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Chameleons or chamaeleons (
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Chamaeleonidae) are a distinctive and highly specialized
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
of
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by thei ...
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...
s with 202 species described as of June 2015. The members of this family are best known for their distinct range of colors, being capable of shifting to different hues and degrees of brightness. The large number of species in the family exhibit considerable variability in their capacity to change color. For some, it is more of a shift of brightness (shades of brown); for others, a plethora of color-combinations (reds, yellows, greens, blues) can be seen. Chameleons are distinguished by their zygodactylous feet, their
prehensile Prehensility is the quality of an appendage or organ that has adapted for grasping or holding. The word is derived from the Latin term ''prehendere'', meaning "to grasp". The ability to grasp is likely derived from a number of different orig ...
tail, their laterally compressed bodies, their head casques, their projectile tongues, their swaying gait, and crests or horns on their brow and snout. Chameleons' eyes are independently mobile, and because of this there are two separate, individual images that the brain is analyzing of the chameleon’s environment. When hunting prey, they focus forward in coordination, affording the animal stereoscopic vision. Chameleons are adapted for climbing and visual hunting. The use of their prehensile tail offers stability when they are moving or resting while on a branch in the canopy; because of this, their tail is often referred to as a "fifth limb." Another character that is advantageous for being arboreal is how laterally compressed their bodies are; it is important for them to distribute their weight as evenly as possible as it confers stability on twigs and branches in the trees. They live in warm habitats that range from
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
to
desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
conditions, with various species occurring in Africa,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
, southern Europe, and across southern Asia as far as
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
. They have been introduced to
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,
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
, and
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and ...
.


Etymology

The English word ''chameleon'' ( ) is a simplified spelling of
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
', a borrowing of the Greek ' (''khamailéōn''), a compound of ' (''khamaí'') "on the ground" and ' (''léōn'') "lion".


Classification

In 1986, the family Chamaeleonidae was divided into two subfamilies, Brookesiinae and Chamaeleoninae. Under this classification, Brookesiinae included the genera '' Brookesia'' and '' Rhampholeon'', as well as the genera later split off from them ('' Palleon'' and '' Rieppeleon''), while Chamaeleoninae included the genera '' Bradypodion'', '' Calumma'', '' Chamaeleo'', '' Furcifer'' and '' Trioceros'', as well as the genera later split off from them ('' Archaius'', '' Nadzikambia'' and '' Kinyongia''). Since that time, however, the validity of this subfamily designation has been the subject of much debate, although most phylogenetic studies support the notion that the pygmy chameleons of the subfamily Brookesiinae are not a
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
group. While some authorities have previously preferred to use this subfamilial classification on the basis of the absence of evidence principle, these authorities later abandoned this subfamilial division, no longer recognizing any subfamilies with the family Chamaeleonidae. In 2015, however, Glaw reworked the subfamilial division by placing only the genera '' Brookesia'' and '' Palleon'' within the Brookesiinae subfamily, with all other genera being placed in Chamaeleoninae.


Change of color

Some chameleon species are able to change their skin coloration. Different chameleon species are able to vary their coloration and pattern through combinations of pink, blue, red, orange, green, black, brown, light blue, yellow, turquoise, and purple. Chameleon skin has a superficial layer which contains pigments, and under the layer are cells with very small (nanoscale)
guanine Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine ( uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is ...
crystals. Chameleons change color by "actively tuning the photonic response of a lattice of small guanine nanocrystals in the s-iridophores". This tuning, by an unknown molecular mechanism, changes the wavelength of light reflected off the crystals which changes the color of the skin. The color change was duplicated '' ex vivo'' by modifying the osmolarity of pieces of white skin. Color change in chameleons has functions in
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
, but most commonly in social signaling and in reactions to temperature and other conditions. The relative importance of these functions varies with the circumstances, as well as the species. Color change signals a chameleon's physiological condition and intentions to other chameleons. Because chameleons are
ectotherm An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life ...
ic, another reason why they change color is to regulate their body temperatures, either to a darker color to absorb light and heat to raise their temperature, or to a lighter color to reflect light and heat, thereby either stabilizing or lowering their body temperature. Chameleons tend to show brighter colors when displaying aggression to other chameleons, and darker colors when they submit or "give up". Some species, particularly those of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
and some African genera in rainforest habitats, have blue
fluorescence Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, tha ...
in their skull
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s, deriving from bones and possibly serving a signaling role. Some species, such as
Smith's dwarf chameleon Smith's dwarf chameleon or the Elandsberg dwarf chameleon (''Bradypodion taeniabronchum'') is a species of lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, ...
, adjust their colors for camouflage by the vision of the specific predator species (bird or snake) by which they are being threatened. Chameleons have two superimposed layers within their skin that control their color and thermoregulation. The top layer contains a lattice of guanine nanocrystals, and by exciting this lattice the spacing between the nanocrystals can be manipulated, which in turn affects which wavelengths of light are reflected and which are absorbed. Exciting the lattice increases the distance between the nanocrystals, and the skin reflects longer wavelengths of light. Thus, in a relaxed state the crystals reflect blue and green, but in an excited state the longer wavelengths such as yellow, orange, green, and red are reflected. The skin of a chameleon also contains some yellow pigments, which combined with the blue reflected by a relaxed crystal lattice results in the characteristic green color which is common of many chameleons in their relaxed state. Chameleon color palettes have evolved through evolution and the environment. Chameleons living in the forest have a more defined and colorful palette compared to those living in the desert or savanna, which have more of a basic, brown, and charred palette.


Evolution

The oldest described chameleon is '' Anqingosaurus brevicephalus'' from the Middle Paleocene (about 58.7–61.7 mya) of China. Other chameleon fossils include '' Chamaeleo caroliquarti'' from the Lower Miocene (about 13–23 mya) of the Czech Republic and Germany, and ''Chamaeleo intermediate'' from the Upper Miocene (about 5–13 mya) of Kenya. The chameleons are probably far older than that, perhaps sharing a common ancestor with iguanids and agamids more than 100 mya (agamids being more closely related). Since fossils have been found in Africa, Europe, and Asia, chameleons were certainly once more widespread than they are today. Although nearly half of all chameleon species today live in Madagascar, this offers no basis for speculation that chameleons might originate from there. In fact, it has recently been shown that chameleons most likely originated in mainland Africa. It appears there were two distinct oceanic migrations from the mainland to Madagascar. The diverse speciation of chameleons has been theorized to have directly reflected the increase in open habitats (savannah, grassland, and heathland) that accompanied the Oligocene period.
Monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
of the family is supported by several studies. Daza ''et al.'' (2016) described a small (10.6 mm in snout-vent length), probably neonatal lizard preserved in the Cretaceous ( Albian-
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...
boundary) amber from
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John Wells explai ...
. The authors noted that the lizard has "short and wide skull, large orbits, elongated and robust lingual process, frontal with parallel margins, incipient prefrontal boss, reduced vomers, absent retroarticular process, low presacral vertebral count (between 15 and 17) and extremely short, curled tail"; the authors considered these traits to be indicative of the lizard's affiliation with Chamaeleonidae. The phylogenetic analysis conducted by the authors indicated that the lizard was a
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
-chamaeleonid. However, Matsumoto &
Evans Evans may refer to: People *Evans (surname) * List of people with surname Evans Places United States *Evans Island, an island of Alaska * Evans, Colorado *Evans, Georgia *Evans County, Georgia *Evans, New York *Evans Mills, New York *Evans Cit ...
(2018) reinterpreted this specimen as an albanerpetontid amphibian. This specimen was given the name ''
Yaksha perettii ''Yaksha perettii'' is an extinct species of albanerpetontid amphibian, and the only species in the genus ''Yaksha''. It is known from three specimens found in Cenomanian aged Burmese amber from Myanmar. The remains of ''Yaksha perettii'' are t ...
'' in 2020, and was noted to have several convergently chameleon-like features, including adaptations for ballistic feeding. While the exact evolutionary history of color change in chameleons is still unknown, there is one aspect of the evolutionary history of chameleon color change that has already been conclusively studied: the effects of signal efficacy. Signal efficacy, or how well the signal can be seen against its background, has been shown to correlate directly to the spectral qualities of chameleon displays. Dwarf chameleons, the chameleon of study, occupy a wide variety of habitats from forests to grasslands to shrubbery. It was demonstrated that chameleons in brighter areas tended to present brighter signals, but chameleons in darker areas tended to present relatively more contrasting signals to their backgrounds. This finding suggests that signal efficacy (and thus habitat) has affected the evolution of chameleon signaling. Stuart-Fox et al. note that it makes sense that selection for crypsis is not seen to be as important as selection for signal efficacy, because the signals are only shown briefly; chameleons are almost always muted cryptic colors.


Description

Chameleons vary greatly in size and body structure, with maximum total lengths varying from in male ''
Brookesia micra ''Brookesia micra'', also known as the Nosy Hara leaf chameleon is a species of chameleons from the islet of Nosy Hara in Antsiranana, Madagascar. At the time of its discovery, it was the smallest known chameleon and among the smallest reptile ...
'' (one of the world's smallest reptiles) to 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. Typical sizes of species of chameleon commonly kept in captivity or as pets are: The feet of chameleons are highly adapted to
arboreal locomotion Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
, 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 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 In biology, dactyly is the arrangement of digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. It comes from the Greek word δακτυλος (''dáktylos'') = "finger". Sometimes the ending "-dactylia" is used ...
or zygodactyl, though neither term is fully satisfactory, both being used in describing 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 Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...
part of the tail to the neck; both the extent and size of the spikes vary 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.


Senses

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 Depth perception is the ability to perceive distance to objects in the world using the visual system and visual perception. It is a major factor in perceiving the world in three dimensions. Depth perception happens primarily due to stereopsis ...
, not stereopsis. Chameleons have the highest magnification (per size) of any vertebrate. Like
snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more ...
s, 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 Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiatio ...
. Chameleons exposed to ultraviolet light show increased social behavior and activity levels, are more inclined to bask, feed, and reproduce as it has a positive effect on the
pineal gland The pineal gland, conarium, or epiphysis cerebri, is a small endocrine gland in the brain of most vertebrates. The pineal gland produces melatonin, a serotonin-derived hormone which modulates sleep patterns in both circadian and seasonal cy ...
.


Feeding

All chameleons are primarily
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
s that feed by ballistically projecting their long
tongue The tongue is a muscular organ in the mouth of a typical tetrapod. It manipulates food for mastication and swallowing as part of the digestive process, and is the primary organ of taste. The tongue's upper surface (dorsum) is covered by taste ...
s 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 tongue apparatus consists of highly modified
hyoid bones The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra ...
, 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 Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperatur ...
, when other
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...
lizards species are still inactive, likely temporarily expanding their thermal
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development * Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
as a result. File:Chameleon gab fbi.png, Tongue structure, with cup-like end File:Chameleon-Stage 04.jpg, Tongue begins strike File:Chameleon-Stage 03.jpg, Capturing prey File: Chameleon-Stage 01.jpg, Bringing prey to the mouth


Bones

Certain species of chameleons have bones that glow when under
ultraviolet light Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiatio ...
, also known as
biogenic A biogenic substance is a product made by or of life forms. While the term originally was specific to metabolite compounds that had toxic effects on other organisms, it has developed to encompass any constituents, secretions, and metabolites of p ...
fluorescence Fluorescence is the emission of light by a substance that has absorbed light or other electromagnetic radiation. It is a form of luminescence. In most cases, the emitted light has a longer wavelength, and therefore a lower photon energy, tha ...
. Some 31 different species of ''Calumma'' chameleons, all native to
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
, displayed this fluorescence in CT scans. The bones emitted a bright blue glow and could even shine through the chameleon's four layers of skin. The face was found to have a different glow, appearing as dots otherwise known as
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s on facial bones. The glow results from
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s, pigments,
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, and other materials that make up a chameleon's skeleton, possibly giving chameleons a secondary signaling system that does not interfere with their color-changing ability, and may have evolved from
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
.


Distribution and habitat

Chameleons primarily live in the mainland of
sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
and on the island of Madagascar, although a few species live in northern Africa,
southern Europe Southern Europe is the southern region of Europe. It is also known as Mediterranean Europe, as its geography is essentially marked by the Mediterranean Sea. Definitions of Southern Europe include some or all of these countries and regions: Alb ...
(Portugal, Spain, Italy, Greece), the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
, southern
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area, the List of countries and dependencies by population, second-most populous ...
,
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
, and several smaller islands in the
western Indian Ocean Western may refer to: Places *Western, Nebraska, a village in the US *Western, New York, a town in the US *Western Creek, Tasmania, a locality in Australia *Western Junction, Tasmania, a locality in Australia *Western world, countries that id ...
. Chameleons inhabit all kinds of
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and mountain rain forests,
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground ...
s, and sometimes
desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
s and
steppe In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate gras ...
s. 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 Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
. Many species of chameleons are threatened by extinction. Declining chameleon numbers are due to habitat loss.


Reproduction

Chameleons are mostly
oviparous Oviparous animals are animals that lay their eggs, with little or no other embryonic development within the mother. This is the reproductive method of most fish, amphibians, most reptiles, and all pterosaurs, dinosaurs (including birds), and m ...
, with some being ovoviviparous. The oviparous species lay eggs three to six weeks after
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
. The female will dig a hole—from , 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 chameleon The veiled chameleon (''Chamaeleo calyptratus'') is a species of chameleon ( family Chamaeleonidae) native to the Arabian Peninsula in Yemen and Saudi Arabia. Other common names include cone-head chameleon and Yemen chameleon. They are born pa ...
s (''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 the species. The eggs of
Parson's chameleon Parson's chameleon (''Calumma parsonii'') is a large species of chameleon in the family Chamaeleonidae. The species is endemic to isolated pockets of humid primary forest in eastern and northern Madagascar. It is listed on CITES Appendix II, ...
(''Calumma parsonic''), a species that is rare in captivity, are believed to take more than 24 months to hatch. Chameleons lay flexible-shelled eggs which are affected by environmental characteristics during incubation. The egg mass is the most important in differentiating survivors of Chameleon during incubation. An increase in egg mass will depend on temperature and water potential. To understand the dynamics of water potential in Chameleon eggs, the consideration of exerted pressure on eggshells will be essential because the pressure of eggshells play an important role in the water relation of eggs during entire incubation period 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.


Diet

Chameleons generally eat
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s, but larger species, such as the common chameleon, may also take other lizards and young
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s. The range of diets can be seen from the following examples: * The veiled chameleon, '' Chamaeleo calyptratus'' from
Arabia The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Pl ...
, 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 eat 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 ''Chamaeleo'' is a genus of chameleons in the family Chamaeleonidae. Most species of the genus ''Chamaeleo'' are found in sub-Saharan Africa, but a few species are also present in northern Africa, southern Europe, and southern Asia east to Indi ...
'', mainly eats
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
s and mantises; such
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s 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,
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
s,
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s, flies, and plant materials such as green leaves, oats, and fruit. * Some chameleons like the
panther chameleon The panther chameleon (''Furcifer pardalis'') is a species of chameleon found in the eastern and northern parts of Madagascar in a tropical forest biome. Additionally, it has been introduced to Réunion and Mauritius. Taxonomy The Nosy Be pa ...
of
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
regulate their
vitamin D3 Cholecalciferol, also known as vitamin D3 and colecalciferol, is a type of vitamin D that is made by the skin when exposed to sunlight; it is found in some foods and can be taken as a dietary supplement. Cholecalciferol is made in the skin ...
levels, of which their insect diet is a poor source, by exposing themselves to
sunlight Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when ...
since its UV component increases internal production.


Anti-predator adaptations

Chameleons are preyed upon by a variety of other animals.
Birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
and
snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more j ...
are the most important predators of adult chameleons. Invertebrates, especially ants, put a high predation pressure on chameleon eggs and juveniles. Chameleons are unlikely to be able to flee from predators and rely on
crypsis In ecology, crypsis is the ability of an animal or a plant to avoid observation or detection by other animals. It may be a predation strategy or an antipredator adaptation. Methods include camouflage, nocturnality, subterranean lifestyle an ...
as their primary defense. Chameleons can change both their colors and their patterns (to varying extents) to resemble their surroundings or disrupt the body outline and remain hidden from a potential enemy's sight. Only if detected, chameleons actively defend themselves. They adopt a defensive body posture, present an attacker with a laterally flattened body to appear larger, warn with an open mouth, and, if needed, utilize feet and jaws to fight back. Vocalization is sometimes incorporated into threat displays. File:Chameleon in malawi 2018-6.webm, (video) Chameleon in
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...
File:Chameleon, Mysore.jpg, Chameleon found in
Mysore Mysore (), officially Mysuru (), is a city in the southern part of the state of Karnataka, India. Mysore city is geographically located between 12° 18′ 26″ north latitude and 76° 38′ 59″ east longitude. It is located at an altitude o ...
, Southern India File:Chamaeleo chamaeleon Frightened thus black.JPG, This common chameleon (''Chamaeleo chamaeleon'') turned black. File: Chameleon-Boomslang-Interaction P Berg.jpg, alt=A flap-necked chameleon defending itself against a boomslang attack, A flap-necked chameleon, ''Chamaeleo dilepis'', attacked by a
boomslang The boomslang (, , or ; ''Dispholidus typus'') is a large, highly venomous snake in the family Colubridae. Taxonomy and etymology Its common name means "tree snake" in Afrikaans and Dutch – ''boom'' meaning "tree", and ''slang'' meaning "sna ...
while crossing a road in Namibia adopts a threatening defense posture. File: Chamaeleo namaquensis (Namib-Naukluft, 2011).jpg, Namaqua chameleon in threat display, Namib-Naukluft National Park, turned black and opened its mouth when an attempt was made to move it off a busy road.


Parasites

Chameleons are parasitized by
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
worms, including threadworms ( Filarioidea). Threadworms can be transmitted by biting insects such as ticks and mosquitoes. Other roundworms are transmitted through food contaminated with roundworm eggs; the larvae burrow through the wall of the
intestine The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans an ...
into the bloodstream. Chameleons are subject to several protozoan parasites, such as ''Plasmodium'', which causes
malaria Malaria is a mosquito-borne infectious disease that affects humans and other animals. Malaria causes symptoms that typically include fever, tiredness, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. ...
, ''Trypanosoma'', which causes sleeping sickness, and ''Leishmania'', which causes
leishmaniasis Leishmaniasis is a wide array of clinical manifestations caused by parasites of the trypanosome genus '' Leishmania''. It is generally spread through the bite of phlebotomine sandflies, ''Phlebotomus'' and ''Lutzomyia'', and occurs most freq ...
. Chameleons are subject to parasitism by
coccidia Coccidia (Coccidiasina) are a subclass of microscopic, spore-forming, single-celled obligate intracellular parasites belonging to the apicomplexan class Conoidasida. As obligate intracellular parasites, they must live and reproduce within an a ...
,Le Berre and Bartlett, p. 109 including species of the genera ''
Choleoeimeria ''Choleoeimeria'' is a genus of alveolate parasites that infect the biliary tracts of reptiles. Morphologically they are similar to the ''Eimeria'', to whom they are closely related. The genus was described in 1989 by Paperna and Landsberg. Gen ...
'', '' Eimeria'', and '' Isospora''.


As pets

Chameleons are popular reptile pets mostly imported from African countries like Madagascar, Tanzania, and Togo. The most common in the trade are the Senegal chameleon (''Chamaeleo senegalensis''), veiled chameleon (''Chamaeleo calyptratus''), panther chameleon (''Furcifer pardalis''), and Jackson's chameleon (''Trioceros jacksonii''). The U.S. has been the main importer of chameleons since the early 1980s accounting for 69% of African reptile exports. However, there have been large declines due to tougher regulations to protect species from being taken from the wild and due to many becoming invasive in places like Florida. They have remained popular though which may be due to the captive-breeding in the U.S. which has increased to the point that the U.S. can fulfill its demand, and has now even become a major exporter as well. In the U.S. they are so popular, that despite Florida having six invasive chameleon species due to the pet trade, reptile hobbyists in these areas search for chameleons to keep as pets or to breed and sell them, with some selling for up to a thousand dollars.


Historical understandings

Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
(4th century BC) describes chameleons in his
History of Animals ''History of Animals'' ( grc-gre, Τῶν περὶ τὰ ζῷα ἱστοριῶν, ''Ton peri ta zoia historion'', "Inquiries on Animals"; la, Historia Animalium, "History of Animals") is one of the major Aristotle's biology, texts on biolo ...
.
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
(1st century AD) also discusses chameleons in his Natural History, noting their ability to change color for camouflage. The chameleon was featured in Conrad Gessner's ''Historia animalium'' (1563), copied from ''De aquatilibus'' (1553) by Pierre Belon. In Shakespeare's ''Hamlet'', the eponymous Prince says "Excellent, i' faith, of the chameleon's dish. I eat the air, promise-crammed." This refers to the Elizabethan belief that chameleons lived on nothing but the air.


References


General bibliography

* Le Berre, François; Bartlett, Richard D. (2009). ''The Chameleon Handbook''. Barron's Educational Series. 3rd Edition. .


Further reading

* * * * * Davison, Linda J. ''Chameleons: Their Care and Breeding.'' Hancock House Publishers, 1997. * * de Vosjoli, Philippe. ''Essential Care of Chameleons.'' Advanced Vivarium Systems, 2004. *


External links

* * {{Authority control Lizards of Africa Afrotropical realm fauna Articles containing video clips Extant Paleocene first appearances Taxa named by Constantine Samuel Rafinesque