Chameleons or chamaeleons (family Chamaeleonidae) are a distinctive and highly specialized clade of Old World lizards with 202 species described as of June 2015. These species come in a range of colors, and many species have the ability to change color.

Chameleons are distinguished by their zygodactylous feet; their very extensive, highly modified, rapidly extrudable tongues; their swaying gait; and crests or horns on their brow and snout. Most species, the larger ones in particular, also have a prehensile tail. Chameleons' eyes are independently mobile, but in aiming at a prey item, they focus forward in coordination, affording the animal stereoscopic vision.

Chameleons are adapted for climbing and visual hunting. They live in warm habitats that range from rain forest to desert conditions, with various species occurring in Africa, Madagascar, southern Europe, and across southern Asia as far as Sri Lanka. They also have been introduced to Hawaii, California, and Florida, and often are kept as household pets.

Description & appearance

Chameleons vary greatly in size and body structure, with maximum total lengths varying from 15 mm (0.59 in) in male Brookesia micra (one of the world's smallest reptiles) to 68.5 cm (27.0 in) in the male Furcifer oustaleti. Many have head or facial ornamentation, such as nasal protrusions, or horn-like projections in the case of Trioceros jacksonii, or large crests on top of their heads, like Chamaeleo calyptratus. Many species are sexually dimorphic, and males are typically much more ornamented than the female chameleons.

The feet of chameleons are highly adapted to arboreal locomotion, and species such as Chamaeleo namaquensis that have secondarily adopted a terrestrial habit have retained the same foot morphology with little modification. On each foot, the five clearly distinguished toes are grouped into two fascicles. The toes in each fascicle are bound into a flattened group of either two or three, giving each foot a tongs-like appearance. On the front feet, the outer, lateral, group contains two toes, whereas the inner, medial, group contains three. On the rear feet, this arrangement is reversed, the medial group containing two toes, and the lateral group three. These specialized feet allow chameleons to grip tightly onto narrow or rough branches. Furthermore, each toe is equipped with a sharp claw to afford a grip on surfaces such as bark when climbing. It is common to refer to the feet of chameleons as didactyl or zygodactyl, though neither term is fully satisfactory, both being used in describing totally different feet, such as the zygodactyl feet of parrots or didactyl feet of sloths or ostriches, none of which is significantly like chameleon feet. Although "zygodactyl" is reasonably descriptive of chameleon foot anatomy, their foot structure does not resemble that of parrots, to which the term was first applied. As for didactyly, chameleons visibly have five toes on each foot, not two.

Some chameleons have a crest of small spikes extending along the spine from the proximal part of the tail to the neck; both the extent and size of the spikes varies between species and individuals. These spikes help break up the definitive outline of the chameleon, which aids it when trying to blend into a background.

Chameleons have the most distinctive eyes of any reptile. The upper and lower eyelids are joined, with only a pinhole large enough for the pupil to see through. Each eye can pivot and focus independently, allowing the chameleon to observe two different objects simultaneously. This gives them a full 360-degree arc of vision around their bodies. Prey is located using monocular depth perception, not stereopsis. Chameleons have very good eyesight for reptiles, letting them see small insects from a 5–10 meter distance.[citation needed] In fact, chameleons have the highest magnification (per size) of any vertebrate.

Like snakes, chameleons do not have an outer or a middle ear, so there is neither an ear opening nor an eardrum. However, chameleons are not deaf: they can detect sound frequencies in the range of 200–600 Hz.

Chameleons can see in both visible and ultraviolet light. Chameleons exposed to ultraviolet light show increased social behavior and activity levels, are more inclined to bask and feed, and are also more likely to reproduce, as it has a positive effect on the pineal gland.

Distribution & habitat

Chameleons primarily live in the mainland of sub-Saharan Africa and on the island of Madagascar, although a few species live in northern Africa, southern Europe, the Middle East, southern India, Sri Lanka, and several smaller islands in the western Indian Ocean. There are introduced, feral populations of veiled and Jackson's chameleons in Hawaii, and isolated pockets of feral Jackson's chameleons have been reported in California, Florida and Texas.

Chameleons inhabit all kinds of tropical and mountain rain forests, savannas, and sometimes deserts and steppes. The typical chameleons from the subfamily Chamaeleoninae are arboreal, usually living in trees or bushes, although a few (notably the Namaqua chameleon) are partially or largely terrestrial. Most species from the subfamily Brookesiinae, which includes the genera Brookesia, Rieppeleon, and Rhampholeon, live low in vegetation or on the ground among leaf litter. Many species of chameleons are threatened by extinction. Declining chameleon numbers are due to habitat loss.

Food

All chameleons are primarily insectivores that feed by ballistically projecting their long tongues from their mouths to capture prey located some distance away. While the chameleons' tongues are typically thought to be one and a half to two times the length of their bodies (their length excluding the tail), smaller chameleons (both smaller species and smaller individuals of the same species) have recently been found to have proportionately larger tongue apparatuses than their larger counterparts. Thus, smaller chameleons are able to project their tongues greater distances than the larger chameleons that are the subject of most studies and tongue length estimates, and can project their tongues more than twice their body length.

The chameleon's tongue apparatus consists of highly modified hyoid bones, tongue muscles, and collagenous elements. The hyoid bone has an elongated, parallel-sided projection, called the entoglossal process, over which a tubular muscle, the accelerator muscle, sits. The accelerator muscle contracts around the entoglossal process and is responsible for creating the work to power tongue projection, both directly and through the loading of collagenous elements located between the entoglossal process and the accelerator muscle. The tongue retractor muscle, the hyoglossus, connects the hyoid and accelerator muscle, and is responsible for drawing the tongue back into the mouth following tongue projection.

Tongue projection occurs at extremely high performance, reaching the prey in as little as 0.07 seconds, having been launched at accelerations exceeding 41 g. The power with which the tongue is launched, known to exceed 3000 W kg−1, exceeds that which muscle is able to produce, indicating the presence of an elastic power amplifier to power tongue projection. The recoil of elastic elements in the tongue apparatus is thus responsible for large percentages of the overall tongue projection performance.

One consequence of the incorporation of an elastic recoil mechanism to the tongue projection mechanism is relative thermal insensitivity of tongue projection relative to tongue retraction, which is powered by muscle contraction alone, and is heavily thermally sensitive. While other ectothermic animals become sluggish as their body temperatures decline, due to a reduction in the contractile velocity of their muscles, chameleons are able to project their tongues at high performance even at low body temperatures. The thermal sensitivity of tongue retraction in chameleons, however, is not a problem, as chameleons have a very effective mechanism of holding onto their prey once the tongue has come into contact with it, including surface phenomena, such as wet adhesion and interlocking, and suction. The thermal insensitivity of tongue projection thus enables chameleons to feed effectively on cold mornings prior to being able to behaviorally elevate their body temperatures through thermoregulation, when other sympatric lizards species are still inactive, likely temporarily expanding their thermal niche as a result.

Chameleons generally eat insects, but larger species, such as the common chameleon, may also take other lizards and young birds.
The range of diets can be seen from the following examples:

-The veiled chameleon, Chamaeleo calyptratus from Arabia, is insectivorous, but eats leaves when other sources of water are not available. It can be maintained on a diet of crickets. They can eat as many as 15–50 large crickets a day.
- Jackson's chameleon (Trioceros jacksonii) from Kenya and northern Tanzania eats a wide variety of small animals including ants, butterflies, caterpillars, snails, worms, lizards, geckos, amphibians, and other chameleons, as well as plant material, such as leaves, tender shoots, and berries. It can be maintained on a mixed diet including kale, dandelion leaves, lettuce, bananas, tomatoes, apples, crickets, and waxworms.
- The common chameleon of Europe, North Africa, and the Near East, Chamaeleo chamaeleon, mainly eats wasps and mantises; such arthropods form over three quarters of its diet. Some experts advise that the common chameleon should not be fed exclusively on crickets; these should make up no more than half the diet, with the rest a mixture of waxworms, earthworms, grasshoppers, flies, and plant materials such as green leaves, oats, and fruit.
- Temperature influences the amount of food eaten.
- Some chameleons like the panther chameleon of Madagascar regulate their vitamin D3 levels, of which their insect diet is a poor source, by exposing themselves to sunlight since its UV component increases internal production.

Reproduction

Chameleons are mostly oviparous, with some being ovoviviparous.

The oviparous species lay eggs three to six weeks after copulation. The female will dig a hole — from 10–30 cm (4–12 in), deep depending on the species — and deposit her eggs. Clutch sizes vary greatly with species. Small Brookesia species may only lay two to four eggs, while large veiled chameleons (Chamaeleo calyptratus) have been known to lay clutches of 20–200 (veiled chameleons) and 10–40 (panther chameleons) eggs. Clutch sizes can also vary greatly among the same species. Eggs generally hatch after four to 12 months, again depending on species. The eggs of Parson's chameleon (Calumma parsonii), a species which is rare in captivity, are believed to take more than 24 months to hatch.

The ovoviviparous species, such as the Jackson's chameleon (Trioceros jacksonii) have a five- to seven-month gestation period. Each young chameleon is born within the sticky transparent membrane of its yolk sac. The mother presses each egg onto a branch, where it sticks. The membrane bursts and the newly hatched chameleon frees itself and climbs away to hunt for itself and hide from predators. The female can have up to 30 live young from one gestation.