Dragons of the Air: An Account of Extinct Flying Reptiles

by H. G. Seeley

Compared with the same aspect of the skull of Rhamphorhynchus

The third opening in the side of the head, counting from before backward, is the orbit of the eye. In this vacuity is often seen the sclerotic circle of overlapping bones formed in the external membrane of the eye, like those in nocturnal birds and some reptiles. The eye hole varies in form from an inverted pear-shape to an oblique or transverse oval, or a nearly circular outline. It is margined by the frontal bone above; the tear bone or lachrymal, and the malar or cheek bone in front; while the bones behind appear to be the quadrato-jugal and post-frontal bones, though the bones about the eye are somewhat differently arranged in different genera.

The eyes were frequently, if not always, in contact with the anterior walls of the brain case, as in many birds, and are always far back in the side of the head. In Dimorphodon they are in front of the articulation of the lower jaw; in Rhamphorhynchus, above that articulation; while in Ornithostoma they are behind the articulation for the jaw. This change is governed by the position of the quadrate bone, which is vertical in the Lias genus, inclined obliquely forward in the fossils from the Oolites, and so much inclined in the Chalk fossil that the small orbit is thrown relatively further back.

Thus far the chief difference in the Pterodactyle skull from that of a bird is in the way in which the malar arch is prolonged backward on each side. It is a slender bar of bone in birds, without contributing ascending processes to border vacuities in the side of the face, while in these fossil animals the lateral openings are partly separated by the ascending processes of these bones. This divergence from birds, in the malar bone entering the orbit of the eye is approximated to among reptiles and mammals, though the conditions, and perhaps the presence of a bone like the post-orbital bone, are paralleled only among Reptiles. The Pterodactyles differ among themselves enough for the head to make a near approach to Reptiles in Dimorphodon, and to Birds in Pterodactylus. In the Ground Hornbill and the Shoebill the lachrymal bones in front of the orbits of the eyes grow down to meet the malar bars without uniting with them. The post-frontal region also is prolonged downward almost as far as the malar bar, as though to show that a bird might have its orbital circle formed in the same way and by the same bones as in Pterodactylus. Cretaceous Ornithosaurs sometimes differ from birds apparently in admitting the quadrato-jugal bone into the orbit. It then becomes an expanded plate, instead of a slender bar as in all birds.

THE TEMPORAL FOSSA

A fourth vacuity is known as the temporal fossa. When the skull of such a mammal as a Rabbit, or Sheep, is seen from above, there is a vacuity behind the orbits for the eyes, which in life is occupied by the muscles which work the lower jaw. It is made by the malar bone extending from the back of the orbit and the process of bone, called the zygomatic process, extending forward from the articulation of the jaw, which arches out to meet the malar bone.

In birds there is no conspicuous temporal fossa, because the malar bar is a slender rod of bone in a line with the lower end of the quadrate bone.

Reptile skulls have sometimes one temporal vacuity on each side, as among tortoises, formed by a single lateral bar. These vacuities, which correspond to those of mammals in position, are seen from the top of the head, as lateral vacuities behind the orbits of the eyes, and are termed superior temporal vacuities. In addition to these there is often in other reptiles a lateral opening behind the eye, termed the inferior temporal vacuity, seen in Crocodiles, in Hatteria, and in Lizards; and in such skulls there are two temporal bars seen in side view, distinguished as superior and inferior. The superior arch always includes the squamosal bone, which is at the back of the single bar in mammals. The lower arch includes the malar bone, which is in front in the single arch of mammals. The circumstance that both these arches are connected with the quadrate bone makes the double temporal arch eminently reptilian.

In Ornithosaurs the lateral temporal vacuity varies from a typically reptilian condition to one which, without becoming avian, approaches the bird type. In skulls from the Lias, Dimorphodon and Campylognathus, there is a close parallel to the living New Zealand reptile Hatteria, in the vertical position of the quadrate bone and in the large size of the vacuity behind and below the eye, which extends nearly the height of the skull. In the species of the genus Pterodactylus, the forward inclination of the quadrate bone recalls the Curlew, Snipe, and other birds. The back of the head is rounded, and the squamosal bone, which appears to enter into the wall of the brain case as in birds and mammals, is produced more outward than in birds, but less than in mammals, so as to contribute a little to the arch which is in the position of the post-frontal bone of reptiles. It is triangular, and stretches from the outer angle of the frontal bone at the back of the orbit to the squamosal behind, where it also meets the quadrate bone. Its third lower branch meets the quadratojugal, which rests upon the front of the quadrate bone, as in Iguanodon, and is unlike Dimorphodon in its connexions. In that genus the supra-temporal bone, or post-orbital bone, appears to rest upon the post-frontal and connect it with the quadrato-jugal. In Dimorphodon the malar bone is entirely removed from the quadrate, but in Pterodactylus it meets its articular end. Between the post-frontal bone above and the quadrato-jugal bone below is a small lunate opening, which represents the lateral temporal vacuity; and so far, this is a reptilian character. But if the thin post-frontal bone were absorbed, Pterodactylus would resemble birds. There is no evidence that the quadrate bone is free in any Ornithosaurs, as it is in all birds, while in Dimorphodon it unites by suture with the squamosal bone. In Ornithostoma the lateral temporal vacuity is little more than a slit between the quadrate bone below, the quadrato-jugal in front, and what may be the post-frontal bone behind (see [Fig. 2, p. 12]).