CHAPTER XII. Evidences of the Animal's Habits From Its Remains

EVIDENCES OF THE ANIMAL'S HABITS FROM ITS REMAINS

Such are the more remarkable characters of the bones in a type of animal life which was more anomalous than any other which peopled the earth in the Secondary Epoch of geological time. Its skeleton in different parts resembles Reptiles, Birds, and Mammals; with modifications and combinations so singular that they might have been deemed impossible if Nature's power of varying the skeleton could be limited. Since Ornithosaurs were provided with wings, we may believe the animals to some extent to have resembled birds in habit. Their modes of progression were more varied, for the structures indicate an equal capacity for movement on land as a biped, or as a quadruped, with movement in the air. There is little evidence to support the idea that they were usually aquatic animals. The majority of birds which frequent the water have their bodies stored with fat and the bones of their extremities filled with marrow. And a bird's marrow bones are stouter and stronger than those which are filled with air. There are few, if any, bones of Pterodactyles so thick as to suggest the conclusion that they contained marrow, and the bones of the extremities appear to have been constructed on the lightest type found among terrestrial birds. Their thinness, except in a few specimens from the Wealden rocks, is marvellous; and all the later Pterodactyles show the arrangement, as in birds, by which air from the lungs is conveyed to the principal bones. No Pterodactyle has shown any trace of the web-footed condition seen in birds which swim on the water, unless the diverging bones of the hind foot in Rhamphorhynchus supports that inference. The bones of the hind foot are relatively small, and if it were not that a bird stands easily upon one foot, might be considered scarcely adequate to support the animal in the position which terrestrial birds usually occupy. Yet, as compared with the length and breadth of the foot in an Ostrich, the toes of an Ornithosaur are seen to be ample for support. These facts appear to discourage the idea that the animals were equally at home on land and water, and in air.

Some light may be thrown upon the animal's habits by the geological circumstances under which the remains are found. The Pterodactyle named Dimorphodon, from the Lias of the south of England, is associated with evidences of terrestrial land animals, the best known of which is Scelidosaurus, an armoured Dinosaur adapted by its limbs for progression on land. And the Pterodactyle Campylognathus, from the Lias of Whitby, is associated with trunks of coniferous trees and remains of Insects. So that the occurrence of Pterodactyles in a marine stratum is not inconsistent with their having been transported by streams from off the old land surface of the Lias, on which coniferous trees grew and Dinosaurs lived.

Similar considerations apply to the occurrence of the Rhamphocephalus in the Stonesfield Slate of England. The deposit is not only formed in shallow water, but contains terrestrial Insects, a variety of land plants, and many Reptiles and other animals which lived upon land. The specimens from the Purbeck beds, again, are in strata which yield a multitude of the spoils of a nearly adjacent land surface; while the numerous remains found in the marine Solenhofen Slate in Germany are similarly associated with abundant evidences of varied types of terrestrial life. The evidence grows in force from its cumulative character. The Wealden beds, which yield many terrestrial reptiles and so much evidence of terrestrial vegetation, and shallow-water conditions of disposition, have afforded important Pterodactyle remains from the Isle of Wight and Sussex.

The chief English deposit in which these fossils are found, the Upper Greensand, has afforded thousands of bones, battered and broken on a shore, where they have lain in little associated groups of remains, often becoming overgrown with small marine shells. Side by side with them are found bones of true terrestrial Lizards and Crocodiles of the type of the Gavial of the Indian rivers, many terrestrial Dinosaurs, and other evidences of land life, including fossil resins, such as are met with in the form of amber or copal at the present day.

The great bones of Pterodactyles found in the Chalk of Kent, near Rochester, became entombed, beyond question, far from a land surface. There is nothing to show whether the animals died on land and were drifted out to sea like the timber which is found water-logged and sunken after being drilled by the ship-worm (Teredo) of that epoch. Seeing the power of flight which the animal possessed, storms may have struck down travellers from time to time, when far from land.

Evidence of habit of another kind may be found in their teeth. They are brightly enamelled, sharp, formidable; and are frequently long, overlapping the sides of the jaws. They are organs which are often better adapted for grasping than for tearing, as may be seen in the inclined teeth of Rhamphocephalus of the Stonesfield Slate; and better adapted for killing than tearing, from their piercing forms and cutting edges, in genera like Ornithocheirus of the Greensand. The manner in which the teeth were implanted and carried is better paralleled by the fish-eating crocodile of Indian rivers than by the flesh-eating crocodiles, or Muggers, which live indifferently in rivers and the sea. As the Kingfisher finds its food (see Fig. 20) from the surface of the water without being in the common sense of the term a water bird, so some Pterodactyles may have fed on fish, for which their teeth are well adapted, both in the stream and by the shore.

A Pterodactyle's teeth vary a good deal in appearance. The few large teeth in the front of the jaw in Dimorphodon, associated with the many small vertical teeth placed further backward, suggest that the taking of food may have been a process requiring leisure, since the hinder teeth adapted to mincing the animal's meat are extremely small. The way in which the teeth are shaped and arranged differs with the genera. In Pterodactylus they are short and broad and few, placed for the most part towards the front of the jaws. Their lancet-shaped form indicates a shear-like action adapted to dividing flesh. In the associated genus Rhamphorhynchus the teeth are absent from the extremity of the jaw, are slender, pointed, spaced far apart, and extend far backward. When the jaws of the Rhamphorhynchus are brought together there is always a gap between them in front, which has led to belief that the teeth were replaced by some kind of horny armature which has perished. In the long-nosed English type of Ornithocheirus the jaws are compressed together, so that the teeth of the opposite sides are parallel to each other, with the margins well filled with teeth, which are never in close contact, though occasionally closer and larger in front, in some of the forms with thick truncated snouts.

It is not the least interesting circumstance of the dentition of Pterodactyles that, associated in the same deposits with these most recent genera with teeth powerfully developed, there is a genus named Ornithostoma from the resemblance of its mouth to that of a bird in being entirely devoid of teeth. It is scarcely possible to distinguish the remains of the toothed and toothless skeletons except in the dentary character of the jaws. There is no evidence that the toothless types ever possessed a tooth of any sort. They were first found in fragments in England in the Cambridge Greensand, but were afterwards met with in great abundance in the Chalk of Kansas, where the same animals were named Pteranodon. A jaw so entirely bird-like suggests that the digestive organs of Pterodactyles may in such toothless forms at least have been characterised by a gizzard, which is so distinctive of Birds. The absence of teeth in the Great Ant-eater and some other allied Mammals has transferred the function which teeth usually perform to the stomach, one part of which becomes greatly thickened and muscular, adapting itself to the work which it has to perform. It is probable that the gizzard may be developed in relation to the necessities which food creates, since even Trout, feeding on the shell-fish in some Irish lochs, acquire such a thickened muscular stomach, and a like modification is recorded in other fishes as produced by food.

Closely connected with an animal's habits is the protection to the body which is afforded by the skin. In Pterodactyles the evidence of the condition of the skin is scanty, and mostly negative. Sometimes the dense, smooth texture of the jaw bones indicates a covering like the skin of a Lizard or the hinder part of the jaw of a Bird. Some jaws from the Cambridge Greensand have the bone channeled over its surface by minute blood vessels which have impressed themselves into the bone more easily than into its covering. Thus in the species of Ornithocheirus distinguished as microdon the palate is absolutely smooth, while in the species named machaerorhynchus it is marked by parallel impressed vascular grooves which diverge from the median line. This condition clearly indicates a difference in the covering of the bone, and that in the latter species the covering had fewer blood vessels and more horny protection than in the other. The tissue may not have been of firmer consistence than in the palate of Mammals. The extremity of the beak is often as full of blood vessels as the jaw of a Turtle or Crocodile.

COVERING OF THE BODY

There is no trace even in specimens from the Solenhofen or Stonesfield Slate of any covering to the body. There are no specimens preserved like mummies, and although the substance of the wings is found there is no trace of fur or feathers, bones, or scales on the skin. The only example in which there is even an appearance suggesting feathers is in the beautiful Scaphognathus at Bonn, and upon portions of the wing membrane of that specimen are preserved a very few small short and apparently tubular bodies, which have a suggestive resemblance to the quills of small undeveloped feathers. Such evidences have been diligently sought for. Professor Marsh, after examining the wing membranes of his specimen of Rhamphorhynchus from Solenhofen, stated that the wings were partially folded and naturally contracted into folds, and that the surface of the tissue is marked by delicate striae, which might easily be taken at first sight for a thin coating of hair. Closer investigation proved the markings to be minute wrinkles on the under surface of the wing membrane. This negative evidence has considerable value, because the Solenhofen Slate has preserved in the two known examples of the bird Archaeopteryx beautiful details of the structure of the larger feathers concerned in flight. It has preserved many structures far more delicate. There is, therefore, reason for believing that if the skin had possessed any covering like one of those found in existing vertebrate animals, it could scarcely have escaped detection in the numerous undisturbed skeletons of Pterodactyles which have been examined.

The absence of a recognisable covering to the skin in a fossil state cannot be accepted as conclusive evidence of the temperature, habits, or affinities of the animal. Although Mammalia are almost entirely clothed with dense hair, which has never been found in a recognisable condition in a fossil state in any specimen of Tertiary age, one entire order, the Cetacea, show in the smooth hairless skins of Whales and Porpoises that the class may part with the typical characteristic covering without loss of temperature and without intelligible cause. That the absence of hair is not due to the aquatic conditions of rivers or sea is proved by other marine Mammals, like Seals, having the skin clothed with a dense growth of hair, which is not surpassed in any other order. The fineness of the growth of hair in Man gives a superficial appearance of the skin being imperfectly clothed, and a similar skin in a fossil state might give the impression that it was devoid of hair. There are many Mammals in which the skin is scantily clothed with hair as the animal grows old. Neither the Elephant nor the Armadillo in a fossil state would be likely to have the hair preserved, for the growth is thin on the bony shields of the living Armadilloes. Yet the difficulty need be no more inherent in the nature of hair than in that of feathers, since the hair of the Mammoth and Rhinoceros has been completely preserved upon their skins in the tundras of Siberia, densely clothing the body. This may go to show that the Pterodactyle possessed a thin covering of hair, or, more probably, that hair was absent. Since Reptiles are equally variable in the clothing of the skin with bony or horny plates, and in sometimes having no such protection, it may not appear singular that the skin in Ornithosaurs has hitherto given no evidence of a covering. From analogy a covering might have been expected; feathers of Birds and hair of Mammals are non-conducting coverings suited to arrest the loss of heat.

With the evidence, such as it is, of resemblance of Ornithosaurs to Birds in some features of respiration and flight, a covering to the skin might have been expected. Yet the covering may not be necessary to a high temperature of the blood. Since Dr. John Davy made his observations it has been known that the temperature of the Tunny, above 90 deg. Fahrenheit, is as warm as the African scaly ant-eater named the Pangolin, which has the body more amply protected by its covering. This illustration also shows that hot blood may be produced without a four-celled heart, with which it is usually associated, and that even if the skin in Pterodactyles was absolutely naked an active life and an abundant supply of blood could have given the animal a high temperature.

The circumstance that in several individuals the substance of the wing membrane is preserved would appear to indicate either that it was exceptionally stout when there would have been small chance of resisting decomposition, or that its preservation is due to a covering which once existed of fur or down or other clothing substance, which has proved more durable than the skin itself.

SHOWING THE SKULL, NECK, BACK AND SOME OF THE LONGER BONES OF THE SKELETON

From a slab in the British Museum (Natural History)]