- Key People:
- Étienne de La Ville-sur-Illon, comte de Lacépède
- Related Topics:
- dinosaur
- lepidosaurian
- anapsid
- Synaptosauria
- Synapsida
Arboreal animals possess groups of anatomical features that help them cling to branches and other substrates. The most common clinging structures in vertebrates are claws; they seem to be the only arboreal adaptations of some lizards, such as the common iguana (I. iguana). Similar structures appear in many geckos (family Gekkonidae), in the anoles (Anolis; family Iguanidae), and in some skinks (family Scincidae).
Other adaptations for climbing include footpads. Pads on the feet consist of wide plates or scales under the fingers and toes. The outer layer of each scale is composed of numerous microscopic hooks formed by the free, bent tips of cells. These minute hooks can catch in the slightest irregularities of a surface, and they enable geckos to run up apparently smooth walls and even upside down on plaster ceilings. Because the hooklike cells are bent downward and to the rear, a gecko must curl its pads upward to disengage them. Thus, when walking or running up a tree or wall, a gecko must curl and uncurl its pad surface with every step.
The giant Solomon Islands skink (Corucia), true chameleons (Chamaeleonidae), arboreal vipers, boas, and pythons use prehensile tails—that is, tails that are capable of supporting most of the weight of the animal or are used habitually for grasping—for clinging to their aerial supports. Still, true chameleons rely mainly on the tonglike arrangement of the digits in their feet. The toes of each foot are united into two opposed bundles—three on the inside and two on the outside of the forefoot, and two on the inside and three on the outside of the hind foot.
Slender vine snakes of several genera of the family Colubridae are capable of extending half the body length in a horizontal plane without support; they do so habitually in bridging the gap between tree branches. Most snakes can reach across an open space. However, all except the vine snakes can extend only a short distance, and that portion of the body invariably sags like a cable. In contrast, the vine snakes bridge an open space like an I-beam. This ability is based partly on their reduced body weight and partly on their deepened and strengthened vertebrae.
Swimming
In water, of course, limb movements—whether bipedal or quadrupedal—that work well in terrestrial environments are not very effective. Aquatic reptiles, with some exceptions, use the same means of propulsion as do fish—that is, lateral undulations of the rear half of the body and tail. Crocodiles and aquatic lizards, such as some monitors (family Varanidae) and the marine iguana (Amblyrhynchus cristatus), undulate their bodies and tails from side to side while holding the limbs against the body. The ancient mesosaurs (order Mesosauria) and ichthyosaurs (order Ichthyosauria) used the same method. The marine ichthyosaurs, which were the reptilian counterpart of the porpoises (family Phocoenidae) in class Mammalia, may have used their flippers as rudders.
The limbless snakes are good swimmers and make lateral undulations similar to those of eels. This fishlike swimming mode requires a flexible body and, usually, a tail of moderate length. Sea snakes have laterally flattened tails that increase their locomotor power. Turtles propel themselves by using their feet as paddles as a part of their quadrupedal limb-movement sequence. Freshwater turtles have webbed feet, whereas the forelimbs of marine turtles are essentially flattened flippers that are moved in a sweeping figure-eight pattern through the water.
Flying
Several groups of reptiles have experimented with flight. One group within the Archosauria (the “ruling reptiles” that include dinosaurs and crocodiles) became highly successful at this means of locomotion and evolved into birds.
Another group of archosaurs, the Pterosauria, developed wings that were supported along the front margin by the arm and an extremely elongated finger. The pterosaur wing was made of skin; since it lacked both internal supports and feathers, it probably lacked the flexibility or durability of a bird wing. Pterosaurs seem to have emphasized soaring and gliding during flight, but they also engaged in flapping flight. It is possible that pterosaurs had clumsy takeoffs like those of the modern albatross (Diomedea). Since most pterosaur remains have been found in marine deposits, it is assumed that many of the species lived along ocean shores, probably roosting on cliffs from which takeoff would have been easier.
Among modern lizards, flying lizards (Draco) are expert gliders. The “wing” of these small reptiles is made up of skin supported by five or six elongated ribs between the forelimbs and hind limbs. At rest the ribs and wings are folded against the sides of the body. During flight the wings form broad semicircles on each side between the limbs. These lizards, which live in the forested country of Southeast Asia and the East Indies, are gliders; in spite of their label as flying lizards, they are not flyers. To glide, the lizard launches itself from a tree into the air toward another tree, turning upward sharply to slow its body just before lighting on the new perch. Since the limbs are not modified, this lizard walks and runs like any other arboreal lizard.
Form and function
External covering
The external covering of reptiles is dry and composed of scales made of keratin. It bears few glands or none at all and differs in this respect from the skin of amphibians and mammals. The epidermis has cycles of growth, and in the outer layer the cells die and harden into a tough and horny surface. Bony plates (osteoderms) develop in the dermis, the layer just below the epidermis, of some reptiles and are best seen on the backs of crocodiles.
Internal features
Skeletal system
The skeletons of reptiles fit the general pattern of vertebrates. They have a bony skull, a long vertebral column that encloses the spinal nerve cord, ribs that form a protective bony basket around the viscera, and a framework of limbs.
Each group of reptiles developed its own particular variations on this major pattern in accord with the general adaptive trends of the group. Snakes, for example, have lost the limb bones, although a few retain vestiges of the hind limbs. The limbs of several types of marine reptiles have been modified into fins or flippers. In other types, such as the extinct marine-dwelling ichthyosaurs and plesiosaurs, the bones of the limbs, which no longer needed to support the weight of the body against the pull of gravity, became much shorter. At the same time, the bones of other reptiles that composed the digits multiplied in number, forming a long flipper.
Groups of reptiles whose modes of life came to depend heavily on passive defense also developed specializations of the skeleton. The bony and horny shells of turtles and the rows of bony plates on the backs of crocodiles and the Ankylosaurus (a genus of dinosaurs that lived between the Early Jurassic Period and the end of the Cretaceous) are cases in point.
Skull and dentition
The skulls of the several subclasses and orders vary in the ways mentioned below. In addition to differences in openings on the side of the skull and in general shape and size, the most significant variations in reptilian skulls are those affecting movements within the skull.
As a group, reptilian skulls differ from those of early amphibians. Reptiles lack an otic notch (an indentation at the rear of the skull) and several small bones at the rear of the skull roof. The skulls of modern reptiles are also sharply set off from those of mammals in many ways, but the clearest differences occur in the lower jaw and adjacent regions. Reptiles have a number of bones in the lower jaw, only one of which, the dentary, bears teeth. Behind the dentary a small bone, the articular, forms a joint with the quadrate bone near the rear of the skull. In contrast, the lower jaw of a mammal is made up of a single bone, the dentary; the articular and quadrate have become part of the chain of little bones in the middle ear. An almost complete transition between these two very different arrangements is known from fossils of early synapsids (order Therapsida).
The dentition of most reptiles shows little specialization in a given row of teeth. A dentition that divides groups of teeth into distinctive bladelike incisors, tusklike canines, and flat-crowned molars occurs in mammals but does not occur in reptiles. Instead, the entire tooth row is usually made up of long conical teeth. Venomous snakes have one or several hollow or grooved fangs, but they have the same shape as most snake teeth. The principal differences between species lie in the number, length, and position of the teeth. Crocodiles among the living forms and dinosaurs among the extinct forms have but a single upper and a single lower tooth row. Snakes and many extinct reptilian groups have teeth on the palatal bones (vomer, palatine, pterygoid) and on the bones of the upper jaw (premaxilla, maxilla). However, only one row of teeth is present on the lower jaw.
Lizards have conical or bladelike bicuspid or tricuspid teeth. Some species have conical teeth at the front of the jaws and cuspid teeth toward the rear, but the latter are not comparable to the molars of mammals in either form or function. (They are neither flat-crowned nor used to grind food.) Turtles, except for the earliest extinct species, lack teeth. Instead, they have upper and lower horny plates that serve to bite off chunks of food.
The teeth of reptiles are also less specialized in function than are mammalian teeth. The larger carnivorous reptiles are equipped only to tear off or bite off large pieces of their prey and swallow them without chewing. Insectivorous lizards, which constitute the majority of all lizards, usually crack the exoskeleton of their insect prey, and then they swallow the prey without grinding it up. Snakes simply swallow their prey whole without any mechanical reduction, although the puncture wounds permit digestive enzymes to enter the prey to aid digestion.
Many reptiles developed joints (in addition to the hinge for the lower jaw) within the skull that permit the slight movement of one part relative to others. The capacity for such movement within the skull, called kinesis, enables an animal to increase the gape of the mouth and thus is an adaptation for swallowing large objects. Apparently some of the large carnivorous theropod dinosaurs (such as Allosaurus) had a joint between the frontal and parietal bones in the roof of the skull. All reptiles of the super order Lepidosauria (lizards, snakes, and tuatara) have kinetic skulls, but they differ from the dinosaurs in that the joint on the floor of the skull occurs at the juncture of basisphenoid and pterygoid bones in lepidosaurians.
The skulls of the lepidosaurians became increasingly kinetic as new groups evolved. The Sphenodontia (which include living tuatara [Sphenodon]) and their antecedents, the Rhynchocephalia, had only the basisphenoidal-pterygoidal joint. The lizards lost the lower temporal bar, thereby freeing the quadrate bone and allowing greater movement to the lower jaw, which is hinged to the quadrate. Finally, in the snakes, this trend culminates in the most kinetic skull among the vertebrates. The skulls of snakes possess the ancestral basisphenoidal-pterygoidal joint, a highly mobile quadrate (which gives even greater mobility to the lower jaw), and upper jaws capable of rotating on their longitudinal axes and moving both forward and backward. Many snake species also have a hinge on the roof of the skull between the nasal and frontal bones that allows the snout to be raised slightly. In short, the only part of a snake’s skull incapable of movement is the braincase.
Nervous system
As in all vertebrates, the nervous system of reptiles consists of a brain, a spinal nerve cord, nerves running from the brain or spinal cord, and sense organs. When compared with mammals, reptiles have proportionately smaller brains. The most important difference between the brains of these two vertebrate groups lies in the size of the cerebral hemispheres, the principal associative centres of the brain. These hemispheres make up the bulk of the brain in mammals and, when viewed from above, almost hide the rest of the brain. In reptiles the relative and absolute size of the cerebral hemispheres is much smaller. The brain of snakes and alligators forms less than 1⁄1,500 of total body weight, whereas in mammals such as squirrels and cats the brain accounts for about 1/100 of body weight.