Form and function

Skull and jaws

The skull is derived from the primitive diapsid condition, but the lower bar leading back to the quadrate bone is absent, however, giving greater flexibility to the jaw. In some burrowers (such as Anniella and the worm lizards) as well as some surface-living forms (such as the geckos), the upper and lower temporal bars have been lost. Small burrowing lizards have thick, tightly bound skulls with braincases that are well protected by bony walls. In most lizards, the front of the braincase is made up of thin cartilage and membrane, and the eyes are separated by a thin, vertical interorbital septum. In burrowing forms with degenerate eyes, the septum is reduced and adds to the compactness of the skull. Most lizard skulls, particularly in the Scleroglossa, are kinetic (that is, the upper jaw can move in relation to the rest of the cranium). Since the anterior part of the braincase is cartilaginous and elastic, the entire front end of the skull can move as a single segment on the back part, which is solidly ossified. This increases the gape of the jaws and probably assists in pulling struggling prey into the mouth.

Dentition

Most lizards eat a variety of arthropods, with sharp, tricuspid teeth adapted for grabbing and holding. In most lizards, teeth are present along the jaw margin (on the maxilla, premaxilla, and dentary bones). However, in some forms, teeth may also be found on the palate. In the embryo, an egg tooth develops on the premaxilla bone and projects forward from the snout. Although it aids in piercing the shell, it is lost soon after hatching. This is a true tooth, unlike the horny epidermal point in turtles and crocodilians.

The teeth of some large predators are conical and slightly recurved. The Komodo dragon (Varanus komodoensis), for example, has serrated teeth that are curved like a scalpel blade; these teeth can cut through the leg muscle of a full-grown water buffalo (Bubalus bubalis) and cause it to bleed to death. In contrast, mollusk and crustacean feeders, such as the caiman lizard (Dracaena), have blunt, rounded teeth in the back of the jaw designed for crushing. Some herbivorous species (such as iguanas) have leaf-shaped tooth crowns with serrated cutting edges. The venomous lizards (Heloderma) have a longitudinal groove or fold on the inner side of each mandibular tooth; these grooves conduct the venom from the lizard to its victim.

The common mode of tooth implantation is pleurodonty, in which the teeth are fused to the inner side of the labial wall. In the other mode, acrodonty, teeth are fused to the tooth-bearing bone, often to the crest of the bone. Acrodont teeth are rarely replaced once a certain growth stage is reached. The dentition of the Agamidae is usually described as acrodont, but most species have several pleurodont teeth at the front of the upper and lower jaws.

Locomotion and limb adaptations

Most lizards are quadrupedal and have a powerful limb musculature. They are capable of rapid acceleration and can rapidly change direction. The racerunners or whiptails (Aspidoscelis) can attain speeds of 29 km (18 miles) per hour, which, in terms of their own body length (less than 50 cm [20 inches] long), puts them in a class with fast terrestrial mammals. A tendency toward elongation of the body is found in some families, and a reduction of limb length or a complete loss of limbs often accompanies such elongation. Such lizards propel themselves entirely by lateral undulations emanating from highly complicated ventral abdominal muscles. Limbless lizards that move quickly on the surface or through sand (such as glass snakes [Ophisaurus]) tend to have elongate tails, whereas the burrowers have extremely reduced tails. Some burrowers (such as the amphisbaenians) dig by ramming the head into the substrate. This is followed by the rotation of the head around the head joint to compress the substrate. Others, like the California legless lizards (Anniella), literally “swim” through the sand.

Many modifications of the toes occur in lizards. Some desert geckos, the iguanid Uma, and the lacertid Acanthodactylus have fringes on the toes that provide increased surface area, preventing the lizard from sinking into loose desert sand. Arboreal geckos and anoles (Anolis) have lamellae (fine plates) on the undersides of the toes. Each lamella is made up of brushlike setae. The tips of each seta divide hundreds of times into tiny spatulae (spoon-shaped strands); the final strand is less than 0.25 micrometre (0.00001 inch) in diameter. (A tokay gecko [Gekko gecko], for example, has about half a million setae on each foot.) These fine hairlike processes greatly enhance the clinging ability of the lizards, allowing some to easily climb vertical panes of glass. Intermolecular forces between spatulae on the gecko’s setae and the surface provide the adhesion.

The true chameleons (family Chamaeleonidae), a predominantly arboreal group, have a different type of highly specialized limb. The digits on each foot are divided into two groups by webs of skin. On each hind limb, three of the toes face away from the body, whereas two face toward the body; on each forelimb, the pattern is reversed. Each foot can thus be divided into an outer and an inner portion, which can be opposed as the branch is gripped. Chameleons and some other lizards have prehensile tails, which also aid in grasping branches.

Several terrestrial lizards are able to run bipedally. Basilisk lizards (Basiliscus) are actually able to run across water for short distances. During bipedal locomotion the tail is held out backward and upward and acts as a counterweight. The frilled lizard (Chlamydosaurus kingii) can also run bipedally.

Some lizards are able to parachute or glide through the air and make soft landings. The most highly adapted of these are the flying lizards (Draco), a group of agamids from Southeast Asia. The “wings” that enable this lizard to glide are extensible lateral folds of skin that are supported by elongate ribs.