Dragons of the Air: An Account of Extinct Flying Reptiles

by H. G. Seeley

It would therefore appear from the vital community of structures with Birds, that Pterodactyles and Birds are two parallel groups, which may be regarded as ancient divergent forks of the same branch of animal life, which became distinguished from each other by acquiring the different condition of the skin, and the structures which were developed in consequence of the bony skeleton ministering to flight in different ways; and with different habit of terrestrial progression, this extinct group of animals acquired some modifications of the skeleton which Birds have not shown. There is nothing to suggest that Pterodactyles are a branch from Birds, but their relation to Birds is much closer, so far as the skeleton goes, than is their relation with the flightless Dinosaurs, with which Birds and Pterodactyles have many characters in common.

On the theory of elimination of character which I have used to account for the disappearance of some Mammalian characters from the Pterodactyle, that loss is seen chiefly in the removal of the parts which have left a Reptilian articulation of the lower jaw with the skull, and the articulation of the vertebræ throughout the vertebral column by a modified cup-and-ball form of joint. The furculum of the Bird is always absent from the Pterodactyle. No specimen has shown recognisable clavicles or collar-bones. Judged by the standard of existing life, Pterodactyles belong to the same group as Birds, on the evidence of brain and lungs, but they belong to a different group on account of the dissimilar modifications of the skeleton and apparent absence of feathers from the skin.

The most impressive facts in the Pterodactyle skeleton, in view of these affinities, are the structures which it has in common with Reptiles. Some structures are fundamental, like the cup-and-ball articulation of the vertebræ, which is never found in birds or mammals. Although not quite identical with the condition in any Reptile, this structure is approximately Lizard-like or Crocodile-like in the cup-and-ball character. It shows that the deepest-seated part of the skeleton is Reptile-like, though it may not be more Reptilian than is the vertebral column of a Mammal, if comparison is made between Mammals and extinct groups of animals known as Reptiles, such as Dinosaurs and Theriodontia.

The orders of animals which have been included under the name Reptilia comprise such different structural conditions of the parts of the skeleton which may be termed reptilian in Ornithosaurs, that there is good reason for regarding the cup-and-ball articulation as quite a distinctive Reptilian specialisation, in the same sense that the saddle-shaped articulation between the bodies of adjacent vertebræ in a bird is an Avian specialisation. From the theoretical point of view the Ornithosaur acquired its Reptilian characters simultaneously with its Avian and Mammalian characters.

There is nothing in the structure of the skeleton of the Dinosauria, to which Ornithosaurs approximate in several parts of the body, which would help to explain the cup-and-ball articulation of the backbone, if the Flying Reptile were supposed to be an offshoot from the carnivorous Dinosaurs.

The elimination of Reptile characters from so much of the skeleton, and the substitution for them of the characters of Birds and Mammals, would be of exceptional interest if there had been any ground for regarding the flying animal as more nearly related to a Reptile than to a Bird. But if the evidence from the form of the brain and nature of the pneumatic organs seen in the limb bones accounts for the Avian features of the skeleton, the Reptilian condition of the vertebral column helps to show a capacity for variation, and that the fixity of type and structure, which the skeleton of the modern Bird has attained, is not necessarily limited to or associated with the vital organs of Birds.

The variation of the cup-and-ball articulation in the neck of a Chelonian, which makes the third vertebra cupped behind, the fourth bi-convex, the fifth cupped in front, and the sixth flattened behind, shows that too much importance may be attached to the mode of union of these bones in Serpents, Crocodiles, and those Lizards which have the cup in front; for while in Lizards the anterior cup, oblique and depressed, is found in most of its groups, the Geckos show no trace of the cup-and-ball structure, and in that respect resemble the Hatteria of New Zealand.

If, therefore, the cup-and-ball articulation of vertebræ in Ornithosauria has any significance as a mark of affinity to Reptiles, it could only be in approximation to those living Reptiles which possess the same character, and would have it on the hypothesis that both have preserved the structure by descent from an earlier type of animal. This hypothesis is negatived by the fact that the cup-and-ball articulation is unknown in the older fossil Reptiles.

Although the articulation for the lower jaw with the skull in Ornithosaurs is only to be paralleled among Reptiles, the structure is adapted to a brain case which is practically indistinguishable from that of a Bird, except for the postorbital arch.

The hypothesis of descent, therefore, becomes impossible, in any intelligible form, in explanation of distinctive character of the skeleton. The hypothesis of elimination may also seem to be insufficient, unless the potential capacity for new development be recognised as concurrent, and as capable of modifying each region of the skeleton, or hard parts of the animal, in the same way that the soft organs may be modified. From which we infer that all structures, which distinguish the several grades of organisation in modern classifications, soft parts and hard parts alike, may come into existence together, in so far as they are compatible with each other, in any class or ordinal division of animals.