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strate the present di

Fig. 105.–Group of Crocodiles. A long-snouted Gharial or Gavial (Gavialis


gangeticus) on the top of Crocodilus acutus; a Nile Crocodile (C. vulgaris) in the
foreground; C. palustris, a "Mugger," in the right upper corner. Observe the
peculiar floating attitude of the young specimen.
{450}

There is the same tendency to transfer the choanae further back, owing to the
formation of a solid secondary roofing in of the mouth, to transform the
amphicoelous into procoelous vertebrae, to reduce the supratemporal foramina, and
to obtain a better development of the dorsal armour, whilst that on the ventral
side is gradually reduced. Lastly, there is a tendency towards a shortening and
broadening of the snout, a condition which has reached its culmination in the
Alligators, while the Gavials are survivals of another branch. The notches in the
premaxilla, for the reception of some of the lower teeth, have also been acquired
independently. Although the recent Crocodilia cannot now, as has been pointed out
by Boulenger, be separated into different families, no valid diagnoses being
possible owing to the existence of Tomistoma, their phylogeny shows them to belong
to at least two heterogeneous groups.

Key to the Genera of recent Crocodilia.

I. Snout very long and slender. The mandibular symphysis extends at least to the
fifteenth tooth, and is partly formed by the splenial bones.

a. Nasal bones very small, and widely separated from the premaxillaries ..........
Gavialis gangeticus, p. 451.

b. Nasal bones long, in contact with the premaxillaries .......... Tomistoma


schlegeli, p. 453.

II. Snout not slender, but triangular or rounded off. The mandibular symphysis does
not reach beyond the eighth tooth, and does not reach the splenial bones.

a. Fourth mandibular tooth fitting into a notch in the upper jaw.

1. Without a bony nasal septum .......... Crocodilus, p. 454.

2. Nasal bones dividing the nasal groove .......... Osteolaemus, p. 466.

b. Fourth mandibular tooth fitting into a pit in the upper jaw.

1. Without a bony nasal septum .......... Caiman, p. 471.

2. Nasal bones dividing the nasal groove .......... Alligator, p. 466.

Fam. 1. Teleosauridae, in the Lias and Oolite of Europe; marine.–Snout very long
and slender. Nasals widely separated from the premaxillae by the maxillaries.
Choanae at the posterior end of the palatines. In front of the eye a small sub-
lacrymal foramen. Supratemporal foramina large. Vertebrae amphicoelous. Anterior
limbs scarcely half as long as the posterior pair. The dermal armour consists of
two rows of broad scutes on the back, while the belly is protected by a shield of
numerous bony scutes, which are connected with each other by sutures. Teeth
numerous and rather slender. General appearance like that of Gavials.
{451}

Teleosaurus of the Middle and Upper Oolite in England and France. Snout very
slender. Nasals narrow and short. The under side is protected by a beautifully
finished armour, consisting of a square breast-shield of four rows of bony scutes,
and a larger, long, oval shield on the belly, with about six longitudinal and
seventeen transverse rows of scutes.

Mystriosaurus, of the Upper Lias in France and Germany, reached a length of 15


feet, and is characterised by an additional series of keeled but smaller caudal
plates running parallel with the middle pairs, which are neatly sutured together.

Fam. 2. Metriorhynchidae, in the Upper Oolite of Europe; marine.–Nasals broad


posteriorly, sometimes extending with a pointed wedge very near the premaxillae.
Without sub-lacrymal foramina. Eyes with a ring of ossifications in the sclerotic.
Dermal armour unknown. Vertebrae and choanae like those of the previous family.
Metriorhynchus and Geosaurus.

Fam. 3. Macrorhynchidae, in the freshwater deposits of the Purbeck, Wealden, and


Greensand of Europe. Snout long and slender. The nasals are narrow, and so
elongated that they meet a similar long extension of the premaxillaries. Choanae
between the palatines and pterygoids. Vertebrae amphicoelous. Dermal armour
consisting of two imbricating dorsal and eight ventral rows, e.g. Pholidosaurus of
the English Wealden.

Fam. 4. Gavialidae.–Snout long and slender. The choanae are situated entirely
within the pterygoids. Vertebrae procoelous. Members of this family make their
first appearance in the littoral marine deposits of the Upper Chalk of Europe and
North America; others are common in tertiary, marine, and freshwater deposits,
whilst only two genera and species occur now in the Oriental sub-region.

Thoracosaurus in the Upper Chalk of New Jersey and France and Belgium is
intermediate between Gavialis and Tomistoma. The prefrontal bones are very small,
while the lacrymals are very long and surround the nasals posteriorly. The nasals
themselves are slender, and reach the posterior likewise long and narrow
prolongations of the premaxillaries.

Gavialis.–The snout is extremely long and slender. The mandibular symphysis is so


long that it comprises a great portion of the splenial bones, and extends backwards
almost to the level of the last teeth and to the palatal foramina. The nasal
{452}bones are very short, and are separated from the premaxillaries by the long
suture of the maxillaries. About twenty-eight upper and twenty-five lower teeth on
each side.

G. gangeticus, the only recent species, is essentially Indian, inhabiting chiefly


the basins of the Ganges, Brahmaputra, and Indus; it occurs also in the Mahanadi of
Orissa and in Arakan, but does not live in the Irrawaddy, nor in the Narbada,
Kistna, and farther south. In spite of its great size, which reaches 20 feet or
even more, it is harmless, and lives entirely upon fish; hence its Hindustani name,
gharial, meaning fish-eater, of which the generic name is a corruption.

The nuchal and dorsal scutes form a continuous shield, but there are two small
postoccipital scutes. General colour, dark olive-brown above; the young are paler,
with dark markings. The male is remarkable for several peculiarities. The nose is
very much swollen, and can be inflated like a bag when the nostrils are closed. In
connexion herewith, probably produced by the recoil of the air in the long narial
passages towards the choanae or posterior nares, there is a pair of hollow globular
swellings, in large specimens of the size of a goose's egg. The shell of these
globes is formed by the dorsal wings of the palatine bones above the floor of the
choanae, and they extend forwards to the right and left of the ethmoid almost to
the vertical downward process of the prefrontals.
fig106

Fig. 106.–Skull of Gavialis gangeticus (the Gharial). × ⅛. F, frontal; J, jugal; L,


lacrymal; Mand, mandible; Mx, maxillary; Na, nasal; Par, parietal; Pm.c,
premaxillary; Prf, prefrontal; Ptf, postfrontal; Qj, quadrato-jugal; Sq, squamosal.

Although the Gharial is common enough, we know next to nothing about its habits,
and in zoological gardens it is rather rare. A. Anderson[139] has, however, made
the following observations. Forty eggs were dug out of the sand, where they were
{453}lying in two tiers, twenty below and twenty above, with a foot of sand
between. The young ran with amazing rapidity the moment they were hatched. Some of
them actually bit his fingers before he had time to remove the shell from their
bodies! The length of these new-born creatures was 15 to 16 inches, 9 of which
belonged to the long and slender tail.

Several fossil species have been described from the Pliocene deposits of the
Sivalik Hills of India; and in the same district occurred the closely allied
Rhamphosuchus crassidens, which reached the gigantic length of about 50 feet!

Tomistoma.–The general configuration of the skull and snout is that of Gavialis,


but the nasal bones are long and reach the premaxillaries, although not the nasal
groove, thereby separating the maxillaries. The first and fourth mandibular teeth
fit into notches of the upper jaw, while most of the others fit into pits between
the teeth of the upper jaw. About twenty upper and eighteen lower teeth on each
side.

T. schlegeli, the only species, reaches a length of 15 feet; it inhabits the rivers
and swamps of Borneo, Malacca, and Sumatra. Fossil specimens of Tomistoma have been
found in the Miocene of Malta and Sardinia. Gavialosuchus of the Miocene of Hungary
is closely allied.

Fam. 5. Atoposauridae.–The few members of this family, Atoposaurus, Alligatorium,


and Alligatorellus, lived in the Upper Oolitic period of France, and were small,
about one foot in length. The vertebrae are amphicoelous. The nasal groove is
divided by a prolongation of the nasal bones. The head is short, and in its general
shape rather like that of a lizard.

Fam. 6. Goniopholidae, in the Purbeck and Wealden of Europe and the corresponding
level of North America. The vertebrae are amphicoelous. The choanae are still
elongated but are situated between the palatines and pterygoids. The premaxillaries
are rather large, and each sends a broad triangular process between the nasal and
maxillary. The nasals are broad and are well separated from the nasal groove. The
splenials help to form the mandibular symphysis.

Goniopholis.–The general configuration of the skull is rather like that of


Crocodilus vulgaris. There is a pair of deep notches in the upper jaw for the
reception of the lower canine teeth. G. simus and G. crassidens in England and
continental Europe, and {454}others in Colorado, were large-sized Crocodiles, some
with a skull 2 feet in length. The dermal armour consisted of a pair of dorsal
rows, a thoracic and an abdominal shield, composed as in the Teleosauridae of six
to eight longitudinal sutured rows.

Fam. 7. Crocodilidae.–Beginning in the Upper Cretaceous period of Europe and North


America, many forms of Crocodiles, Alligators, and Caimans existed in the Tertiary
period in America, Europe, and India; persisting in Europe until the Plistocene.
The vertebrae are procoelous. The choanae are completely surrounded by the
pterygoids. The nasals reach the nasal groove, except in Crocodilus cataphractus.
The orbits are larger than the small supratemporal fossae, and always continuous
superficially with the lateral temporal fossae, the postfronto-jugal bridge not
reaching the surface. The dorsal armour consists of more than one pair of
longitudinal rows, while the ventral armour is much reduced in thickness or absent.
Diplocynodon.–Common in the Oligocene and Miocene of Europe, e.g. D. hastingsiae.
The skull resembles that of the Alligators, but has a pair of lateral notches in
the premaxilla for the reception of the third, and sometimes also of the fourth
mandibular tooth. The ventral armour is still rather strong.

Crocodilus.–The fourth mandibular tooth fits, as a rule, into a notch in the upper
jaw. The other teeth are more or less interlocked with those of the other jaw. The
fifth upper tooth is the largest. The nasal bones form the posterior border of the
nasal groove, but do not extend into it as a septum. The bony scutes of the dorsal
shield are keeled, and stand closely together, being rarely united by suture; and
they form from four to six principal rows.

Crocodiles have occurred since the U

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