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Palæontological Report of the Princeton Scientific Expedition of 1877

REPTILIA.

Reading edition

CROCODILIA.

CROCODILUS.

Crocodilus aptus, Leidy.

Contributions to Extinct vertebrate fauna of the Western Territories, p. 126.

Henry's Fork.

Crocodilus Grinnelli, Marsh.

American Journal of Science and Arts, vol. i., p. 465.

From Cottonwood Creek.

Crocodilus Elliotii, Leidy.

Cont. to Ex. Vert. Fauna, p. 126.

Represented by a perfect skull, and several vertebræ.

This fossil exhibits a form of skull which shows characters of both crocodile and alligator; the latter to a less marked degree. That it properly belongs to the former genus is shown by the notch in the upper jaw which receives the canine of the lower.

The entire skull is remarkably flat on its upper surface, the face and cranium being nearly in the same plane without the descent at the frontals usual in these reptiles. The jaw is deeply notched at the sutures between the maxillaries and pre-maxillaries, and the second maxillary notch is well marked. All the bones of the upper surface of the head are deeply pitted.

The borders of the cranium are rounded as they approach the orbits; the superior temporal orifices are almost perfectly circular, the fore-and-aft diameter exceeding the transverse by only one millimetre. This effect may, in some degree, be due to distortion.

To give a more detailed account of the several elements of the skull, we take up first the basioccipital. This bone is remarkably long and straight, tapers gradually downwards, and becomes quite narrow at the distal end. It is smooth throughout, and exhibits no rugose muscular attachments, such as are sometimes seen in other members of the order. The condyle is large and nearly spherical, but with median groove distinctly marked. In size and shape it is more like that of the alligator than of the ordinary crocodile, but it is somewhat different from either. It differs from the former, in not having so long a neck distinctly marked by a constriction; and from the latter, in not having additional articular faces on each side of the condyle proper. As far as can be judged, no portion of it is formed by the exoccipitals. Below the condyle the basioccipital is perforated by two small vascular foramina; the spheno occipital canal occupies the usual place, and is very large.

The exoccipitals are large, of very great width, but rather low from above downwards. The position of the foramina which perforate these bones is peculiar; it resembles more the arrangement seen in the skull of the alligator than in that of the crocodile, but it has an additional foramen. There are, then, two small venous foramina near the condyle; while along the lateral margin of the occiput, are placed in a vertical line the foramina for the hypoglossal and pneumogastric nerves, and the internal carotid artery. The foramen for the facial nerve, etc., is situated in the usual place, and is of the usual size. The foramen magnum is heart-shaped, low, wide above, contracting below. The paroccipital processes are long and slender, and project strongly backwards.

The supra-occipital is very small. It shows to some extent on the upper surface of the skull, wedged in between the parietals. As a whole, the occiput is of remarkable shape: it is perfectly vertical, as in all other crocodilians; is remarkably high from above downwards, and is very broad at top, becoming extremely narrow distally. This latter feature is owing to the peculiar shape of the pterygoids; which, when viewed from behind, do not appear to reach the basioccipital.

The basisphenoid is so destroyed by crushing, that nothing can be said of its shape or of its foramina. It was, however, evidently very stout. The alisphenoid is large and smooth and is not ridged; the foramen ovale is small, and is not produced into an anterior notch. The suture between the alisphenoid and the pro-otic are quite distinct, but the latter is so firmly anchylosed to the quadrate that its limits are indeterminable. None of the other periotic bones are visible.

The quadrate is very long and broad. In shape it resembles the corresponding bone of the alligator, but is somewhat broader. The lower surface is divided into two unequal parts by a conspicuous ridge, which runs to within an inch of the articular surface. This surface has a more decidedly grooved or trochlear appearance than in the common crocodile; but not to the same degree as in the specimen described by Dr. Leidy. It is more like the Mississippi alligator in this respect than any living species of which we have specimens.

The pterygoids are of peculiar shape; they are long, slender, and pointed, and meeting the basioccipital on the median line in front, they project downwards and backwards. Their posterior border is very deeply emarginate, so that they seem to have no connection with each other, or with the basioccipital; instead of having the broad, plate-like appearance of these bones in recent species. The suture between these on the palatal surface is long; and the processes of the pterygoids, which bound the posterior nares, are long and stout. The posterior nares have the position which they take in the recent species; they are visible in the occipital surface, and are directed backwards as well as downwards. They are rather small, and appear to have no septum between them, but this cannot be said with any certainty.

The transpalatine is also somewhat peculiar in shape; the process which joins the pterygoid is of great length, being nearly as long as that bone. The other limbs are more normal in length. The three processes are connected at the usual angle.

The palatals are long and narrow, becoming wider anterior to the foramina. The suture with the maxillaries is rounded, and there are no forward processes as in the true crocodiles; but at the same time, these bones are not of the shape exhibited in the alligators. The palatal foramina are of immense length; they are more than one third as long as the entire bony palate. Their width is also considerable.

The maxillaries are long and very broad; the alveolar border is of about the same shape as in the crocodile, but less decidedly sinuous; and the posterior part passes in below the alveolus of the lower jaw. The convexity of the upper surface of the maxillaries is not so well marked as in the true crocodiles. It is nearly as flat as in the alligator. The palatine plates of the maxillaries are short, broad, and nearly flat, arching slightly to form the alveolus, and perforated along this border by rows of foramina.

The premaxillaries are very short; they curve strongly outwards from the notch and enclose the large anterior narial opening, which is distinctively crocodilian (as distinguished from other genera) in shape. The muzzle ends quite sharply. The palatine processes are short and convex in both directions, and the incisive foramen is heart-shaped. The alveolus is quite regular in outline; it is pitted in front for the first mandibular teeth, but there is no perforation for them.

The teeth are short, stout, compressed so as to form cutting-edges, and are somewhat obtuse. They are finely striated from base to tip. The premaxillary held four teeth; these are all broken off, but from their fangs it appears that they formed an uninterrupted row, and were subequal in size. The maxillary accommodated fifteen teeth, of which the fifth is the largest, and forms a very prominent canine. The posterior maxillary teeth are proportionately larger, and more equal than in either crocodile or alligator.

The nasals are rather broad; they send processes into the anterior narial opening, which tend to divide it, but this division was probably not complete. The limits of the prefrontals and lachrymals are so obscure as to preclude description.

The frontals are long and very narrow, though wider than in the alligator. They are smooth and flat, exhibiting no concavity on top. They expand considerably at the posterior part of the orbit, at the sutures with the post-frontals. These bones are long, stout, and curved very strongly outwards.

The mastoids are of remarkable size; they project far backward along the tympanic, and encroach largely upon the occipital region. It is partly owing to this that the occiput is so high.

The parietals are short, and very narrow between the temporal orifices, behind these they expand considerably. The orbits are large and of irregular shape.

The malar is long and rather slender.

Measurements.

M.

Length from occipital border to end of muzzle ·455

Breadth of cranium at postorbital angles ·0935

Breadth of cranium between temporal orifices ·019

Breadth of forehead between orbits ·036

Breadth of temporal orifices ·037

Fore-and-aft diameter of the same ·038

Fore-and-aft diameter of the orbits ·073

Length of face in advance of the orbits ·3055

Breadth of face outside of the fifth maxillary tooth ·176

Breadth of muzzle as formed by premaxillaries (about) ·114

Breadth of muzzle at notch for canine ·093

Length of premaxillaries to notch ·080

Estimated length of entire alveolar border ·283

Breadth of articular surface of quadrate ·059

Vertical height of occiput ·124

Vertical diameter of foramen magnum ·016

Transverse diameter of foramen magnum ·025

Vertical diameter of condyle ·023

Transverse diameter of condyle ·028

Length from palatine foramen to end of pterygoid ·120

Length of palatals ·1295

Length of palatine foramen ·142

Greatest breadth of palatine foramen ·051

Length of bony palate from incisive foramen to posterior nares ·350

The mandible is long, rather shallow, but very thick. The symphysis is very long, extending as far as the seventh tooth. The chin is quite sharp, but becomes broad, as the rami diverge quite rapidly. The alveolar border is rounded and comparatively straight, the median enlargement of the dentary is in thickness rather than in height. The two rami diverge at an angle which is more open than in the true crocodiles, and less so than in the alligator. The mandibular foramen is smaller than in the latter genus, but corresponds with it in position; it has its long diameter parallel with, and not oblique to, the alveolus. The mandibular fossa is extremely large and deep. The splenial is long and stout, ends obtusely, and does not reach the symphysis. The articular cavity is broad from side to side, but rather shallow; it is not divided into two distinct facets as in the alligator. The post-glenoid process is very stout; it projects but slightly upwards, less so than in either crocodile or alligator. It has no median ridge as in the former genus, and is tuberous at the end.

The teeth are much like those of the upper jaw, but are somewhat sharper and more conical; they are not recurved, and have no distinct constriction, as is found in most of the recent species. There are about eighteen teeth to each ramus; the first is large and sharp, and is followed by two small ones and then by the canine. The remaining teeth do not exhibit any great differences of size.

Measurements.

M.

Length of rami (straight) ·618

Width of lower jaw outside of glenoid cavities ·316

Length of symphysis ·110

Width of jaw at second enlargement ·057

Depth at oval foramen ·073

Greatest width at symphysis ·110

Space occupied by teeth ·340

Breadth of glenoid cavity ·072

Length of post-glenoid ·065

Vertebræ.

Cervicals.—The centra are long and nearly cylindrical, expanding slightly near the anterior face. The hypapophyses are short and broad; they project forwards, and are somewhat compressed. The vascular foramina correspond in size and position to those of the alligator. The neural canal is small and subcircular, having comparatively short but very stout neurapophyses, which are perforated behind the diapophyses. These are short and stout, and are developed from the neurapophyses alone. The zygapophyses are long and flat; the anterior ones project almost vertically. The neural spines are stout, and of greater antero-posterior extent than in the alligator.

Dorsals.—A few of the anterior dorsals have strong hypapophyses. The centra of all are long and stout. The neural canal is smaller than in the cervical region; it has strong neurapophyses, which develop long and depressed diapophyses. The neural spines are low and broad. The zygapophyses are developed on the laminæ, and do not project upwards as in the cervical region.

The lumbars are much like the dorsals, except that they are longer, and have very large and depressed diapophyses, which project somewhat upwards.

Measurements.

M.

Length of centrum of a posterior cervical ·049

Height of neural canal ·015

Fore-and-aft diameter of neural spine ·023

Length of diapophysis ·023

Dorsal.

M.

Length of centrum ·0555

Length of hypapophysis ·018

Fore-and-aft diameter of neural spine ·030

Lumbar.

M.

Length of centrum ·057

Fore-and-aft diameter of neural spine ·0285

Diameter of diapophysis ·029

The dermal scutes are long ellipses, deeply pitted on one side, but without a trace of a keel. Their edges show no signs of sutural union.

The bones described indicate a reptile about fifteen feet long. They were found near Smith's Fork, Wyoming.

Crocodilus parvus, sp. nov.

A small reptile represented by sixteen vertebræ and a portion of the pelvis.

The cervicals have short centra, with very deep articular cups and hemispherical heads: the latter have a prominent rim around the base. The hypapophyses are short, stout, and very nearly vertical in direction; the sides of the centra are channelled by a deep vertebraterial canal; the parapophyses are developed very low down, and are very prominent. The diapophyses are developed partly from the centra and partly from the neurapophyses; they are stout and very short. The facets for the ribs are developed in the usual place. The neural arch is rather high, and forms a small, narrow canal; the proportions of the neurapophyses and neural spines are about as in Alligator mississippiensis, but the zygapophyses are not so prominent.

The dorsals show a considerable increase in size over the cervicals; the centra become elongate, and the articular cups shallower and transversely oval. Several of the anterior vertebræ retain large hypapophyses. In the dorsal region the diapophyses are developed from the neurapophyses alone, and at a considerable height above the neuro-central suture; they are long and depressed. The zygapophyses and neural spines present no peculiarities of structure.

In the lumbar region the vertebræ regain their cylindrical form, and become still more elongate; but the cups are comparatively shallow and the heads low; there is no distinct shoulder. The neural canal is small, with low, broad neurapophyses, from which are developed very broad and depressed diapophyses, which project outwards in a horizontal plane. The neural spines are thick, and broad antero-posteriorly, but are so broken that their height cannot be determined.

In both dorsal and lumbar region the neurapophyses are deeply notched on their posterior edges, and close to the neuro-central sutures, for the passage of the spinal nerves.

The ilium is very high compared with most crocodiles; its vertical diameter is considerably more than half of the antero-posterior diameter. The construction of the bone is very much like that in the modern Crocodilia; but the suprailiac border is more regular, and the anterior tuberosity is not so thick or so much everted as in these forms. The acetabulum is small, shallow, directed downwards and outwards, and situated considerably forward of the median line. The iliac surface is smooth, but very irregular, being very deeply concave above the acetabulum, and convex behind it. The sacral surface is rugose, rises above the sacrum, and shows attachments for two sacral vertebræ. Thus the construction of the entire pelvis shows but very little variation from the modern type.

Measurements.

M.

Antero-posterior diameter of ilium ·095

Vertical diameter of ilium ·061

Antero-posterior diameter of acetabulum ·029

Length of centrum of third (?) cervical vertebra (from edge of cup to tip of ball) ·032

Height of neural canal, third (?) cervical ·0095

Length of hypapophysis of third cervical ·0085

Length of centrum of a posterior dorsal ·040

Length of diapophysis (about) ·032

Length of centrum of a lumbar ·046

Vertical diameter of a lumbar ·029

Breadth of neurapophysis of a lumbar ·027

Crocodilus heterodon, Cope.

Alligator heterodon, Proc. American Philosophical Society, 1872, p. 544.

Represented by a single tooth from the posterior part of the mandibular series. It agrees very closely with the corresponding tooth of Alligator mississippiensis, but presents some differences. The crown is very low, obtuse, and finely striate; it is compressed and shows a cutting-edge. Its longest diameter is fore-and-aft; both this and the transverse diameter are proportionately greater than in the modern species. The constriction of the neck is very decided; the fang is large, and especially thick.

Measurements.

M.

Fore-and-aft diameter of crown ·010

Transverse diameter of crown ·007

Vertical height of crown

Crocodilus clavis, Cope.

U. S. Geol. Survey of Terrs. 1872, p. 612.

This species is indicated by the remains of a crocodilian larger than Crocodilus Elliotii, but very different from it. The pitting of the dermal scutes, and of all the cranial bones, is deeper than in any other of the Bridger crocodilians in our possession, and very strongly resembles that in Alligator mississippiensis.

The mandible has a long symphysis, and the alveolar border shows the sudden deep depression just behind it, which is so marked in the ordinary alligator. The rami, however, do not diverge at such an open angle. The jaw is very stout, and is deeper and thinner than in Crocodilus Elliotii; and is of about the same proportionate thickness, but deeper than in the alligator. The splenial ends obtusely, and approaches the symphysis somewhat more closely than in the alligator, but does not enter into its formation. The teeth are stout and obtusely conical, not striate, and implanted very close together; they exhibit about the same alternation of size as do those of the alligator. What appears to be the canine is small; it is preceded by a much smaller, and succeeded by a slightly smaller tooth. The dermal scutes are quadrate in shape; they are deeply and irregularly pitted, and have a low but distinct longitudinal keel; they are suturally united by their lateral borders.

CHELONIA.

The Bridger beds have yielded a great abundance and variety of land, marsh, and fresh-water chelonians. They present much variation of form and size; and, owing to the great pressure in the strata, scarcely any two specimens of the same species are alike; occasionally, however, an undistorted specimen is obtained.

The most abundant remains of turtles are those of a species of the recent genus Emys, which are found in all the beds, and are, perhaps, the commonest fossils of the basin. The genera Trionyx and Hadrianus (Testudo, Leidy) follow next in order.

HADRIANUS, Cope.

Resembles Testudo in form, but has two analscuta, as have most Emydidæ.

Hadrianus allabiatus, Cope.

U. S. Geol. Survey of Territories, 1872, p. 630.

Represented by two ungual phalanges which resemble the claws of Testudo, but are proportionately shorter, broader, and not so pointed at the ends. They are oval in section, both longitudinally and transversely; the articular faces are deep and subinferior.

From Henry's Fork.

Hadrianus octonarius, Cope.

Geol. Survey of Terrs., 1872. p 630.

Represented by the right humerus of a large individual.

This bone is intermediate in character between the corresponding bones of Testudo and Emys. The head is longer and narrower than in the former genus; it is implanted on a short, distinct neck, and is strongly convex in both directions. The articular surface extends to the base of the internal tuberosity. The tuberosities, especially the external one, are very large and thick. The external rises above the head, and resembles that of Emys in shape. The fossa separating the tuberosities is comparatively narrow. The shaft is much like that of Emys, but is somewhat more strongly curved; not so much, however, as in Testudo, and the distal end is not so much flattened. The anconeal fossa is wider and deeper than in either genus. The trochlea is broad, and divided by an obscure groove into a small and convex radial face, and a nearly plane (transversely) ulnar face. The condyles are prominent and rugose.

Found near Dry Creek.

EMYS.

Emys wyomingensis, Leidy.

Cont. to Ex. Vert. Fauna, p. 140.

Represented by six perfect shells from Cottonwood and Dry Creeks, and Henry's Fork, and a shoulder girdle from Cottonwood Creek, which is provisionally referred to this species.

This species, when adult, was upward of a foot in length, and in the composition of the shell, number of scutes, etc., resembled the living species of the genus.

The Shoulder Girdle has the form characteristic of the Emydidæ, but presents some peculiarities. The scapula is short, not being as long as the coracoid, and is not so straight or cylindrical as is usual in Emys; but is compressed, and somewhat recurved, and tapers towards the upper extremity, which is marked by a smooth rounded facet for the cartilaginous supra-scapula. The precoracoid is long, sub-cylindrical at its origin, but becomes broad and flat distally; at the extremity it is longitudinally striated. The proximal end is flexed forward at an obtuse angle, and is expanded to form the long suture with the scapula and coracoid. It contributes considerably to the glenoid cavity. The coracoid is remarkably long, and after leaving the sutures with the other elements of the girdle, becomes very broad and thin. This is most marked distally. The anterior border is thickened, and the bone is slightly concave on both surfaces.

The three parts of the shoulder girdle are united by distinct bony symphyses; they differ from some of the modern species, in which there is only a ligamentous union between the coracoid and the other parts. It presents a further difference in the long and slender neck which is formed by the flexing of the precoracoid, and in the shallow and subcircular glenoid cavity. The short scapula and long coracoid would indicate a species with a low, broad carapace, as was probably the case in E. wyomingensis, though much difficulty has been experienced in determining this point, owing to the distortion of the specimens.

Measurements.

M.

Length of neck to point between coracoid and precoracoid ·0345

Length of coracoid (about) ·133

Length of precoracoid ·114

Length of scapula (about) ·126

BAENA, Leidy.

Baena undata, Leidy.

Cont. to Ex. Vert. Fauna, p. 160.

Three nearly perfect shells of different ages, from Dry and Cottonwood Creeks.

TRIONYX, Geoffr.

Trionyx uintaensis, Leidy.

Cont. to Ex. Vert. Fauna, p 176.

Cottonwood Creek.

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