Chameleons or chamaeleons (family Chamaeleonidae) are a distinctive and highly specialized clade of Old World with 200 species described as of June 2015. The members of this family are best known for their distinct range of colours, being capable of colour-shifting camouflage. The large number of species in the family exhibit considerable variability in their capacity to change colour. For some, it is more of a shift of brightness (shades of brown); for others, a plethora of colour-combinations (reds, yellows, greens, blues) can be seen.
Chameleons are also distinguished by their feet, their prehensility tail, their laterally compressed bodies, their head casques, their projectile tongues used for catching prey, their swaying gait, and in some species crests or horns on their brow and snout. Chameleons' eyes are independently mobile, and because of this the chameleon's brain is constantly analyzing two separate, individual images of its environment. When hunting prey, the eyes focus forward in coordination, affording stereoscopic vision. Chameleons are Diurnality and adapted for visual hunting of invertebrates, mostly insects, although the large species also can catch small vertebrates. Chameleons typically are arboreal, but there are also many species that live on the ground. The arboreal species use their prehensile tail as an extra anchor point when they are moving or resting in trees or bushes; because of this, their tail is often referred to as a "fifth limb". Depending on species, they range from rainforest to desert conditions and from lowlands to highlands, with the vast majority occurring in Africa (about half of the species are restricted to Madagascar), but with a single species in southern Europe, and a few across southern Asia as far east as India and Sri Lanka. They have been introduced to Hawaii and Florida.
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.
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(a) Reversible colour change is shown for two males (m1 and m2): during excitation (white arrows), background skin shifts from the baseline state (green) to yellow/orange, and both vertical bars and horizontal mid-body stripe shift from blue to whitish (m1). Some animals (m2) have their blue vertical bars covered by red pigment cells.
(b) Red dots: time evolution in the CIE chromaticity chart of a third male with green skin in a high-resolution video; dashed white line: optical response in numerical simulations using a face-centred cubic (FCC) lattice of guanine crystals with lattice parameter indicated with black arrows.
(c) Haematoxylin and eosin staining of a cross-section of white skin showing the epidermis (ep) and the two thick layers of iridophores.
(d) TEM images of guanine nanocrystals in S-iridophores in the excited state and three-dimensional model of an FCC lattice (shown in two orientations).
(e) TEM image of guanine nanocrystals in D-iridophores.
Scale bars, 20 mm (c); 200 nm (d, e).]]
Colour change in chameleons has functions in camouflage, but most commonly in social signalling and reactions to temperature and other conditions. The relative importance of these functions varies with the circumstances, as well as the species. Colour change signals a chameleon's physiological condition and intentions to other chameleons. Because chameleons are , another reason why they change colour is to regulate their body temperatures, either to a darker colour to absorb light and heat to raise their temperature, or to a lighter colour to reflect light and heat, thereby either stabilizing or lowering their body temperature. Chameleons tend to show brighter colours when displaying aggression to other chameleons, and darker colours when they submit or "give up". Most chameleon genera (exceptions are Chamaeleo, Rhampholeon and Rieppeleon) have blue fluorescence in a species-specific pattern in their skull , and in Brookesia there is also some in tubercles on the body. The fluorescence is derived from bones that only are covered in very thin skin and possibly serves a signaling role, especially in shaded habitats.
Some species, such as Smith's dwarf chameleon and several others in the genus Bradypodion, adjust their colours for camouflage depending on the vision of the specific predator species (for example, bird or snake) by which they are being threatened.Young, Emma (2008) Chameleons fine-tune camouflage to predator's vision. New Scientist In the introduced Hawaiian population of Jackson's chameleon, conspicuous colour changes that are used for communication between chameleons have increased, whereas anti-predator camouflage colour changes have decreased relative to the native source population in Kenya, where there are more predators.
Chameleons have two superimposed layers within their skin that control their colour 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 colour, which is common for many chameleons in their relaxed state. Chameleon colour palettes have evolved through evolution and the environment. Chameleons living in the forest have a more defined and colourful palette compared to those living in the desert or savanna, which have more of a basic, brown, and charred palette.
The chameleons are probably far older than that, perhaps sharing a common ancestor with Iguanidae 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.
Daza et al. (2016) described a small (10.6 mm in snout-vent length), probably neonatal lizard preserved in the Cretaceous (Albian-Cenomanian boundary) amber from Myanmar. 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-chamaeleonid. However, Matsumoto & Evans (2018) reinterpreted this specimen as an albanerpetontid amphibian. This specimen was given the name Yaksha perettii in 2020, and was noted to have several convergently chameleon-like features, including adaptations for ballistic feeding. While the exact evolutionary history of colour change in chameleons is still unknown, there is one aspect of the evolutionary history of chameleon colour 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 colours.
Typical sizes of species of chameleon commonly kept in captivity or as pets are:
about 5 |
5–10 |
about 5 (2–3 for birthing females) |
about 3–5 |
3–5 |
5–11 |
The feet of chameleons are highly adapted to arboreal locomotion, and species such as Chamaeleo namaquensis that have secondarily adopted a terrestrial habit have retained the same foot morphology with little modification. On each foot, the five 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 Dactyly or Dactyly, 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 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.
Like , 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.Le Berre and Bartlett, p. 31
Chameleons can see in both visible and ultraviolet light. Chameleons exposed to ultraviolet light show increased social behavior and activity levels, are more inclined to bask, feed, and reproduce as it has a positive effect on the pineal gland.
The tongue apparatus consists of highly modified Hyoid, tongue muscles, and Collagen. 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 force. 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 Ectotherm become sluggish as their body temperatures decline, due to a reduction in the contractile velocity of their muscles, chameleons are able to project their tongues at high performance even at low body temperatures. The thermal sensitivity of tongue retraction in chameleons, however, is not a problem, as chameleons have a very effective mechanism of holding onto their prey once the tongue has come into contact with it, including surface phenomena, such as wet adhesion and interlocking, and suction. The thermal insensitivity of tongue projection thus enables chameleons to feed effectively on cold mornings prior to being able to behaviorally elevate their body temperatures through thermoregulation, when other sympatric lizards species are still inactive, likely temporarily expanding their thermal Ecological niche as a result.
Chameleons are found only in tropical and subtropical regions and inhabit all kinds of lowland and mountain forests, woodlands, shrublands, , and sometimes , but each species tends to be a restricted to only one of a few different habitat types. 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. The genus Brookesia, which comprises the majority of the species in the subfamily Brookesiinae, live low in vegetation or on the ground among leaf litter. Many chameleon species have small distributions and are considered threatened. Declining chameleon numbers are mostly due to habitat loss.
The oviparous species lay eggs three to six weeks after copulation. 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 ( 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 ( Calumma parsoni) typically take 400 to 660 days 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.
Chameleons are subject to several protozoan parasites, such as Plasmodium, which causes malaria, Trypanosoma, which causes sleeping sickness, and Leishmania, which causes leishmaniasis.
Chameleons are subject to parasitism by coccidia,Le Berre and Bartlett, p. 109 including species of the genera Choleoeimeria, Eimeria, and Isospora.
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.
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.
As pets
Historical understandings
General bibliography
Further reading
External links
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