Chameleons

(Chamaeleonidae)

Class: Reptilia

Order: Squamata

Suborder: Sauria

Family: Chamaeleonidae

Thumbnail description
Very small to large, arboreal, semiterrestrial, and terrestrial reptiles with pincer-like feet, prehensile tails, and long projectile tongues

Size
0.75–28 in (19–711.2 mm)

Number of genera, species
6 genera; 180 species and subspecies

Habitat
Forest, savanna, and desert

Conservation status
Vulnerable: 4 species

Distribution
Southern Portugal and Spain, Sicily, Malta, southern Greece (southern Peloponnese) Samos, Chios, Crete, southern Turkey, Cyprus, Syria, Lebanon, Jordan, Israel, Sinai Peninsula, Yemen, Saudi Arabia, Iraq, Iran, Pakistan, India, Sri Lanka, Africa and neighboring islands, Fernando Póo, Canary Islands, Socotra, Pemba, Zanzibar, Mafia, Seychelles, Comoro Islands; Madagascar and adjacent islets, Nosy Be, Nosy Boraha (Île Sainte-Marie-de-Madagascar), Nosy Faly, Nosy Ambariovato, Nosy Mangabe, Nosy Tanikely, Nosy Alanana (Île aux Prunes), and Nosy Sakatia. Introduced to Hawaii, Réunion, and possibly Mauritius and several other islands near Africa, Madagascar, and Greece.

Evolution and systematics

The fossil record is sparse, but the history of chameleons may be more than 60 million years old. Although chameleons are believed to have originated in Africa or Madagascar, the oldest known fossil, 26 million years old, is Chamaeleo caroliquarti from western Bohemia. Based on the known fossil record, chameleons were distributed in Africa but also in parts of the world where they are not found today, such as China, Bavaria, and western Bohemia. Chameleons eventually disappeared from the latter three regions, perhaps as a consequence of changing climatic conditions that favored cooler temperatures and lower humidity.

New areas were inhabited as chameleons radiated to more hospitable climates and evolved into new forms. Mountains, forests, and savannas isolated some species, and their morphologic characteristics evolved to include rows of scales called crests that were high, wavy, or spiky on the back (dorsal crest), throat (gular crest), or belly (ventral crest). A number of chameleons developed one, two, three, four, or six bony horns of different shapes and sizes, flexible extensions on the snout, movable flaps of skin on the side of the head, and other differentiating characteristics, such as patterns, coloration, and body shape and size. All chameleons retained certain prominent features, however, that in combination distinguish them from all other lizards, including projectile tongues used to capture prey, large protruding eyes encased in an eyelid with a tiny aperture referred to as eye turrets, toes fused in bundles of two and three to form grasping pincers, and a prehensile tail.

The classification of this diverse group of lizards has undergone many revisions in genera, families, subfamilies, species, and subspecies throughout the nineteenth and twentieth centuries. The naturalist John E. Gray introduced 16 genera in 1865, and Franz Werner later reduced the number to three genera but also created three new families in 1902. The numbers of species discovered and described increased over the years, but some of them later were eliminated as synonyms of taxa already described and named by previous authors. Werner recognized 70 species in 1902 and raised the number to 88 by 1911; by 1981 Vincent A. Wager recognized 113. By 1986 the number of recognized species had risen to 128, according to Charles Klaver and Wolfgang Böhme, who revised the entire phylogeny of the family based on the morphologic features of the male sexual organs and the lung morphology, bone structure, and chromosome characteristics. While elements of this classification system are not finalized and may be subject to change, it has been accepted worldwide as best representing the relationships within the family Chamaeleonidae. At the last published revision of the system in 1997, Klaver and Böhme recognized no subfamilies, six genera, and two subgenera (Chamaeleo and Trioceros). Within these groupings they cited a total of 171 forms (species plus subspecies):

• Bradypodion: 27
• Brookesia: 24
• Calumma: 25
• Chamaeleo (Chamaeleo): 24
• Chamaeleo (Trioceros): 37
• Furcifer: 20
• Rhampholeon: 14

A few subspecies were elevated to full species after 1997, and several new species were discovered and described, primarily from Madagascar. By 2002 the total number of valid species and subspecies was 180, but this number is likely to change in the future.

Physical characteristics

Chameleons are best known for their ability to change colors. The palette of any species is limited to only certain colors, however. In the case of members of the genera Brookesia and Rhampholeon, the palette consists mainly of shades of tan, brown, and black. The coloration of juvenile chameleons is usually more cryptic than adults of the same species, which may help conceal them from predators. The most dramatic and varying coloration probably belongs to the panther chameleon, Furcifer pardalis, from Madagascar. Within their wide geographic range in the northern third of the island, the coloration of adult males varies significantly from locale to locale, with numerous distinctive color palettes, such as pink and blue, green and red, aqua blue and green, red-orange and white, and turquoise and navy blue.

Chameleons display variations of their color palettes in response to psychological or physiological stimuli and to communicate, not to match their background as was once believed. Although chameleons lack vocal chords, some species are capable of vibration that produces an audible sound or can expel air forcibly from the lungs to generate a hissing or squeaking noise. Chameleons also are known to make sounds in a frequency inaudible to the human ear, but they cannot hear very well, because they lack eardrums and external ear openings.

A male communicating his intentions to a mate often sports the most vivid colors at his disposal. A female likewise will display coloration to communicate her willingness to mate. Calm, subdued colors may indicate receptivity, where dark, intense coloration warns her suitor to stay away. In a few species, such as Calumma boettgeri and C. nasuta, females show striking purplish blue spots, called "threat spots," on the head to deter males. Competing males exhibit bright and intense colors, but the loser usually changes to drab coloration and slinks away to indicate that the contest is over. Chameleons that have a range of colors in their palette may manifest them on different parts of the body, such as the legs, throat, or head. Some are capable of showing stripes and patterns that recede when the chameleon is not in an excited or stressed state. Calm chameleons typically display the least vivid colors. An ill chameleon may become dark or pale in coloration, and sleeping chameleons are often very pale. Color also plays a part in thermoregulation; dark colors absorb the sun's rays when chameleons are cold, and paler colors deflect sunlight.

The epidermis does not grow, and the chameleon sheds completely from time to time as it outgrows this layer of skin. Two cell layers beneath the transparent outer layer can contain red and yellow cells containing pigment granules called chromatophores, and, beneath them, there are cell layers that reflect blue and white light. Below these layers are black or brown pigment called melanin. As these layers of cells expand, contract, and overlap due to result of stimulation (or the lack of it), the chameleon can rapidly change color.

Sex determination in adult chameleons is not a difficult matter for most chameleon species, because they most often are sexually dimorphic, meaning males and females are different in form or size. For example, in the majority of species where males have horns, females lack horns. Males are usually larger than females, except in the genera Brookesia and Rhampholeon. Species that are not sexually dimorphic may be different in coloration, or sexually dichromatic, such as Furcifer pardalis. Females of this species are typically a reddish orange or tan marked with brown or black, regardless of geographic locale. Determining the sex of species in the genera Brookesia and Rhampholeon depends primarily on the presence of a bulge at the base of the tail created by paired sexual organs called hemipenes. It is much more difficult to ascertain the sex of juveniles of most species from birth to about six months of age, or whenever the first indication of adult coloration, horns, crests, or a hemipenial bulge becomes apparent.

The most important physical feature of a chameleon is its large and protruding eyes. A chameleon can move its eyes independently and is able to process two images at once. This ability is the chameleon's best defense against predators, because its hearing is very poor. It scans the surrounding environment with telescopic vision that enables it to plan and execute a defense (usually concealment or flight) well in advance of the predator's approach. Phenomenal eyesight also facilitates locating prey from a great distance. As chameleons target prey, two separate images merge into one to gauge distance. Then chameleons engage their most fascinating feature—the tongue.

In 2000 a group of researchers published the results of a study on the mechanics of prey prehension in chameleons that unraveled the mysteries of how a chameleon's tongue really works. The hyoid bone is a piece of cartilage that extends into the mouth from the throat bones (called the hyolingual apparatus) and is attached to a chameleon's long tongue. This is where the tongue rests when it is not in use. The tongue is launched from the hyoid bone with the use of ringed muscles in the tongue. This highly complex structure, composed of cartilage, muscles, nerves, glands, and tissue, is used this way in prey capture:

• The central cylindrical accelerator muscle is responsible primarily for projecting the tongue for prey capture.
• The tip of the tongue, or tongue pad, sits atop the accelerator muscle, connected by several pairs of muscles. As the chameleon launches the tongue pad at prey, it turns inside out (evaginates) and actively reverses (invaginates) to form a pouch immediately before prey contact.
• The pouch engulfs the prey, and wet adhesion and interlocking maintain grip while suction created by the largest paired muscles, the pouch retractor, transfers prey deeper into the pouch.
• The tongue retractors attached to the accelerator muscle return the tongue to the resting position on the hyoid bone, and the prey is in the chameleon's mouth to be crushed and swallowed.

Before this study, the capture of prey often was attributed only to adhesion to the tongue pad. The withdrawal of the tongue pad to form a pouch not only creates suction forces on the prey but also increases the adhesive properties of the tongue. Suction accounts for more than two-thirds of the total force generated by a chameleon's tongue. This permits capture of larger prey, such as lizards and birds, than is possible using just adhesion. Chameleons also employ the tongue pad to lap drinking water from leaves or other surfaces.

The skeletal structure of chameleons is remarkable for it's flexibility. They can compress their bodies to bask in sunlight or inflate their lungs and expand the rib cage to bluff potential predators. Chameleon feet are designed to grasp, with five toes on each foot fused in bundles of two and three toes to form a pincer. On the front feet the bundle of three toes is on the inside of the foot, and the bundle of two toes is on the outside. This is reversed on the rear foot, giving them a secure and strong grasp and allowing them to navigate horizontally or vertically on a wide variety of vegetation or structures. Sharp claws on each toe help them climb and grip surfaces that they cannot grasp tightly, such as tree trunks, and are used by females to excavate tunnels to lay eggs.

In the genera Bradypodion, Calumma, Chamaeleo, and Furcifer, tail length is roughly equal to or slightly longer than body length. These species can use their prehensile tail as a fifth limb and to anchor themselves while launching their long tongue at prey. Some species, such as Parson's chameleon (C. parsonii parsonii), use their tails to communicate. Males engaged in ritualistic threat displays repeatedly coil the tips of their long tails tightly, curl them up and over the back, and whip them forward. Sleeping chameleons often roll their long tails into a perfect coil like a watch spring, and chameleons may intertwine tails during copulation. Members of the genera Brookesia and Rhampholeon have much shorter, less prehensile tails that nevertheless can be used as a grasping hook in some species.

Distribution

Chameleons occur naturally only in the Old World. Africa (including offshore islands) has the highest concentration of species and subspecies, with all 27 members of the genus Bradypodion, all 14 forms in the genus Rhampholeon, and 59 forms in the genus Chamaeleo, totaling 99 species and subspecies. Forty percent of the world's species inhabit Madagascar and offshore islands, including 19 of the 21 members of the genus Furcifer, all 27 forms in the genus Brookesia, and 28 forms in the genus Calumma, for a total of 73 species and subspecies. The eight remaining forms are from Yemen (one),

Saudi Arabia (two), India and Sri Lanka (one), the Comoro Islands (two), Socotra (one), and the Seychelles (one).

One species, Chamaeleo chamaeleon, or the common chameleon, is found in Europe, the Middle East, Greece, northern Africa, southwestern Saudi Arabia, and Yemen. This makes up the widest range of distribution of any chameleon species. Jackson's chameleon (Chamaeleo jacksonii) has the dubious distinction of being the first chameleon species to be introduced and become well established in the New World. A few dozen specimens imported for the pet trade in the 1970s escaped into the wilderness in Hawaii, creating a large feral population that has continued to thrive more than 30 years later.

Habitat

Chameleon habitat is as varied as the species in this diverse family of reptiles. One species, Chamaeleo namaquensis, lives in one of the most inhospitable regions on earth, the Namib Desert in Africa. This sturdy and aggressive chameleon tolerates extremely high temperatures by day and near freezing temperatures at night and lives a terrestrial existence near the sparse vegetation of sand dunes. Other species are far less tolerant of such extreme temperatures and require high humidity, particularly species that are montane or rainforest specialists. These species may not survive the loss of their complex environmental niches in the future, whether from deforestation, modification, or climatic change. Some chameleon species have adapted to the degraded vegetation that invariably accompanies the burgeoning human population in underdeveloped countries. Unprotected natural forest is frequently burned or cut for grazing, agriculture, fuel, and housing. Sometimes agriculture, such as fruit or coffee trees, provides alternate habitat for arboreal creatures like chameleons, but such crops as rice do not.

There are chameleon species that utilize vegetation in or near virtually every forest type, including lowland evergreen broadleaf rainforest, semi-evergreen moist broadleaf forest, deciduous or semi-deciduous broadleaf forest, thorn forest, upper and lower montane forest, cloud forest, disturbed natural forest, and exotic or native species plantations. Other species live in grassland, scrub, or semidesert conditions and can be found from sea level all the way up to elevations of nearly 15,000 ft (4,500 m).

Behavior

Chameleons are diurnal, and many species begin the day at dawn by seeking a spot to bask in the sun to increase body temperature and metabolism. Chameleons are ectothermic (cold-blooded) and must regulate their temperature by exposing their body to sun or shade. Once they reach a comfortable temperature, they may begin seeking prey or lap dew or rain to quench thirst. A typical day is spent resting or seeking prey while keeping a watchful eye for predators as they move about in the environment. All moving objects must be analyzed as a potential threat. If the object advances in their direction, chameleons may move off into foliage or attempt to conceal themselves by swiveling behind their perches, known as "squirreling." The main predators of chameleons are birds and snakes.

At dusk chameleons seek a place to roost and sleep and often return to the same location every night. Many species roost at the ends of branches to sleep. Small species may wrap their tails around the stems and drape their bodies on top of large leaves, with their heads pointed down. If a predator touches the branch, leaf, or plant, they release their grip and slide or drop to the ground and play dead or scurry into the underbrush.

Suitable chameleon habitat, whether disturbed or undisturbed, must include enough space to support a viable population of these often asocial and territorial reptiles. Males are usually intolerant of other males of the same species within visual proximity, especially during mating season. Females normally avoid males when they are gravid or unreceptive to mating. Habitat partitioning by height and life stage may relieve some of these pressures in dense populations within a restricted area of habitat, but conflicts can result in injuries or even death.

Feeding ecology and diet

Chameleons consume a wide variety of flying and crawling insects, butterflies, moths, larvae, snails, and spiders in nature, and larger species consume some vertebrates as well. Chameleons prey on smaller chameleons, lizards, birds, and even snakes. Captive chameleons will accept young mice, but it is unlikely that this is a natural prey. Chameleons also ingest vegetation, including leaves, flowers, and fruits. Other organic matter, such as bark, twigs, moss, and soil are sought out and consumed by chameleons, but the nutritional or medicinal value of some of these items is unknown. Chameleons are sit-and-wait ambush predators, but many species are quite mobile and travel long distances seeking prey along the way, while others are much more sedentary and utilize a smaller range. There is anecdotal evidence that chameleons congregate in areas where insects appear only at certain times of the year, such as when insects are attracted to coffee blooming or when cicadas hatch. When these food supplies are no longer present, the chameleons disperse.

Reproductive biology

Reproduction in chameleons typically begins with ritualistic courtship displays by males. In many species this entails the display of bright colors and a series of jerking or bobbing head movements while advancing on a female. Some males advance slowly with a halting or jerky gait, but others move very quickly and can be aggressive toward females. Females that are unreceptive or gravid may flee or may face the suitor with gaping mouth while hissing, rearing up on the hind legs, and rocking to discourage the male's advances. Females are known to approach males and grasp their forelegs or horns to stop their pursuit. In some cases, unreceptive or gravid females attack males and inflict bite wounds that can be fatal.

If the female remains passive to the courtship of the male, he will mount the female by grasping her flanks and position himself on the right or left side of her body. He then everts the nearest of his two sexual organs while inserting it in her cloaca, the common chamber into which the intestinal, urinary, and generative canals discharge in reptiles, and copulation ensues. Some species copulate for a few minutes and others for as long as several hours, after which they usually go their separate ways. A few species form pair bonds for a period of time during the mating season.

The majority of chameleon species are oviparous, meaning they produce offspring by laying eggs in tunnels or pits in the ground or under rocks or leaves after a gestation period that can last a few weeks or several months, depending on the species. Females excavate tunnels and pits by digging with their front feet and then back into them to deposit eggs. When they are finished, they bury the eggs, fill in the tunnel or pit, and stomp the soil down to conceal the location of the nest. Some females drag leaves and twigs over the site. This is the final act of motherhood for a chameleon, and her young will be independent at birth. Incubation times vary according to species and according to the stage of embryonic development at the time the eggs are deposited. The shortest known incubation period is around one month, and the longest is 18 months. The young emerge by slitting a star-shaped opening in the end of the eggshell with the egg tooth, a sharp, calcified protrusion on the tip of the upper jawbone that later falls off.

Ovoviviparous species (those with eggs that hatch within the mother's body, or immediately after being laid) are found primarily in climates that experience greater extremes of cold and likely represent a reproductive strategy to increase the survival rate of neonates where eggs deposited in the ground might not hatch. Basking gravid females often position themselves so that sunlight is directed on their swollen abdomens to warm the developing babies. A female paces nervously while giving birth to her young, which emerge encased in thin, transparent membranes. The neonates wriggle free of the membrane and begin moving and climbing about immediately, usually seeking food within 24 hours. They instinctively disperse, perhaps to avoid predation by the mother; this rarely occurs, however, in the confines of a cage in captive births. While live-bearing females are in contact with their offspring at birth (unlike egg-laying females), they do not nurture them in any way.

Conservation status

In 1996 three chameleon species, Furcifer campani, F. labordi, and F. minor, were classified as Vulnerable by the IUCN, based on a 20% population decline in 10 years, or three generations. A fourth species, Brookesia perarmata, was classified as Vulnerable for this reason and also because they occupy an area of less than 39 sq mi (100 sq km) and fewer than five locations. All chameleons in the genera Bradypodion, Calumma, Chamaeleo, and Furcifer are listed on CITES Appendix II, indicating that they are threatened with extinction unless commercial trade is tightly controlled. A moratorium on importation for commercial trade of all but four species of chameleons (F. pardalis, F. lateralis, F. oustaleti, and F. verrucosus) from Madagascar was imposed by CITES in 1995, owing to escalating levels of trade and concerns that extinction might result. This moratorium remained in effect in 2002. Although Brookesia perarmata is included on the IUCN Red List as Vulnerable, no members of the genera Brookesia or Rhampholeon received formal protection from CITES to prevent unsustainable commercial trade as of 2002.

The main threats to chameleons are ongoing loss, modification, and fragmentation of acceptable habitat and collection for the legal and illegal commercial pet trade. When the extent of occurrence is small, the number of known sites is few, the distribution structure is fragmented, and the species is a specialist within a declining habitat, the risk of extinction is high. This paradigm is applicable to numerous species of chameleons worldwide. The majority of chameleon species have not survived or reproduced in captivity and should not be considered candidates for captive breeding projects aimed at preservation of the species. Habitat preservation and conservation management in the wild for these vulnerable species are critical to preventing future extinction.

Significance to humans

There are relatively few traditional uses for chameleons by local people within their range of distribution, but these uses generally involve burning or killing chameleons for folk medicine or to ward off evil spirits. Some cultural traditions dictate that chameleons must not be harmed. Chameleons are not used very often as food. The major consumption of wild chameleons is for the international commercial live pet trade that reached its apex in the 1990s, when more than 260,000 chameleons were exported from Madagascar and 345,000 from Africa and Yemen. The major consumers are the United States, western Europe, and Asia. The commercial trade in reptiles, particularly those captured in the wild, has been criticized by conservation, scientific, and animal rights organizations as inhumane and because it is detrimental to the survival of wild populations. It has been estimated that less than 1% of chameleons taken in the wild live longer than a few months in captivity. This is primarily the result of captivity-related stress, injury, diseases, parasites, and failure or inability to meet the highly specialized environmental and nutritional requirements necessary for survival in a captive setting. For humane and conservation reasons, chameleons should not be considered appropriate as pets.

Species accounts

Resources

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World Conservation Union (IUCN). Rue Mauverney 28, Gland, 1196 Switzerland. Phone: 41 (22) 999-0000. Fax: 41 (22) 999-0002. E-mail: mail@hq.iucn.org Web site:

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[Article by: Ardith L. Abate]

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