateral wings of the pterygoids are connected by
separate bones, the ectopterygoids = transpalatines = transverse bones, with the
maxillaries, and in recent forms also with the jugals. Thus an extensive, very firm
bony palate is produced; and the large palatal foramina, between the palatines,
maxillaries, ectopterygoids and pterygoids, are closed by the same dense mucous
membrane which cover the whole roof of the mouth.

The opisthotic and epi-otic bones fuse early with the lateral and with the supra-
occipital bones; only the pro-otic remains longer as a separate element, perforated
anteriorly by a large hole for the exit of the third branch of the trigeminal
nerve. The basisphenoid is scarcely visible, being covered by the pterygoids. The
presphenoid is large, continued forwards and upwards into the usually cartilaginous
interorbital septum. Near the anterior and upper margin of the presphenoid is a
large notch on either side for the passage of the optic nerve, the three eye-muscle
nerves and the first branch of the trigeminal nerve. There are no separate orbito-
sphenoids, their place being taken by membrane or cartilage in continuation with
the interorbital septum, but the alisphenoids are large, abutting upwards against
the frontals. Each prefrontal sends down a vertical process which joins the
palatine of its side.

The configuration of the snout varies much. There are two parallel lines of
development since the Jurassic epoch, namely, long-snouted creatures, of which two
still survive as Gavialis and Tomistoma, and more broad and short-snouted members
like the rest of the Crocodiles and Alligators. In opposition to the Parasuchia the
elongation of the snout is effected by the maxillaries. The length of the nasals
varies much, mostly in conformity with that of the maxillaries. As a rule they
reach the premaxillaries but not always the nasal groove. In Gavialis they are
short, far separated from the premaxillaries by the maxillaries, which meet in the
dorso-median line. The orbit is bordered by the frontals, which at an early age
fuse into an unpaired piece, and by the prefrontal, lacrymal, jugal, and
postfrontal. {436}At a deeper level the orbit is partly divided from the lateral
temporal fossa by a strong column which is formed by the meeting of a downward
process of the postfrontal with an inner process of the jugal, and an ascending
process of the ectopterygoid (cf. Fig. 108, p. 458). This arrangement adds
considerably to the strength of the skull. The lateral temporal fossa is bordered
in front by the column just described; below by the jugal and the quadrato-jugal,
which is firmly wedged in between the jugal and quadrate; behind by the quadrate;
above by the postfrontal, which forms a strong superficial bridge with the
squamosal. This rests upon and often fuses with the quadrate and an intervening
transverse wing-like extension of the lateral occipital bone. By this squamoso-
postfrontal bridge part of the original temporal fossa is divided into the lateral
one just described, and a dorsal fossa. The latter is bordered by the postfrontal,
squamosal, and united parietals. This dorsal temporal fossa is consequently not
homologous with that of the Parasuchia, a vestige of which is however present in
many, especially in young skulls of Crocodiles, in the shape of a narrow passage
which extends backwards from the dorsal fossa, bridged over by the junction of the
parietal with the squamosal, and bordered below by the occipitals.

The size of the upper temporal fossae stands in an inverse ratio to that of the
lateral fossae. In the older Eusuchia the upper were the larger of the two. The
temporo-mandibular muscle which lifts or shuts the lower jaw arises from the walls
of the upper fossa, passes beneath the jugal arch, and is inserted into the supra-
angular portion of the lower jaw. In the more recent Crocodiles this muscle is more
and more superseded by the pterygo-mandibular muscle, which, arising chiefly from
the dorsal surface of the much broadened-out pterygoid bone, fills the widened
space between the latter and the quadrate, and is inserted into the outer surface
of the os angulare of the lower jaw. This muscle, owing to its general disposition,
is capable of much more powerful development and leverage than the temporo-
maxillary muscle, which latter, being more reduced, allows the dorsal fossae to be
more and more closed up by the surrounding bones.

The fossae are still comparatively large in the long-snouted genera Gavialis and
Tomistoma, which live entirely upon fish and scarcely chew their food, whilst these
holes almost completely {437}disappear in some of the Alligators, namely in the
broad- and short-snouted members, which, having a varied diet, taken from every
available group of the animal kingdom, chew their prey.

The quadrate extends obliquely backwards, and is immovably wedged in and partly
fused with the quadrato-jugal, the squamosal, and the lateral occipital wings.
Between the latter and the quadrate remains a slit-like canal, well visible from
behind, through which passes the continuation into the mandible of the columellar
or ossicular chain of the auditory apparatus. Intricate passages, used as
additional enlargements of the space of the middle ear, pervade the proximal
portions of the quadrate and the roof of the cranium beneath the parietal bridges
mentioned above, the two sides communicating with each other. The supra-occipital
bone is visible from behind; its top is covered and partly fused with a
continuation of the parietals, which are, like the frontals, fused into an unpaired
mass. The occipital condyle is formed entirely by the basi-occipital bone, so far
as the articulating facet is concerned, but it is supported on either side by a
lamella from the lateral occipitals.

The two halves of the lower jaw form a symphysis of very variable length. Each half
is composed of six bones. (1) The articulare, perforated in its upper, posterior,
inner corner by a canal for the reception of the siphonium, a narrow tube of
connective tissue, which connects the cavities of the middle ear with the large
empty space enclosed within the lower jaw; (2) the angulare; (3) the dentary, which
alone carries the teeth; (4) the splenial, a long splint-like bone on the surface
of the inner or median side of the jaw, of variable length; (5) the operculare, the
counterpart of the splenial on the outer side; (6) the supra-angulare, which forms
the dorsal border of the lower jaw between the dentary and the angulare.

The teeth, which are more or less conical or compressed laterally, are deeply
implanted in separate sockets. They are often shed throughout life, the successors
lying on the median side, and with their caps partly fitting into the wide, open
roots of the teeth to be expelled. The number of teeth in the premaxilla is
universally five on either side in recent forms, but in a few species, e.g.
Crocodilus niloticus and C. porosus, the second pair is lost with maturity and is
not replaced. In the broad-snouted {438}kinds, especially in the Alligators, most
of the upper teeth overlap laterally those of the lower jaw. In most species of
Crocodilus the overlapping is less marked and the teeth partly interlock, but the
fourth mandibular tooth, generally the strongest and longest, is received into a
lateral notch at the junction of the premaxillary and maxillary. Frequently those
of the longer lower teeth which fit into pits of the upper jaw, gradually transform
the pits into holes by continued pressure upon the bone, and in old specimens the
tip of the lower tooth may even perforate and stand out above the skin of the
snout.

The vertebrae are solid, but remnants of the notochord persist for a long time in
the middle of the centra. These are still amphicoelous in the Jurassic Eusuchia,
and there were probably considerable intervertebral portions of the notochord. From
the Lower Chalk onwards the vertebrae are procoelous, with the exception of the
first caudal vertebra, which has a knob at either end, so that naturally the
posterior of the two sacral vertebrae is opisthocoelous. This peculiar formation of
the first caudal is probably correlated with the flexibility of the tail.

Cartilaginous intercentral rings, pads or menisci, occur regularly throughout the

vertebral column, unless they are abolished by fusion of adjoining vertebrae. It is
most instructive to follow the attachment of the ribs in one and the same
individual. The position of the capitulum, vertically below the tuberculum in the
neck, changes in the thorax into one in which the capitulum lies anterior to the
tuberculum and in the same horizontal plane with it. Moreover, whilst on the
cervical vertebrae the capitulum is carried by the centrum (enclosing with the
tuberculum a typical transverse canal for the vertebral artery, etc.), further back
it moves its point of attachment upwards, lying right upon the neuro-central suture
on the tenth and eleventh vertebrae. From the twelfth vertebra backwards both
capitulum and tuberculum are carried by the transverse process or diapophysis of
the neural arch. The ribs of the five or six lumbar vertebrae are merely vestigial
or absent. The ribs of the two sacral vertebrae are very stout, fusing in the adult
with both centrum and neural arch. Some of the anterior caudal vertebrae also carry
ribs, attached across the neuro-central suture; long before maturity they fuse with
their vertebrae, and then look like transverse processes. Most of the caudal
vertebrae carry also a {439}pair of chevron-bones, and these are continuous with
the intercentral rings of cartilage.

The atlas and the epistropheus or axis are of supreme interest. Crocodiles are, in
fact, the only animals in which these two vertebrae retain all their constituent
hard parts in an almost undisturbed primitive condition (Fig. 103, 1-4). The basal
piece of the atlas-ring, the first basiventral or intercentrum, carries a pair of
long ribs attached by their capitular portions. A small knob near the dorsal edge
of the rib occurs in many specimens, and is the last remnant of the tubercular
portion. The latter was still complete in Jurassic Crocodiles, for instance in
Metriorhynchus (Fig. 103, 2, t1). The first centrum joins that of the second
vertebra as its so-called odontoid process, not directly, however, but by the
intercalation of the complete second basiventral, represented by a cartilaginous
disc, and by a large unpaired pyramidal piece (Fig. 103, 3, 2). This, serially
homologous with the ventral half of the atlas-ring, is the second basiventral
intercentrum, wedged in from below between the odontoid process and the second
centrum, with which it soon fuses. Moreover, it carries the capitulum of the second
rib (2, Cp2), the tuberculum of which is articulated with a facet of the second
neural arch in Jurassic Eusuchia (t2). In recent Crocodiles this tubercular portion
is much reduced, and, curiously enough, is attached to a knob which belongs to the
odontoid piece or first centrum. This shifting explains the apparently anomalous
condition that "the atlas of the Crocodiles carries two pairs of ribs, the second
vertebra none." To complete the account of the atlas we have to mention the
separate unpaired piece which lies upon the two neural arches. It is the detached
neural spine, and not the remnant of a "pro-atlas."

The first and seco