|Carolina anole (above),
Crested anole (below)
Dactyloidae are a family of lizards commonly known as anoles (US: // ( listen)) and native to warmer parts of the Americas, ranging from southeastern United States to Paraguay. Instead of treating it as a family, some authorities prefer to treat it as a subfamily, Dactyloinae, of the family Iguanidae. In the past they were included in the family Polychrotidae together with Polychrus (bush anoles), but the latter genus is not closely related to the true anoles.
Anoles are small to fairly large lizards, typically green or brownish, but their color varies depending on species and many can also change it. In most species at least the male has a dewlap, an often brightly colored flap of skin that extents from the throat/neck and is used in displays. Anoles share several characteristics with geckos, including details of the foot structure (for climbing) and the ability to voluntarily break off the tail (to escape predators), but they are only very distantly related, anoles being part of Iguania. Anoles share many anatomical features with tree-living iguanas.
Anoles are active during the day and feed mostly on small animals such as insects, but some will also take fruits, flowers and nectar. They are fiercely territorial. After mating the female lays an egg on the ground and she may repeat this every few weeks.
Anoles are widely studied in fields such as ecology, behavior and evolution, and some species are commonly kept in captivity as pets. Anoles can function as a biological pest control by eating insects that are harmful to plants, but represent a serious risk to small native animals and ecosystems if introduced to regions outside their home range.
Distribution and habitat
Anoles are a very diverse and plentiful group of lizards. They are native to tropical and subtropical South America, Central America, Mexico, the West Indies and southeastern United States. A particularly high species richness exists in Cuba, Hispaniola, Mexico, Central America, Colombia and Ecuador. Fewer live in eastern and central South America (for example, less than 20 species are known from huge Brazil, compared to more than 75 and 40 in Colombia and Ecuador respectively), Contiguous United States (1 native species, compared to Mexico's more than 45), and the Lesser Antilles (about 25 species, compared to more than 60 and 55 in Cuba and Hispaniola respectively). However, the Lesser Antilles are relatively rich compared to their very small land area and their species are all highly localized endemics, each only found on one or a few diminutive islands. In South America, the diversity is considerably higher west of the Andes (Tumbes-Chocó-Magdalena region) than east (Amazon basin), as well illustrated in Ecuador where about 2⁄3 of the anole species live in the former region and 1⁄3 in the latter.
The only species native to the contiguous United States is the Carolina (or green) anole, which ranges at least as far west as central Texas and north to North Carolina. Several anole species have been introduced to the contiguous US, mostly Florida, but also other Gulf Coast states and California. The most prevalent of these introductions is the brown anole. In contrast to the contiguous United States, Puerto Rico and the Virgin Islands are home to 16 native species, all endemic.
Anoles inhabit a wide range of habitats, from highlands, to the coast and rainforest to desert scrub. Some species live close to humans and may use fences or walls of building as perches, even inhabiting gardens or trees along roads in large cities like Miami. Most anoles are arboreal or semi-arboreal, but there are also terrestrial and semi-aquatic species. They are often grouped into several ecomorphs—crown giant, trunk crown, trunk, trunk ground, twig and grass bush—that inhabit specific niches.
Appearance and behavior
Anoles vary in size. Males generally reach a larger size than females, but in a few species it is the other way around. Many anoles are between 8 and 20 cm (3–8 in) in total length, including a tail that often is longer than the head-and-body. Some larger species can surpass 30 cm (12 in). The largest is the knight anole where males can reach about 50 cm (20 in) in length.
Underneath an anole's toes are pads that have several to a dozen flaps of skin (adhesive lamellae) going horizontally and covered in microscopic hairlike protrusions (setae) that allow them to cling to many different surfaces, similar to but not quite as efficient as a gecko. The extent of these structures and clinging ability varies, being more developed in anole species that live high in the tree canopy than ones living at lower levels.
To escape predators, their tails have the ability to break off at special segments, which is known as autotomy. The tail continues to wriggle strongly for some minutes after detaching. Eventually the tail is regenerated. A few semi-aquatic species will attempt to escape from predators by diving into water or running bipedally across it, similar to basilisks. A wide range of animals will eat anoles, such as large spiders, centipedes, predatory katydids, large frogs, lizards (also cannibalism), birds, monkeys and carnivoran mammals. Some species of snakes, notably the Caribbean Alsophis racers, feed mostly on lizards like anoles.
Most anoles are brownish or green, but there are extensive variations depending on the exact species. The majority can change their color depending on things like emotions (for example, aggression or stress), activity level and as a social signal (for example, displaying dominance), but evidence showing that they do it in response to temperature (thermoregulation) or the color of the background (camouflage) is lacking. The extent and variations of this ability differ widely throughout the individual species. For example, the Virginia (or green) anole can change its color from a bright, leafy green to a dull brown color, while the brown anole can only change its shade, ranging from pale gray-brown to very dark brown.
Disregarding color change, minor individual variations in the basic color, mostly related to sex or age, are common. In some anole species this variation is more pronounced and not only related to sex and age. Examples of this are the basic color of the Cayman blue-throated and leopard anoles, which vary geographically, roughly matching the main habitat at a location. In the Puerto Rican giant anole, a species only able to perform minor color changes (essentially lightness/darkness), juveniles are gray-brown and adults typically green, but an uncommon morph maintains a gray-brown color into adulthood.
Many—but not all—anole species have dewlaps, made of erectile cartilage and covered in skin, that extend from their neck/throat areas. The size, shape and colors of the dewlap vary extensively depending on species, and often it differs between the sexes, being smaller (in some essentially absent) and/or less colorful in females. In a few species, including the Carolina and bark anoles, it varies depending on subspecies. The striped anole is the only species where it is asymmetrically colored, being brighter on one side than the other. The dewlap serves as a signal for species recognition (when more than one anole species lives in a region), attracting mates, territoriality, deterring predators and communicating condition.
Most species of anoles exhibit sexual dimorphism, which allows one to discern between males and females fairly easily with the naked eye. This typically involves the overall size (males larger, in some species as much as three times the mass of females), overall color pattern or the dewlap (larger and more colorful in males), but may also involve things like the size of the crest along the nape/back/tail or the shape of the nose. In the green anole, the female is characterized by a pale dorsal stripe extending from the neck to the tail, a generally smaller body, and a smaller head with a shorter snout. Female brown anoles share these characteristics, although their dorsal stripes are often wider, with diamond-shaped or squiggly edges. This stripe may be present sometimes in males, especially young ones not yet fully grown, but it is always fainter with less-defined edges. Some females have small dewlaps (pale and much smaller than those of the males).
Territoriality and breeding
Anoles are highly territorial, and will fan their dewlap, bob their head and perform "push-ups" to scare away potential competitors. If this does not scare off the intruder, a fight proceeds in which the two anoles attempt to bite each other. Females maintain a feeding territory. Males maintain a larger breeding territory, which overlaps with the feeding territory of one or several females. The home range is generally larger in males than in females, and larger in large anole species than in smaller.
The breeding period varies. In species living in highly seasonal regions it is generally relatively short, typically during the wet season. It is prolonged, often even year-round, in species living in regions with less distinct seasons. In many where it is prolonged but not year-round, it begins in the spring and ends in the fall. Males attract females by extending their dewlap and bobbing their heads. The female may mate with multiple males, but is also able to store sperm inside her body for several months. Anole eggs are soft-shelled and yellowish-white. The female lays one egg (occasionally two) on the ground per time, often placed casually among leaf-litter or in a small hole. She produces one egg in each ovary, and may lay an egg every one to four weeks.
Feeding and activity
Anoles are opportunistic feeders, and may attempt to eat any attractive meal that is of the right size. They primarily feed on insects like flies, crickets, caterpillars, moths and butterflies, and arachnids like spiders. Several species will also eat small vertebrates such as mice, small birds (including nestlings), lizards (including other anoles species and their own) and frogs, the slow-moving Cuban "false chameleon" anoles (formerly in genus Chamaeleolis, now considered a part of Xiphosurus) have blunt teeth and are specialized snail-eaters, and a few semi-aquatic species like the Cuban stream anole may catch prey in water such as shrimp and fish. Many will chase down/sneak up to a potential prey item, while others are sit-and-wait predators that pounce on prey when it gets close to the anole. Many anole species will take plant material, notably fruits, flowers and nectar, and overall they are best described as omnivorous. Some fruit-eating species, like the knight anole, may function as seed dispersers. Anoles are vulnerable to drying out and generally need access to water for drinking.
Evolution and adaptability
On the website Anole Annals, Martha Munoz calls attention to a 2012 paper ("Rapid Change in the Thermal Tolerance of a Tropical Lizard") by scientists Alex Gunderson and Manuel Leal of Duke University, in which they examine whether physiology has evolved in the invasive populations of Anolis cristatellus (the Puerto Rican crested anole) in Florida:
Anolis lizards are a model system for studies of evolutionary ecology because they are remarkably adaptable creatures. We know from long-term studies...that anoles can rapidly adapt their behavior and morphology over ecological timescales. For example, the presence of a ground-dwelling predator (Leiocephalus carinatus) forged a strong selective gradient in favor of A. sagrei with longer hindlimbs within a single generation. Interestingly, in a follow-up study the long-term effect of this predator is that A. sagrei evolves shorter hindlimbs, as they will tend to perch higher off the ground, where the perch diameter is narrower than near the ground. ...
A notable exception to Anolis 'evolvability,' however, is thermal physiology... Although native to the Puerto Rico, A. cristatellus has made a permanent home in Florida... In a previous study, Jason Kolbe and colleagues demonstrated that the thermal niches (i.e., available thermal habitat) are quite divergent between Puerto Rico and Florida, and showed a rapid evolutionary response in acclimation to thermal environment in one of the invasive populations. This sets up the possibility that invading anoles will evolve their thermal physiology to match their new environment... [A]lthough thermal physiology is predominantly conserved among lizards, anoles are a notable exception to this rule.
In a recent study from the University of Texas, one species of anole were shown to be able to evolve rapidly in response to the invasion of another species. A. carolinensis are native to the southeastern United States as well as small islands off the coast of Florida. A. sagrei is a species native to Cuba that has been slowly invading A. carolinensis habitats. In response to the presence of A. sagrei, A. carolinensis moved to higher perches and adapted larger toepads better suited for those perches. The adaptation that as observed occurred in just 20 generations. This study showed that anoles have the ability to evolve in a remarkably short time frame in response to changing environmental conditions.
Several family names have been used for the anoles in recent decades. Initially they were placed in Polychrotidae together with Polychrus (bush anoles), but genetic studies have shown that the latter genus is closer to Hoplocercidae than the true anoles. The true anoles are closer to Corytophanidae (basilisks and relatives). The true anoles have therefore been transferred to their own family Dactyloidae, alternatively listed as subfamily Dactyloinae of family Iguanidae. The name Anolidae (Cope, 1864) has sometimes been used, but it is a junior synonym of Dactyloidae (Fitzinger, 1843).
More than 425 species of true anoles are known. New species are regularly described (12 new species in 2016 alone), and most of the recent discoveries have been from the mainland of the Americas, with fewer new anoles described from the comparably better-known Caribbean Islands. Traditionally, all the true anoles were included in the genus Anolis and some continue to use this treatment, in which case it is the largest genus of reptile. Genetic studies have revealed eight clades and it has been proposed that these should be recognized as separate genera. Some of these can be further subdivided.
|The eight proposed genera in Dactyloidae|
Relationship with humans
Some anole species are commonly kept in captivity as pets and especially the Carolina (or green) anole is often described as a good "beginner's reptile", but it too requires specialized care.
Anoles can function as a biological pest control by eating pest insects that harm plants. Anole abundances can be considerably higher in diversified agroecosystems (multiple different plant types) than high-intensity agroecosystems (typically only one or very few plant types, and regular use of agrochemicals), making the former particularly suitable for this type of pest control. However, because of their potential of becoming invasive species, releasing anole species outside their native range is strongly discouraged and often illegal (even if said species occurs elsewhere in a country; for example, it is illegal to release Carolina anoles to California, as its native range is in Southeastern United states).
The willingness of many anoles of living close to humans in heavily altered habitats have made them common. However, there are also many species restricted to specific habitats (for example, primary rainforest) or small islands, making them more vulnerable. Typical threats to these are habitat loss from both humans and extreme weather, or competition/predation by introduced species. Only a small percentage of the many anole species have been rated by the IUCN, but this includes several recognized as threatened. For example, A. amplisquamosus, an endangered species only known from highland forest in the Cusuco National Park region of Honduras, was common in the early 2000s, but by 2006 it had experienced a drastic decline and was only infrequently encountered. A clear explanation for this is lacking, although it may be related to habitat loss. The Finca Ceres anole, a critically endangered species only known from a single location in Matanzas Province, Cuba, has suffered habitat loss both due to hurricanes and expanding agricultural land. The strikingly colored blue anole from Colombia's Gorgona Island is threatened from deforestation and predation by the introduced western basilisks.
Anoles overall do not appear to have experienced the widespread extinctions and extirpations prevalent among larger Caribbean reptiles, although one species, the Culebra Island giant anole, possibly is extinct, and others, at least the Morne Constant anole, do not grow as large today as they once did.
As introduced species
When introduced to regions outside their native range by humans (deliberately or by mistake), anoles may become invasive and represent a threat to small local animals. In the contiguous United States, the Carolina anole has been introduced to California, the brown anole has been introduced to the Gulf Coast states and California, and the knight, bark, Jamaican giant, large-headed, Puerto Rican crested, Cuban green and Hispaniolan green anoles have been introduced to Florida. The Barbados and Morne Constant anoles have also been recorded in Florida, but do not appear to have become established. There are indications that the invasive brown anole is displacing the native Carolina anole in Florida and Texas by outcompeting it and eating its young, although some studies suggest that the latter remains common but has been forced to occur higher in trees (making it less visible to humans). Regardless, the Carolina anole is common and widespread overall, and it has itself been introduced to several regions outside its native range, including California, Hawaii, Guam, Palau, some islands in the West Indies, Tamaulipas in Mexico, isolated locations in Spain and Japan's Ogasawara Islands. In Japan's Ogasawara Islands, the introduced Carolina anoles have caused declines in native lizards and insects, including the near-extinction of the endemic Celastrina ogasawaraensis butterfly and five dragonfly species. In addition to Florida, the Cuban green anole has been introduced to the Dominican Republic, São Paulo (Brazil) and Tenerife (Spain). In Florida and the Dominican Republic it competes with native anoles (Carolina anole and Hispaniolan green anole, respectively) and it is feared that something similar may happen in São Paulo. The same pattern can be seen in Dominica where the introduced Puerto Rican crested anole locally has displaced the endemic Dominican anole. The brown anole and Graham's anole have both been introduced to Bermuda where they threaten the rare Bermuda rock skink. Outside the Americas, the brown anole has been introduced to Hawaii, Tenerife, Singapore and Taiwan, and it is known to be able to change ant communities on the last island.
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