The upper figures show the side and back of a dorsal vertebra of Ornithocheirus compared with corresponding views of the side and back of a dorsal vertebra of a Crocodile
There are two remarkable modifications of the early dorsal vertebræ in some of the Cretaceous Pterodactyles. First, in the genus Ornithodesmus from the Weald the early dorsal vertebræ are blended together into a continuous mass, like that which is found in the corresponding region of the living Frigate-bird, only more consolidated, and similar to that consolidated structure found behind the dorsal vertebræ, known as the sacrum, made by the blending of the vertebræ into a solid mass which supports the hip bones. Secondly, in some of the Cretaceous genera of Pterodactyles of Europe and America the vertebræ in the front part of the back are similarly blended, but their union is less complete; and in genera Ornithocheirus and Ornithostoma—the former chiefly English, the latter chiefly American—the sides of the neural spines are flattened to form an oval articular surface on each side, which gives attachment to the flattened ends of their shoulder-blade bones named the scapulæ. This condition is found in no other animals. Three vertebræ appear to have their neural arches thus united together. The structure so formed may be named the notarium to distinguish it from the sacrum.
SACRUM
For some mysterious reason the part of the backbone which lies between the bones of the hips and supports them is termed the sacrum. Among living reptiles the number of vertebræ in this region is usually two, as in lizards and crocodiles. There are other groups of fossil reptiles in which the number of sacral vertebræ is in some cases less and in other cases more. There is, perhaps, no group in which the sacrum makes a nearer approach to that of birds than is found among these Pterodactyles, although there are more sacral vertebræ in some Dinosaurs. In birds the sacral vertebræ number from five to twenty-two. In bats the number is from five to six. In some Solenhofen species, such as Pterodactylus dubius and P. Kochi and P. grandipelvis, the number is usually five or six. The vertebræ are completely blended. The pneumatic foramina in the sacrum, so far as they have been observed, are on the under sides of the transverse processes; while in the corresponding notarial structure in the shoulder girdle the foramina are in front of the transverse processes. Almost any placental mammal in which the vertebræ of the sacral region are anchylosed together has a similar sacrum, which differs from that of birds in the more complete individuality of the constituent bones remaining evident. The transverse processes in front of the sacrum are wider than in its hinder part; so that the pelvic bones which are attached to it converge as they extend backward, as among mammals. The bodies of the vertebræ forming the sacrum are similar in length to those of the back. Each transverse process is given off opposite the body of its own vertebra, but from a lower lateral position than in the region of the back, in which the vertebræ are free.
FIG. 27. SACRUM OF RHAMPHORHYNCHUS
Showing the complete blending of the vertebræ and ribs as in a bird, with the well-defined Iliac bones, produced chiefly in front of the acetabulum for the head of the femur.
The hip bones are closely united with the sacrum by bony union, and rarely appear to come away from the sacral vertebræ, as among mammals and reptiles, though this happens with the Lias Pterodactyles. In the Stonesfield Slate and Solenhofen Slate the slender transverse processes from the vertebræ blend with the ilium of the hip girdle, and form a series of transverse foramina on each side of the bodies of the vertebræ. In the Cambridge Greensand genera the part of the ilium above the acetabulum for the articular head of the femur appears to be always broken away, so that the relation of the sacrum to the pelvis has not been observed. This character is no mark of affinity, but only shows that ossification obliterated sutures among these animals in the same way as among birds.
The great difference between the sacrum of a Pterodactyle and that of a bird has been rendered intelligible by the excellent discussion of the sacral region in birds made by Professor Huxley. He showed that it is only the middle part of the sacrum of a chicken which corresponds to the true sacrum of a reptile, and comprises the five shortest of the vertebræ; while the four in front correspond to those of the lower part of the back, which either bear no ribs or very short ribs, and are known as the lumbar region in mammals, so that the lower part of the back becomes blended with the sacrum, and thus reduces the number of dorsal vertebræ. Similarly the five vertebræ which follow the true sacral vertebræ are originally part of the tail, and have been blended with the other vertebræ in front, in consequence of the extension along them of the bird's hip bones. This interpretation helps to account for the great length of the sacrum in many birds, and also explains in part the singular shortness of the tail in existing birds. The Ornithosaur sacrum has neither the lumbar nor the caudal portions of the sacrum of a bird.