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Chameleons (Chamaeleonidae)

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

List of species

KwaZulu-Natal Midlands dwarf chameleon
Armored chameleon
Parson's chameleon
Veiled chameleon
Common chameleon
Jackson's chameleon
Jeweled chameleon
Minor chameleon
Panther chameleon
Short-tailed chameleon

KwaZulu-Natal Midlands dwarf chameleon

Bradypodion thamnobates

taxonomy

Bradypodion thamnobates Raw, 1976, Nottingham Road, Natal, South Africa.

other common names

Afrikaans: Natalse Middelveld.

physical characteristics

This species is 6–7.5 in (152–191 mm) in length. Females are marginally smaller than males. Male coloration is dark green or blue-green with white markings on the head and a prominent gular crest composed of lobed scales. There is also a spiky dorsal crest. Large conical or rounded scales, which are white, blue, or reddish in color, cover the body and legs. Female coloration is olive green, and the markings, scalation, and crests are less pronounced. Juveniles are brownish, yellowish, or light green.

distribution

The species occurs in South Africa in the southeastern province of KwaZulu-Natal in the midlands between the Mooi River, Bulwer, Howick, and Dargle at an elevation of 3,000–4,200 ft (915–1,281 m).

habitat

These chameleons live among shrubs, bushes, and anthropogenic vegetation, including gardens and hedges in residential areas.

behavior

The species is calm and docile, but males are aggressive toward other males.

feeding ecology and diet

The dwarf chameleon preys upon small crawling and flying insects.

reproductive biology

Males court females by head bobbing while approaching a female. Unreceptive females display defensively or flee. Receptive females remain passive and allow males to copulate. Gestation is from five to eight months, depending on ambient climate, and females give live birth to as many as 18 neonates.

conservation status

Listed as Near Threatened by the IUCN. Limited numbers have been exported to zoos and a few private people as of 2002. The range of distribution of this rare species is poorly protected, continues to erode, and is under serious threat, which may affect its abundance in the future.

significance to humans

These chameleons are used occasionally in traditional medicine by indigenous people in Africa.


Armored chameleon

Brookesia perarmata

taxonomy

Brookesia perarmata Angel, 1933, Antsingy at an elevation of 984 ft (300 m), Province du Ménabé, Madagascar.

other common names

English: Antsingy leaf chameleon.

physical characteristics

At 4–6 in (102–152 mm), this is the largest and most easily identifiable member of the genus Brookesia. It is reddish brown, brown, and tan in coloration. The most remarkable physical feature is a row of pointed scales projecting outward from the spine that continue onto the tail, diminishing in size. The remainder of the body and tail bear numerous thorny scales, giving it the armored appearance for which it is named.

distribution

The only known locale is l'Antsingy d'Antsalova in the Tsingy de Bemaraha Nature Reserve, Madagascar.

habitat

The armored chameleon inhabits bushes, shrubs, and leaf litter in or near dense, dry, deciduous forest, where it is primarily terrestrial. According to E. R. Brygoo, three specimens were collected in 1952 "on the trail from Antsalova to Tsiandro, at the level of the Ambodiriana clearing, in the rocks to the north of the road, among the vegetation between the stones."

behavior

The species is docile, secretive, and often sedentary.

feeding ecology and diet

The armored chameleon feeds on invertebrates within its prey size range.

reproductive biology

Little is known of the reproductive biology of this species.

conservation status

This species is classified as Vulnerable by the IUCN based on a 20% population decline in 10 years, or three generations, owing to levels of exploitation and because the population is characterized by an acute restriction in area of occupancy and number of sites. Despite this listing, it receives no formal protection from CITES to prevent unsustainable commercial trade. It has been exported regularly to the pet trade beginning around 1993, but quantities of harvested specimens were unrecorded as of 2002. Protection from commercial trade and preservation of habitat are critical to preventing extinction.

significance to humans

The only member of the genus Brookesia, the armored chameleon is in high demand for the commercial pet trade. Longevity and reproduction in captivity are poor.


Parson's chameleon

Calumma parsonii

taxonomy

Calumma parsonii Cuvier, 1832, Madagascar. Three geographically discrete forms are recognized, based on size, eye turret color, and/or color of the scales outlining the mouth. One sub-species is recognized.

other common names

French: Caméléon de Parson; German: Parsons Chamäleon; Spanish: Camaleón Parson; Malagasy: Ranotoetrabe, ranotohitra, taintrotro, tampamaly.

physical characteristics

This species grows to 12–28 in (305–71l mm) in length. The nominate form is one of the three largest and the heaviest of all species of chameleons, at weights up to 24.5 oz (700 g). Males are tan, turquoise, or blue-green, with white, tan, or yellow scales outlining the edge of the mouth, orange or blue-green eye turrets, and two robust bony horns protruding from the snout. Females are red-brown, green, or blue-green. Three or more prominent dark stripes curve down from the dorsum toward the anterior of the body. There is no dorsal crest. Juveniles are orange, blue-green, or tan. The subspecies, Calumma parsonii cristifer, is 10–18 in (254–457 mm). Males are deep blue-green with a rust blotch on the flank and a complete dorsal crest. Females are tan, orange, or green.

distribution

C. p. parsonii occurs in Ifanadiana and Ikongo, southeastern Madagascar, and Vavatenina, Nosy Boraha, and Maroantsetra, northeastern Madagascar. C. p. cristifer is found in Andasibe.

habitat

C. p. parsonii has been described as originating from primary rainforest, but as of 2002 little intact forest remained within the known range of distribution. Observations after the 1960s have been limited to fragments of disturbed forest and shady mature tree plantations, such as coffee, lychee, and mango. This species preferentially selects dense vegetation and avoids direct sunlight. The C. p. cristifer habitat is largely within the boundaries of a well-protected national park retaining undisturbed moist montane rainforest as of 2002.

behavior

Parson's chameleon is sedentary and shy. Males engage in dramatic territorial displays but are otherwise passive. Females are easily stressed and display with yellow blotches distributed over the head and body.

feeding ecology and diet

The species consumes a wide variety of vertebrate and invertebrate prey, including small lizards and birds. Malagasy residents report consumption of flowers, fruits, and dry leaves as a component of the total diet.

reproductive biology

No detailed long-term research has been performed on the reproductive biology of C. p. parsonii, but data from captive management indicate that sexual maturity and adult size are attained between three and five years of age, substantially later than any other species of Chamaeleonidae. Courtship and copulation begin from two to four months after winter brumation (dormancy) ends, and the mating season lasts approximately eight weeks. Gestation is from three to five months, and incubation has varied from 13 to 24 months in captivity. Clutch size is 20–60 eggs, and the interval between clutches is one year. Less is known of the reproductive biology for C. p. cristifer. In 1997 one clutch of 37 eggs produced 35 live hatchlings in captivity after 13 months of incubation.

conservation status

Not listed by the IUCN, but listed on Appendix II of CITES; proposals to uplist it to Appendix I were submitted in 1998 and 2001. The 1995 CITES moratorium on importation failed to halt sporadic commercial trade, at least through 2002. The continuing levels and patterns of exploitation place small populations at risk of extirpation and decreased reproductive potential. As of 2002 C. p. parsonii had not been documented in a protected area for more than 30 years. This species is at risk of extinction because it is known from relatively few sites, the distribution structure is highly fragmented, and it is a specialist of a rapidly declining habitat. C. p. cristifer is vulnerable to extinction because population densities are low, and it is known only from a few sites. Conservation is dependent on preservation of suitable habitat and prevention of commercial trade.

significance to humans

From 1986 to 1995, 19,000 wild-taken specimens of C. parsonii were exported legally for the commercial pet trade, and sporadic illegal trade continued subsequent to the 1995 CITES moratorium on importation, owing to high international demand and escalating retail selling prices. Despite long life expectancy, fewer than 1% of specimens imported before 1995 remained alive by 2002, and fewer than 300 neonates had hatched alive in captivity.


Veiled chameleon

Chamaeleo calyptratus

taxonomy

Chamaeleo (Chamaeleo) calyptratus Duméril & Bibron, 1851, Yemen and southwestern Saudi Arabia. One subspecies is recognized.

other common names

English: Cone-head chameleon, Yemen or Yemeni chameleon; German: Jemen-Chamäleon.

physical characteristics

The species grows to 10–24 in (254–610 mm) in length. The most prominent feature is a high, prominent casque that is much larger in males than females and the tallest of any chameleon species. Male coloration is shades of green, turquoise, yellow, orange, white, and black with bold stripes and spots. Nongravid females are green with pale patterning, but gravid females display vivid yellow, blue, and green spots and patterns against a dark background. Juveniles are green at birth, and their sex is distinguished easily by the presence of a short, fleshy projection called a tarsal spur on the hind feet of males. It produces audible sounds.

Although the subspecies Chamaeleo calyptratus calcarifer (Peters, 1871) was still considered valid in 1997, there is anecdotal evidence that it is a hybrid of Ch. calyptratus and Ch. arabicus (Matschie, 1893); this requires further investigation.

distribution

Ch. c. calyptratus occurs in Yemen, centered around Ta'izz and Ibb. Ch. c. calcarifer inhabits Saudi Arabia and possibly Yemen.

habitat

The veiled chameleon primarily utilizes Acacia species in areas of heavy cultivation and exotic plantations, such as Catha edulis. The climate is arid, but the habitat is the greenest part of the Arabian Peninsula, and these chameleons usually are found near a water source.

behavior

The veiled chameleon is considered very aggressive and defensive toward members of its species (conspecifics) and humans. Males can inflict serious injuries or death in territorial or courtship disputes. An adult at the Dallas Zoo attacked a mirror and regularly coiled and uncoiled his tail while threatening his image.

feeding ecology and diet

The species feeds on insects, some vertebrates, and significant amounts of vegetation, particularly acacia.

reproductive biology

Courtship consists of head bobbing, intensified coloring, and approaching the female with a jerky gait. Females unreceptive to mating may gape, swing from side to side, hiss, or even attack the males. Receptive females slowly move off but allow the male to mount and copulate. Males may become aggressive and are known to head butt or bite, sometimes causing grave injury to females.

conservation status

Not listed by the IUCN, but listed on CITES Appendix II. In 2001 it was not considered rare within its range and is adapted to highly disturbed habitat in close proximity to humans. Reports of increasing use of agricultural pesticides may have a detrimental impact on future abundance.

significance to humans

Fewer than 10,000 wild-taken specimens were exported for the commercial pet trade from 1985 to 1999. The species reproduced consistently in captivity and is well established in herpetoculture.


Common chameleon

Chamaeleo (Chamaeleo) chamaeleon

taxonomy

Chamaeleo (Chamaeleo) chamaeleon Linnaeus, 1758, Europe, Middle East, Greece, northern Africa, Egypt (Sinai Peninsula), southwestern Saudi Arabia, and Yemen. Two subspecies are recognized.

other common names

English: European chameleon, Mediterranean chameleon; French: Caméléon commun; German: Europäisches Chamäleon, Gemeines Chamäleon.

physical characteristics

The species attains a length of 8–15 in (200–400 mm). Females are often larger than males. Coloration varies but includes green, yellow, gray, and brown with numerous stripes and spots forming consistent patterns on each form.

distribution

Ch. c. chamaeleon occurs in southern Portugal, southern Spain, Canary Islands, Sicily, Malta, southern Turkey, Cyprus, Syria, Lebanon, Jordan, Israel, southern Peloponnese, Samos, Chios, Crete, western Sahara, Morocco, Algeria, Tunisia, Libya, and Egypt. Ch. c. musae inhabits the Sinai Peninsula and Egypt. Ch. c. orientalis is found in southwestern Saudi Arabia and Yemen.

habitat

Owing to a wide distribution, the species is found in numerous habitat types at elevations up to 8,500 ft (2,600 m), including forests, plantations, and scrub in semidesert or coastal regions, usually near a source of water.

behavior

This chameleon is fairly aggressive toward conspecifics.

feeding ecology and diet

The common chameleon feeds on a wide range of invertebrate and vertebrate prey, including young birds and reptiles, and some vegetation.

reproductive biology

Specimens exposed to seasonal temperatures near freezing experience a dormant period (brumation) until temperatures become warmer. As they return to normal activity, males seek females for mating. Unreceptive females may gape, inflate with air, and butt males. Receptive females remain passive and permit copulation. Several copulations may occur per day until the female adopts dark coloration with orange markings and rejects the male's advances. Females produce up to 60 eggs after a gestation period of two months. Young emerge six to 11 months later, depending on climatic conditions and length of diapause, a dormant period in embryonic development.

conservation status

The species is listed on Appendix II of CITES. In Greece it is strictly protected from collection, killing, abuse, wounding, captivity, and export by a presidential decree, and it is listed in the Greek Red Data Book of Threatened Vertebrates. Populations on Chios and Crete are believed to have been extirpated as of the late 1980s. It is included in the "European Community Habitats Directive" in Appendix IV as a priority species, requiring establishment of protected areas. The species has been proposed for an IUCN classification of Lower Risk: Conservation Dependent in Andalusia, Spain, and the European Union, with an additional classification of Vulnerable in the provinces of Huelva and Cádiz, Spain, based on a study conducted from 1993 to 1999. Major threats are habitat destruction and modification, road mortality, and translocation. The conservation status is unknown in northern Africa, the Middle East, and the Saudi Arabian Peninsula.

significance to humans

The common chameleon was exported from Algeria, Egypt, Jordan, and Morocco in limited numbers from 1986 to 1999 for the commercial pet trade. Life span and reproduction in captivity are considered very poor. Large numbers of specimens are collected and sold at markets in northern Africa for purposes of traditional folk medicine and to ward off bad luck. A common practice in Morocco is to throw live chameleons into a fire. Hundreds of dried chameleons are strung in groups of 30–50 using stout twine and hung in the food markets in Morocco.


Jackson's chameleon

Chamaeleo (Trioceros) jacksonii

taxonomy

Chamaeleo (Trioceros) jacksonii Boulenger, 1896, Uganda, later amended to Kikuyu, near Nairobi, Kenya. Two subspecies are recognized.

other common names

English: Mt. Meru chameleon, three-horned chameleon; French: Caméléon de Jackson; German: Ostafrikanisches Dreihornchamäleon.

physical characteristics

This species grows to 6–14 in (152–356 mm) in length. Ch. j. xantholophus is the largest, followed by Ch. j. jacksonii and then Ch. j. merumontanus. Males have three annulated (composed of rings) horns, two preorbital and one nasal in all three forms. Females may have no horns, a single nasal horn, or three well-developed horns that are slightly smaller than those of the males, depending on the subspecies. The body coloration is shades of green or brown with a dark red, yellow, or blue wash on the head, flanks, or tail. Juveniles are brown, black, and off-white and show an infusion of adult coloration at about the age of six months.

distribution

Ch. j. jacksonii occupies the highland of central Kenya, except the eastern slopes of Mount Kenya. Ch. j. xantholophus lives on the southern, eastern and northeastern faces of Mount Kenya and in Kenya and has been introduced to Hawaii. Ch. j. merumontanus inhabits Mount Meru in Tanzania at an elevation of 7,500–9,000 ft (2,288–2,745 m).

habitat

This is a montane, arboreal species that utilizes various habitat types, including moist forest, exotic plantations, and dense bushes.

behavior

Jackson's chameleon is docile and gentle. Males settle territorial and mating disputes by engaging in pushing contests with their horns, but they rarely resort to biting or inflict injury. Females generally are very calm and tolerant of conspecifics.

feeding ecology and diet

Jackson's chameleon preys on a wide variety of invertebrates.

reproductive biology

Males engage in ritualistic head bobbing to court females. Nonreceptive females become dark, rock, gape, and may grasp the forelegs or horns of an advancing male. If the female is receptive, she lifts her tail to facilitate copulation. Gestation varies from five to nine months, depending on ambient temperatures. The species is ovoviviparous (live-bearing), with births of three to 50 young per clutch, depending on the sub-species.

conservation status

Jackson's chameleon is listed on CITES Appendix II. Kenya ceased exporting specimens of Ch. j. jacksonii and Ch. j. xantholophus in the early 1980s. Hawaii considers feral Jackson's chameleons detrimental to the ecosystem. The species lives mainly in highly degraded habitat and exotic plantations, where it is believed to have adapted successfully to living in close proximity to humans.

significance to humans

The species was exported from Kenya in very high numbers for the commercial pet trade beginning in the 1960s, which was accompanied by very high levels of morbidity and mortality through the 1980s, when Kenya stopped exportation. From 1993 to 1999 more than 12,000 Ch. j. merumontanus individuals were exported from Tanzania, but it is unknown as of 2002 whether this has had a detrimental effect on population densities. This subspecies did not adapt well to captivity, and reproduction is poor. Several generations of Ch. j. xantholophus originating from Hawaii were bred in captivity in the 1990s but only by a very few people worldwide.


Jeweled chameleon

Furcifer campani

taxonomy

Furcifer campani Grandidier, 1872, Massif de l'Ankaratra, Madagascar.

other common names

English: Campan's chameleon; Malagasy: Kamora, soamarandrana.

physical characteristics

This species attains a length of 4–5.5 in (107–133 mm). There are three longitudinal white or yellowish stripes on each flank, and rows of colored dots cover the body. Red markings outline the ridge above the eyes. The body color in calm females is green; they are black when excited or stressed. Males are brown.

distribution

The species inhabits the high plateaus south of Antananarivo, Madagascar in Ambatolampy, Ambohimitombo, Andringitra, Ankaratra, Antobeba, Ibity, and Manjakatompo.

habitat

The jeweled chameleon inhabits grasslands, shrubs, and savanna and is semiterrestrial.

behavior

This is a small and shy species.

feeding ecology and diet

The jeweled chameleon preys upon small invertebrates.

reproductive biology

The reproductive biology is unknown.

conservation status

The species is classified as Vulnerable by the IUCN based on a 20% population decline in 10 years, or three generations. It is listed on CITES Appendix II and included in the 1995 CITES moratorium on importation, but specimens are known to have been imported for the commercial pet trade at least through 2002.

significance to humans

More than 15,000 wild-taken specimens were exported legally to the commercial pet trade from 1986 to 1996. There was no documented long-term survivability or reproduction in captivity.


Minor chameleon

Furcifer minor

taxonomy

Furcifer minor Gönther, 1879, Fianarantsoa, Betsileo, southeastern Madagascar.

other common names

English: Lesser chameleon; Malagasy: Sakosotoha.

physical characteristics

The species is 6–10 in (140–254 mm) in length, half of which is tail. Males have paired rostral horns, and females are hornless. Coloration in nongravid females is green with faint yellow and pink markings. Gravid females are spectacularly colored with vivid yellow bands and spots against a dark green or black background, with two purple spots adorning the anterior flank and a reddish-pink hue on the top of the head. Males are tan or blue with rust or dark brown stripes and blue eye turrets; they exhibit an orange patch bordered in black at the shoulder. Juveniles are green at birth.

distribution

The species is distributed across south-central Madagascar at elevations up to 5,414 ft (1,650 m) in western Ambositra, Ambatofinandrahana, Ambatomenaloha, Itremo, and northern Vinanitelo.

habitat

Extensive deforestation within the known range of distribution has reduced the habitat of this species to sparse anthropogenic vegetation, including fruit trees and bushes in close proximity to human habitation.

behavior

These chameleons may be aggressive toward conspecifics, but they are otherwise docile.

feeding ecology and diet

The minor chameleon preys upon invertebrates.

reproductive biology

Females lay 10–16 eggs as many as three times annually; eggs incubate approximately nine months before the young emerge.

conservation status

The species is classified as Vulnerable by the IUCN based on a 20% population decline in 10 years, or three generations. It is listed on CITES Appendix II and included in the 1995 CITES moratorium on importation, but specimens are known to have been imported for the commercial pet trade through 2002.

significance to humans

Before 1993, net exports of wild-taken specimens equaled five, but 2,400 of the species had been harvested for the pet trade before the 1995 CITES moratorium on importation went into effect. Limited captive reproduction occurred with a second generation of offspring bred in captivity, but by 1998 no legal specimens remained alive in captivity.


Panther chameleon

Furcifer pardalis

taxonomy

Furcifer pardalis Cuvier, 1829, Madagascar.

other common names

French: Caméléon panthère; German: Pantherchamäleon; Malagasy: Amboalava, tana.

physical characteristics

This species grows to 8–17 in (305–381 mm) in length. The coloration of adult males can vary significantly between geographic locales, but usually it includes two or three predominant colors from the following palette: pink, red, maroon, orange, yellow, green, turquoise, cobalt blue, brown, tan, gray, black, and white. Excited coloration may produce additional colors on various parts of the body that are not obvious in a calm male. There is a broken white, bluish, or cream lateral stripe midflank. Regardless of geographic locale, females are shades of reddish orange or tan marked with brown or black. Juveniles are reddish brown or gray and black at birth. These color variants are not considered valid subspecies as of 2002, but there is anecdotal evidence that interbreeding specimens from distant locales results in sterile offspring. The snout terminates in a short bony process that is more prominent in males than females, and the chameleons possess complete dorsal, ventral, and gular crests.

distribution

The species occupies roughly the northern third of Madagascar, from Toamasina on the central-eastern coast to Ambanizana in the Masoala Peninsula, Sambava on the northeastern coast, Antsiranana in the north, and Ankaramy on the northwestern coast. They also inhabit islets adjacent to Madagascar: Nosy Be, Nosy Komba, Nosy Tanikely, Nosy Faly, Nosy Mangabe, and Nosy Boraha. They have been introduced to the island of Réunion.

habitat

The species is arboreal and primarily inhabits degraded or anthropogenic vegetation, including forest and scrub as well as the undisturbed edges of rainforest on Nosy Be, Nosy Mangabe, and the Masoala Peninsula.

behavior

Males are active and highly mobile and frequently are seen crossing roads. Females are more sedentary and secretive. The species is not particularly aggressive toward conspecifics or humans.

feeding ecology and diet

The panther chameleon feeds on invertebrate and vertebrate prey typical for medium-size to large chameleons as well as some vegetation.

reproductive biology

Panther chameleons lay 10–46 eggs about 45 days after copulation and can produce more than four clutches of eggs annually. The young hatch four to nine months later, depending on climatic conditions. Growth is rapid, and sexual maturity occurs between the ages of six and nine months.

conservation status

This species is listed on Appendix II of CITES. Before 1999 commercial trade was unrestricted, and more than 34,000 wild-taken specimens were exported from Madagascar in 1998 alone, raising concerns that local populations of distinctive color variants would be extirpated. An annual export quota of 2,000 specimens was established by CITES in 1999, to reduce trade volume. Conservation is not dependent on preserving natural habitat, owing to the mostly successful adaptation of this species to heavily degraded habitat.

significance to humans

The panther chameleon is captured extensively for the international commercial pet trade. More than 100,000 specimens were exported between 1986 and 1999. Captive breeding programs experienced limited success by producing several generations, but there were no established commercial closed-cycle operations as of 2002 where only captive-born adult animals lay eggs to provide the stock that is subsequently sold.


Short-tailed chameleon

Rhampholeon brevicaudatus

taxonomy

Rhampholeon brevicaudatus Matschie, 1892, Derema, Usambara Mountains, Tanganyika (Tanzania).

other common names

English: Bearded pygmy chameleon.

physical characteristics

The species is 2–3 in (51–76 mm) in length, most of which is the snout-to-vent length. Females are brownish-orange, tan, or yellow, and males are cream or brown with darker brown stripes. The interior of the mouth is blue-gray. A fleshy spike protrudes from the chin, and the tail is very short in females and slightly longer, with a prominent hemipenial bulge, in males.

distribution

This species is found in Derema in the Usambara Mountains of East Tanzania, Africa, at elevations up to 4,200 ft (1,300 m).

habitat

The short-tailed chameleon lives in grasses, bushes, and leaf litter in humid forests. It is semiterrestrial.

behavior

This is one of a number of species that vibrate rapidly and audibly when they are disturbed or stressed, which may act as a deterrent to predators.

feeding ecology and diet

The short-tailed chameleon preys upon very small invertebrates.

reproductive biology

Males have been observed courting females several days before mating. Male courtship display may include pursuing the receptive female, rearing up on the hind legs, and approaching the female with a rhythmic gate. Copulation lasts only a few minutes, but it can be repeated several times over subsequent days. Clutch size is two to five eggs, which are laid in a shallow pit after a gestation period of three to four weeks. The young emerge about three months later.

conservation status

This species receives no formal protection from CITES to prevent unsustainable trade. Habitat status and population densities were unknown as of 2002.

significance to humans

The short-tailed chameleon has been captured for the commercial pet trade since at least 1990, but recording export quantities has not been required, and the total number exported is unknown as of 2002.


Resources

Books

Brady, L. D., and R. A. Griffiths. Status Assessment of Chameleons in Madagascar. Cambridge: International Union for the Conservation of Nature and Natural Resources, 1999.

Brygoo, E. R. Faune de Madagascar. Vol. 33, Reptiles Sauriens Chamaeleonidae: Genre Chamaeleo. Paris: ORSTOM et CNRS, 1971.

——. Faune de Madagascar. Vol. 47, Reptiles Sauriens Chamaeleonidae: Genre Brookesia et Complement pour le Genre Chamaeleo. Paris: ORSTOM et CNRS, 1978.

Franke, Joseph, and T. M. Telecky. Reptiles as Pets: An Examination of the Trade in Live Reptiles in the United States. Washington DC: Humane Society, 2001.

Glaw, F., and M. Vences. A Fieldguide to the Amphibians and Reptiles of Madagascar. 2nd ed. Cologne: M. Vences & F. Glaw, 1994.

Klaver, Charles J., and Wolfgang Böhme. A Compilation and Characterization of the Recent Animal Groups: Chamaeleonidae. Part 112. Berlin, New York: Das Tierreich, 1997.

Mellado, J., L. Gimenez, J. J. Gomez, et al. El Camaleón en Andalucía: Distribución actual y amenazas para su supervivencia. Rota: Fundacion Alcalde Zoilo Ruiz-Mateos, 2001.

Nečas, P. Chameleons: Nature's Hidden Jewels. Malabar, FL: Krieger Publishing Co., 1999.

Periodicals

Abate, Ardith. "Assessing the Health of Wild-Caught Chameleons." Chameleon Information Network 31 (Spring 1999): 9–17.

——. "Reports from the Field: Parson's Chameleon." Chameleon Information Network 29 (Fall 1998): 17–25.

——. "The Exportation of Chameleons from Madagascar: Past and Present." Chameleon Information Network 32 (Summer 1999): 9–17.

——. "The Fate of Wild-Caught Chameleons Exported for the Pet Trade." Chameleon Information Network 42 (Winter 2001): 15–18.

Abate, Ardith, and K. Kalisch "Chamaeleon Profile: Chamaeleo (T.) jacksonii." Chameleon Information Network 14 (Winter 1994): 19–28.

Deas, J. "In Search of the Veiled or Yemeni Chameleon, Chamaeleo calyptratus calyptratus." Chameleon Information Network 43 (Spring 2002): 10–20.

Dimaki, M. "Reports from the Field: The European Chameleons." Chameleon Information Network 41 (Fall 2001): 11–13.

Green, G. M., and R. W. Sussman. "Deforestation History of Eastern Rain Forests of Madagascar from Satellite Images." Science 248 (April 1990): 212–215.

Herrel, A., J. J. Meyers, P. Aerts, et al. "The Mechanics of Prey Prehension in Chameleons." Journal of Experimental Biology 203 (2000): 3255–3263.

Kalisch, K. "The Captive Care and Breeding of Chamaeleo [Calumma] parsonii cristifer." Chameleon Information Network 27 (Spring 1998): 15–17.

Raxworthy, Christopher J., and Ronald A. Nussbaum. "Extinction and Extinction Vulnerability of Amphibians and Reptiles in Madagascar." Amphibian and Reptile Conservation 2, no. 1 (2000): 15–23.

Risley, T. "Chameleon Profile: Brookesia and Rhampholeon." Chameleon Information Network 31 (Spring 1999): 21–30.

——. "Field Observations on the Panther Chameleon, Chamaeleo [Furcifer] pardalis Cuvier, 1829." Chameleon Information Network 24 (Summer 1997): 17–29.

——. "Preliminary Observations on a Small Population of Chamaeleo [Calumma] parsonii parsonii Cuvier, 1824 in Eastern Madagascar." Chameleon Information Network 25 (Fall 1997): 25–28.

Organizations

The Chameleon Information Network. 13419 Appalachian, Way, San Diego, CA 92129 USA. Phone: (858) 484-2669. Fax: (858) 484-4757. E-mail: [email protected] Web site: <http://www.animalarkshelter.org/cin>

World Conservation Union (IUCN). Rue Mauverney 28, Gland, 1196 Switzerland. Phone: 41 (22) 999-0000. Fax: 41 (22) 999-0002. E-mail: [email protected] Web site: <http://www.iucn.org>

Other

Convention on International Trade in Endangered Species (CITES). "Notification to the Parties No. 833.: Significant Trade in Animal Species Included in Appendix II. Recommendations of the Standing Committee." [cited June 8, 2002]. <http://www.cites.org>

United Nations Environmental Program—World Conservation Monitoring Centre (UNEP-WCMC). "Significant Trade in Animals, Net Trade Outputs 1994–1999." Report directed to the CITES Animals Committee regarding the implementation of Resolution Conf. 8.9. [cited June 8, 2002]. <http://www.cites.org/eng/cttee/animals/17/sigtrade2001.shtml>

UNEP/WCMC/UNESCO. "World Heritage Sites. Tsingy de Bemaraha Strict Nature Reserve." ([cited June 8, 2002]. <http://www.wcmc.org.uk/protected_areas/data/wh/bemaraha.html>

World Wildlife Fund (WWF). "Maputaland-Pondoland Bushland and Thickets (AT1012)." [cited June 8, 2002]. <http://www.worldwildlife.org/wildworld/profiles/terrestrial/at/at1012_full.html>

Ardith L. Abate

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