"Mystery is the most beautiful thing that we can experience.

It is the fountain of all true art and science."

    Albert Einstein (1879-1955)

    German-born American physicist, Nobel Prize for Physics in 1921

 

Saint Christopher's Bank tree anole, Anolis bimaculatus, male. Oualie Beach, north-western Nevis, Lesser Antilles.



 

Looking Down on Us

 

Family Dactyloidae: Anoles


    Among lizards, there is a family that is particularly species-rich in the West Indies, and that is Dactyloidae: the anoles. Until recently, dactyloids were considered to be members of another family (Polychrotidae) allied to the iguanids, and the three groups are indeed related. However, there are enough peculiarities in common among the members of this assemblage of small to medium-sized lizards to warrant their own separate status.


    Anoles are perhaps the most commonly seen reptiles in the Caribbean. So abundant and varied are they, that in this website they need an entire section only for them. Ubiquitous from sea level to the highest mountains, in grasslands, forests, deserts, swamps, and near water courses, they often are the first native animals - together with birds - that tourists notice as soon as they leave the airports or disembark from the cruise ships docked at the havens of the West Indies.

 

    The genus is almost certainly polyphyletic and so, from a phylogenetic standpoint, it is a somewhat artificial entity. (It does not include the most recent common ancestor of these organisms and, thus, it is an aggregation of "cousins" instead of true "sister" species). The genus Anolis has been subdivided at times into several more genera: Anolis properly, as well as Norops, Chamaelinorops, Chamaeleolis, Semiurus, Dactyloa, and others. Some of these genera are probably valid from a phyletic and genetic standpoint. However, because their relationships are not as yet clear, all Caribbean anoles are presently considered to belong in the single genus Anolis which, as such, is one of the largest genera among vertebrates. Its adaptive radiation and species-richness is unrivaled among Antillean terrestrial vertebrates except perhaps by another huge - and largely sympatric - genus: the Eleutherodactylus rainfrogs. The variety of species and morphologies makes anoles prime subjects for biological studies of many sorts and, indeed, they have received an immense amount of attention by taxonomists, geneticists, and ecologists.

 

    With relatively minor variations of a basic bodily plan, anoles have been able to fill and exploit arboreal and terrestrial niches and sub-niches in the West Indies and in the American continents to a degree unparalleled among the reptiles of the Western Hemisphere. Other reptilian groups partially fill "anoline niches", as happens with the Phelsuma geckoes and a few agamid genera, all in the Old World. Even if such is the case, no other group of lizards surpasses anoles in their adaptability, success, and number of species.

 

    Anoles exhibit a wide variety of colors. They can be of almost any shade and combination of white, gray, black, brown, yellow, green, blue and purple. Some may be somber, while others are arrayed in almost fluorescent hues. While some are basically monochromatic, others may show astonishingly complex patterns of dots, bands, stripes, reticulations, saddle- or hourglass-shaped spots. Additionally, most have the capacity to change their colors dramatically. Green species can turn brown, gray, or black, and although most brown or gray species cannot turn green, they can still alter the shade of their color to lighter or darker, with or without spots and other marks. These responses are related to the mood of the animal, like fear or anger. Species that normally are light in color may turn much darker when they are cold, and that may help them warm up by absorbing more sunlight. In spite of a common misconception, these color changes are not an attempt to blend with their background, although the basic color pattern and posture of many species can indeed be cryptic. (The ability to change their color has earned anoles the name of "American chameleons". The fact is that they are not closely related to the true chameleons of the Eastern Hemisphere).

 

Puerto Rican emerald anoles, Anolis evermanni, malea. The first one is half way through its metachrosis from green to brown. El Yunque National Forest, north-eastern Puerto Rico.
The second one has undergone complete metachrosis. Carite State Forest, east-central Puerto Rico.



Cuban green anoles, Anolis porcatus.

First photograph: male. From a population introduced into Santo Domingo, southern Hispaniola.
Second photograph, female. Hicacos Peninsula, north-western Cuba.  (Courtesy of Mr. Joseph Burgess).


The bold patterns of many anoles turn out to be disruptive of the animal's silhouette when viewed against its natural background.

Thus, they are rendered less obvious to both predators and prey.

This is a Ponce small-fanned anole, Anolis poncensis, female.

Camp Santiago, Salinas, south-eastern Puerto Rico.



These two photographs of a male Anolis gundlachi were taken seconds apart.
They show how the animal changes colors to better mimic its substrate after being alarmed by my camera.
  El Yunque National Forest, north-eastern Puerto Rico.


Many anolines assume a cryptic position when they feel threatened, like flattening themselves against their perches.
Hispaniolan grass anole, Anolis semilineatus. National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.
(Photograph courtesy of Mr. Mel Jose Rivera).


    There are two morphological characters typical of anoles. One is the presence of a lengthwise fold of skin under the throat, commonly called a "dewlap". The other consists of rows of specialized, transversal scales under their toes which allow them to climb with ease.

 

    The dewlap is especially well-developed in males, although females of most species have a reduced version of it. (Both sexes of a few species lack a dewlap altogether, while females of many giant species have dewlaps that can be as large as those of males). The dewlap can be extended at will, like a flag, when the animal forces out and down a cartilage on its edge. The cartilage itself is attached to the hyoid apparatus under the tongue. This fold of skin frequently possesses bright colors and serves as an intraspecific means of recognition during courtship, or as a means to dissuade a predator or potential rival, since it makes the animal look much larger than it really is. By means of the dewlap, many species will send their visual epigrams even to other species, including humans.


    While it extends its dewlap, an anole may increase the overall effect of the bluff or courtship by adopting stereotyped postures and effecting certain motions, like push-ups, standing high on all four legs, flattening its body laterally, raising dorsal crests, waving its tail, and extruding its tongue.

 


A male Saint Croix's anole, Anolis acutus, shows one of the main traits of its group: a protractile dewlap.
Barrenspot, central Saint Croix, United States Virgin Islands.

 

A male opal-bellied anole, Anolis opalinus, threatens me with its dewlap. Blue Mountains, east-central Jamaica.

 


A male Grenadian bush anole, Anolis aeneus, extends its dewlap. Tempe, central Grenada, Lesser Antilles.

 

A male Dominican anole, (highland race, Anolis oculatus montanus), threatens the photographer with its dewlap.

Morne Trois Pitons National Park, south-central Dominica, Lesser Antilles.

(Photograph courtesy of Mr. Mel Jose Rivera).

 

A male Puerto Rican crested anole, (mainland Puerto Rican race, Anolis cristatellus cristatellus), extends its dewlap.

Camp Santiago, Salinas, south-eastern Puerto Rico.

 

A male Barahona giant anole (western race, Anolis barahonae mulitus), tries to discourage being handled by extending its dewlap.

Bahoruco National Park, south-western Dominican Republic, Hispaniola.



A male Puerto Rican twig anole, Anolis occultus, extends its dewlap. El Yunque National Forest, north-eastern Puerto Rico.
(Photographs courtesy of Mr. Alfredo Colon Archilla).



A male Anolis cooki sticks its tongue at me in a threatening fashion. Guanica State Forest, south-western Puerto Rico.


    All anoles are diurnal. Since they have excellent eyesight, and can see well into the ultraviolet range of the electromagnetic spectrum, their world looks to them very different from ours. What an anole sees in the dewlap of a mate or rival might not be the same as what we humans see. In fact, the most ultraviolet-reflective dewlaps are possessed by species that inhabit open areas, where direct sunlight and blue skies allow for large quantities of the appropriate electromagnetic wavelengths. Species that live in dense forest possess little or no ultraviolet-reflective tissue in their dewlaps although, as happens with most species, they still have such reflective tissue in the corners of their mouths. Since opening their jaws in a threatening display is common in most species, this visual signal can still be seen by individuals engaged in combat. At any rate, all anoles seem capable of detecting ultraviolet light reflected by potential prey or predators.

 

With its sharp vision, a female Puerto Rican twig anole, Anolis occultus, surveys its universe.

Toro Negro State Forest, central Puerto Rico.

 

A male Guanica pallid anole, Anolis cooki prepares to pounce on its prey. Guanica State Forest, south-western Puerto Rico.



This female Puerto Rican giant anole, Anolis cuvieri, shows its binocular capacity as it rotates its eyes forward.
Sabana Seca, north-eastern Puerto Rico.



Deliberate and chameleon-like, a female Anolis cuvieri patrols its territory in search of prey.
Guajataca State Forest, north-western Puerto Rico.


    The other trait, already mentioned above, that is common to all anoles can be noticed when one looks closely at their toes. These have lateral expansions near their tips. Under such are found series of modified scales (collectively called "lamellae") which, like those of geckoes, allow them to take a better hold on their substrate, making most of them agile climbers. Indeed, most anoles are to some degree arboreal in their habits. A few continental species of anole have secondarily lost their lamellae.

 

Southern Hispaniolan green anole (nominate race, Anolis coelestinus coelestinus) male. Polo, south-western Dominican Republic, Hispaniola.

This photograph shows the two traits common to almost all anoles: a dewlap and toe lamellae.

 

A gravid female Puerto Rican crested anole (nominate race, Anolis cristatellus cristatellus), sunbathes in a cool morning.

Guayanilla, south-western Puerto Rico.

 

    Like lizards of many other families, anoles can autotomize their tails, loosing them as a defensive maneuver against enemies. The wriggling appendage is left behind for the predator to entertain itself with, while its former owner flees to safety. In a matter of weeks, the anole will develop another tail which will, however, have a simpler scalation and color pattern, and no true vertebrae, but only cartilage in their place.

 

    Although usually silent, many anoles chirp, squeal, or growl when they are fighting or are attacked by a predator. The noise produced can be clearly heard in the case of some species, when one captures and holds them in hand. These, for example, are the high-pitched growls of a Jamaican giant anole when it is held in hand.



Many anoles eat their own old skin when they shed it. The tissue is too rich in proteins to be discarded.
Puerto Rican crested anole (nominate race, Anolis cristatellus cristatellus), male. Guajataca, north-western Puerto Rico.


Emerald anole, Anolis evermani, lick the nectar of the melastome Micranium latifolium. El Yunque National Forest, north-eastern Puerto Rico.
Some anoles are thus pollinators of a number of plants.
(First photograph: male. Last two photographs, female. Courtesy of Mr. Alfredo Colón Archilla).


    Interestingly, the relationship between bodily ornaments and voice use is inversely proportional. Species that are most reluctant or not known to utter vocal sounds tend to be those with the most elaborate visual ornaments, like complex color patterns, high dorsal and caudal crests, and especially large or gaudy dewlaps. Those with proportionally small dewlaps, no crests, and rather simple color patterns tend to be the most vocal. As an example of this phenomenon, Hispaniolan species save for the three crown giants in that island lack caudal crests and, among them, many squeal persistently when they are held in hand. By comparison, most Puerto Rican species possess tail crests, sometimes comparatively enormous, and only Anolis occultus, (small, drab, and without crests) utters an alarm call when captured. The more ornate species only chirp angrily when individuals fight between themselves - in other words, they rely on voice only when visual deterrence between rivals fails to settle the dispute.

 

A female Puerto Rican twig anole, Anolis occultus, shows the use of its prehensile tail as it climbs up its perch and then turns back down.

The reptile slightly changes color in fear, as it notices my presence. Unlike the more commonly seen species, twig anoles are slow and chameleon-like in their movements.

El Tayonal Private Nature Reserve, Arecibo, northern Puerto Rico.

 

    In the continent, anoles are found from the southern United States and throughout Central America to southern South America. However, it is in the Antilles where this group reaches its pinnacle of adaptive evolutionary radiation.

 

    All species are mainly predators of insects and arachnids although some, particularly the giant species, are also adapted to feed on other prey like snails, which shells they break with their powerful jaws. The giants are also capable of feeding on small vertebrates, like other lizards, mice, and even birds. Additionally, a good number of species will take fruit or nectar from the plants around them. The dentition of anoles, like those of other iguanians, is heteromorph. The front teeth are conical and sharp while those in the back of the jaws have multiple cusps and partially function like mammalian molars, to crush hard objects.

 

The anoline dentition reflects the capacity to take on different food items.

Compare the front and back teeth of this male yellow-chinned anole, Anolis gundlachi.

El Yunque National Forest, north-eastern Puerto Rico.

 

Saint Vincent's tree anole, Anolis griseus, male, showing its teeth. Layou Valley, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.

 

The teeth of a Barahona giant anole (western race, Anolis barahonae mulitus). Bahoruco National Park, south-western Dominican Republic, Hispaniola.



The huge jaw muscles bulging below the ear of this male Barbudan Bank tree anole,  Anolis leachii, are not just a bluff.
Some anolines have powerful bites.
From a population introduced into Charlotte Amalie, Saint Thomas, Unites States Virgin Islands.


    West Indian anolines, especially those of the Greater Antilles, have segregated and differentiated from their congeners, radiating and co-evolving in the same island by specializing in the use of different niches within their range. The result is that frequently unrelated species, each usually living on a different island bank, look astonishingly similar in size, shape, and even color.


    Some, the grass anoles, have slender bodies and extremely long tails, are grayish or yellow-brown in color, and camouflage themselves among the grasses on which they live. Others, the tree-crown giants - largest of all anolines - inhabit the upper branches of trees, are usually some shade of green, have large toe pads, huge heads and powerful jaws. Trunk-ground anoles are brown or gray, have short and stocky bodies, tall heads, and long limbs, and live on the ground, lower tree trunks, and rocks. Still others, like the twig anoles, are also usually some shade of gray or brown, but instead have elongated and flattened bodies and heads, and very short tails and limbs, suited for a life spent crawling on small branches, twigs, and on leaves. And so on, and so forth.

 

    The results of this convergent evolution among different islands banks are called "ecomorphs", unrelated but very similar groups of habitat specialists that are found across islands, due to their adaptations to the same structural niche.

 

    Within the same island, their mutual segregation - through divergent evolution among sympatric (and often closely related) species - into the diverse structural niches of their three-dimensional environment allows for a multitude of species to share the same general habitat with a minimum of competition.

 



    The concept of "ecomorph" might be better understood with some visual aid.

 

    The next four photographs show related Puerto Rican anoles that underwent a process of divergent evolution within the same island that allows them to live in close proximity (in "sympatry") by using different niches. For this reason, although they are closely related, they look quite different. In a morphological and ecological sense, they "moved away" from one another, and thus ended up belonging to different ecomorphs.

 

Puerto Rican twig anole, Anolis occultus, female. Toro Negro State Forest, central Puerto Rico.

This species belongs to the twig dwarf ecomorph. The perch on very thin branches and their appearance and behavior help them to pass unnoticed by both foes and prey.

Such species have short legs and tails, apt for crawling slowly but safely on their usual substrate.

 


Yellow-chinned anole, Anolis gundlachi, male. El Yunque National Forest, north-eastern Puerto Rico.

This species belongs to the trunk-ground ecomorph. These anoles can be distinguished by their habit of perching close to the ground,
their large and deep heads, short and stout bodies, long limbs used for sprinting and jumping between
their perches and the ground, and brown or gray colors that blend well with bark and leaf litter.

 

Olive bush anole, Anolis krugi, male. Florida, central Puerto Rico.

This is a species belonging to the grass-bush ecomorph. Such anoles are slender, have very long tails and legs for balancing on thin
grass blades and branches of shrubbery, and frequently have lengthwise, cryptic color patterns of browns, yellows and whites.

 

Puerto Rican emerald anoles, Anolis evermanni.
First photograph: male. Aguas Buenas, east-central Puerto Rico.
Second photograph: Florida, central Puerto Rico.

This is one of the trunk-crown anoles, noted for their habit of frequently perching high on the trunks and crowns of trees, their long and flattened bodies,
pointy and flattened heads and snouts, rather short legs for slow crawling, large toe-pads to cling securely to the substrate and avoid falls, and their usually green color.

 

     Now, let us take the last species shown above, namely Anolis evermanni, and compare it with three others. Together, these four species evolved and live in four different islands, and normally are never found together (they live in "allopatry"), yet underwent a process of convergent evolution. Morphologically and ecologically, they "moved toward" one another. In such way, although they are unrelated and live in different islands, they look very much alike because they evolved to occupy the same ecological niche. All four belong to the trunk-crown ecomorph. Notice how the traits emphasized above for Anolis evermanni apply to all four. In fact, they are so similar to one another that a person not familiar with them could even think that they belong to the same species.

 

Puerto Rican emerald anole,  Anolis evermanni, male. Carite State Forest, east-central Puerto Rico.

 


Hispaniolan northern green anole (nominate race, Anolis chlorocyanus chlorocyanus) male.

Los Haitises National Park, north-eastern Dominican Republic, Hispaniola.

 

Jamaican turquoise anole, Anolis grahami, male. Windsor, north-central Jamaica.

 


Cuban green anole, Anolis porcatus, male.

From a population introduced in Santo Domingo, southern Dominican Republic, Hispaniola.


    Oftentimes, anoles belonging to the same ecomorph are found within the same island. Now, if the very concept of "ecomorph" seems to presuppose convergent evolution of unrelated species living in different islands, how can this be? The answer lies in the fact that once an ecomorph has evolved, a species within it can give rise to two or more sibling species still belonging in the same ecomorph, adapted to the same basic structural niche, yet each further adapted to different degrees of humidity, temperature, sunlight, and other abiotic factors, or they differ in size from each other.


    The next three species of anole of the grass-bush ecomorph diverged from a common ancestor but still occupy the same structural niche: shrubbery and grasses, sometimes in the same general geographic area (in which case they are "sympatric"). However, Anolis krugi occupies the shadiest and most humid areas with the densest vegetation. Anolis pulchellus occupies relatively shady but more open and less humid areas. Anolis poncensis is found in the sunniest, most exposed and driest areas. In that way, although they can live in the same general area, their use of different sub-niches avoids competition between the three. Thus, while they may be sympatric, they are seldom "syntopic" (they do not live side by side).

 


Olive bush anole, Anolis krugi, male. Carite State Forest, east-central Puerto Rico.

 

Puerto Rican bush anole, Anolis pulchellus, male. Bosque del Milenio, San Juan, north-eastern Puerto Rico.


Ponce small-fanned anole, Anolis poncensis, male. Guayanilla, south-western Puerto Rico.


    Anoles that live in sympatry and belong to different, and even the same, ecomorph further avoid competition by having different temperature needs. Within the same general area, heliothermic species inhabit open, sunny microhabitats. They can frequently be seen basking on exposed perches, and are most active in warm, clear weather. On the other hand, non-heliothermic species prefer shady areas, and frequently are thermoconformers, as well. That is, they have adapted to depend much less on direct sunlight for warming up through basking, and indeed can remain active in cool, even chilly, weather conditions. Several species that live anywhere below the canopy of montane rain forests and cloud forests belong to this type of anoline.

 

The following three species of anole are sympatric in some areas and diverged from a common ancestor. The first two belong to the same ecomorph (bush) but, in their case, they diverged to occupy niches that are the same in terms of structure but different in temperature: Anolis pulchellus is a species of rather open, warm areas, while Anolis krugi prefers the cool shade of the forests. The third species, Anolis gundlachi, is also related to them, but diverged to occupy a different structural niche (and thus, belongs in a different ecomorph: trunk-ground). Thus, although it has similar thermal needs as Anolis krugi it still avoids competition with it by making use of a different part of its tri-dimensional habitat. Its chances of competition are even less with A. pulchellus, since it is not only spatially separated from it in the same way it is from A. krugi, but also by very different sunlight, humidity, and temperature needs.

 


Puerto Rican bush anole, Anolis pulchellus, male, balancing itself on an inflorescence of Piscidia carthagenensis.

Virgin Islands National Park, Saint John, United States Virgin Islands.

This is a heliothermic anoline of sunny, warm areas.

Grass-bush ecomorph.

 

Sister species to Anolis pulchellus, this female olive bush anole, Anolis krugi, occupies the same structural habitat, but prefers shadier and cooler places.

El Yunque National Forest, north-eastern Puerto Rico.

Grass-bush ecomorph.

 


Related to the two previous species, and sympatric with them through some of its range,

this is a male yellow-chinned anole, Anolis gundlachi. El Yunque National Forest, north-eastern Puerto Rico.

This anole is a non-heliothermic lizard that inhabits exclusively the ground and lower vegetation levels of dark, closed-canopy mesic forests.

This most thermoconformist of Puerto Rican anolines is never seen basking in the sunlight, and prefers a dark and cool microhabitat.

It avoids competition with its two congeners by occupying the shadiest, coolest areas of the forest (vis a vis A. pulchellus)

and a different structural niche (vis a vis both A. pulchellus and A. krugi).

Trunk-ground ecomorph.




    According to the combination of their size, morphology, and preferred niche within their tri-dimensional habitat the Antillean anoline ecomorphs are:


- grass-bush: small-sized species, with long legs and snouts and very long tail used for balance, occupying grasses and low woody vegetation
- trunk-ground: medium-sized, long-legged (jumper) species with shot snouts, found on the lower parts of tree trunks, ground, and rocks
- trunk: medium-sized, with flatted bodies and rather short legs and snouts, which squirrel on the middle section of trunks
- trunk-crown: medium sized, with short legs and longs snouts, occupying all sorts of perches in higher tree trunks and canopy
- crown giant: large to very large species with long and powerful jaws that usually perch high in the canopy
- twig: minute to very large species with very short legs and tails and long snouts that occupy perches of small diameter at all heights; they are unique in their locomotion in that they move in a slow, chameleon-like way


    In Addition to the normal six ecomorphs, some anolines in the Greater Antilles conform to none of such types. Some anoles are aquatic or specialize in perching on rocks or cliffs. Others, expecially among species that exist without congeners in their islands, are habitat generalists, though all are still arboreal to some degree or another.

Aside from segregating according to species, many anoles do the same according to sex and or size class.

Male olive bush anoles, Anolis krugi, usually choose perches that are larger and higher above ground,

like shrubs and saplings, than females of the same species, which are confined to grasses, sedges and ferns.

Male and female: El Yunque National Forest, north-eastern Puerto Rico.

 

    All being said, it should be borne in mind that although many West Indian anoles (and a number of continental species) can be generally grouped within one or another ecomorph, no two species within the same ecomorph are exactly alike. For example, the Cuban Anolis luteogularis, the Jamaican A. garmani, the Hispaniolan A. barahonae, the Puerto Rican A. roosevelti, are tree-crown giants. However, they do not exactly behave like, have the same size as, or fill the same niche as each other's. Minor variations occur among species within the same ecomorph, due to differing - if only slightly - geological, climatic, and ecological conditions among islands. Such diverse conditions always determine the precise evolutionary and speciation courses and patterns shown by the species that are subjected to them.

 

    Size extremes vary among islands depending also on how many species inhabit each particular island. In the Greater Antilles Cuba, with the most species, has the largest of all crown giants (Anolis noblei) and the smallest of the dwarves (A. ophiolepis). Jamaica, with the least number of species (seven) has the smallest of all crown giants (A. garmani) and the largest of all dwarves (A. valencieni). Hispaniola and Puerto Rico, with a number of species intermediate between those of the other two islands, accordingly have giant and dwarf anoles which size could be considered, thus, "medium". This phenomenon is due to the fact that the more species live in sympatry, the more they tend to differ in size in order to avoid competition among themselves.

 


As a group, twig anoles are the smallest members of the genus Anolis. This is an adult female Anolis occultus, placed on a human hand for scale.

Barranquitas, central Puerto Rico.

Twig ecomorph.

 


Big-fanned trunk anole, Anolis christophei, male.

Central Mountain Range, Dominican Republic, Hispaniola.
The "trunk" anoline ecomorph is found only in Cuba and Hispaniola.

This well-camouflaged species can be difficult to see against its habitual background of bark and lichens.

 

Big-fanned trunk anole, Anolis christophei, female. Ebano Verde Nature Reserve, central Dominican Republic, Hispaniola.

 

Cuban trunk anole, Anolis argenteolus, female. Cabo Cruz, Granma Province, south-eastern Cuba.

(Photograph courtesy of Dr. Richard C. West).

 


Hispaniolan gracile anoles, Anolis distichus, subspecies dominicensis, male and female. Central Mountain Range, Dominican Republic, Hispaniola.

Trunk-bush ecomorph.

 


Anolis distichus ignigularis. Santo Domingo, southern Dominican Republic, Hispaniola. Trunk ecomorph.



Anolis distichus ravitergum, male. Azua, south-western Dominican Republic, Hispaniola.

 

Gonave gracile anole, Anolis caudalis, male. Cote de Archadins, north-western Haiti, Hispaniola. Trunk ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Yellow-bellied desert anole, Anolis websteri, male. Near Gonaives, north-western Haiti, Hispaniola. Trunk-ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Desert gracile anole (nominate race Anolis brevirostris brevirostris), male. Enriquillo National Park, south-western Dominican Republic, Hispaniola. Trunk-ecomorph.


Desert gracile anole, Anolis brevirostris, subspecies wetmorei, female.

Santa Cruz de Barahona, south-western Dominican Republic, Hispaniola.

The species substitutes A. distichus in some xeric regions.

Trunk ecomorph.

 

Jacmel gracile anole, Anolis marron, male. Jacmel, south-eastern Haiti, Hispaniola.

Trunk ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Alto Velo gracile, anole, Anolis altavelensis, male. Alto Velo Island, south-western Dominican Republic, Hispaniola.

Trunk ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Banded red-bellied anole, Anolis rupinae, adult male, juvenile male, and female. Massif de La Hotte, south-western Haiti, Hispaniola.

Grass-bush ecomorph.

(Photographs courtesy of Mr. Eladio M. Fernandez).

 

Artibonite bush anole, Anolis rimarum, male and female.

First photograph near Gonaives, north-western Haiti, Hispaniola.

Second photograph near Marmalade, north-western Haiti, Hispaniola.

Grass-bush ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Hispaniolan hopping anole, Anolis barbouri, male. Bahrouco Mountains, south-western Dominican Republic, Hispaniola.

This is a truly terrestrial species, living on the leaf litter of mesic forests.

(Photographs courtesy of Mr. Marcos Rodriguez).

 

Jamaican giant anoles, Anolis garmani, males.

First photograph: Negril, western Jamaica. Second photograph: Port Antonio north-eastern Jamaica.

Trunk-crown ecomorph.

 

    No Caribbean island possesses all the anoline ecomorphs. The Greater Antillean island banks have by far the most complex anoline faunae in the region, with the Cuban and Hispaniolan insular banks having the most complete arrays of ecomorphs. They also have the highest number of species (more than 50 are found in each). They are followed by the Puerto Rican bank, with 13 species. The Jamaican bank has the simplest Greater Antillean anoline fauna, with six native species (or seven, if the Cuban brown anole, Anolis sagrei, turns out to be a natural invader, something that is still not clear). Several species closely related to those of the Greater Antilles inhabit The Bahamas and other, nearby insular banks (Mona Island, Navassa, the Cayman Islands, etc.).

 

Some Greater Antillean islands have peculiar anolines which equivalents are not found in others.

For example, semi-aquatic anoles are only found in Cuba and Hispaniola.

This is a black stream anole, Anolis eugenegrahami, male. Near Gonaives, north-western Haiti, Hispaniola.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 


Cuban stream anoles, Anolis vermiculatus, male, female and juvenile. Sierra del Rosario, western Cuba.

(Photographs courtesy of Mr. Joseph Burgess).


Cuban twig anoles, (Cuban, Anolis angusticeps angusticeps).
First photograph: male. Cuban National Gardens, Havana, north-western Cuba.

Second photograph: male. Matanzas, north-western Cuba.
Third photograph: female. Sierra del Organos, western Cuba.

Twig ecomorph.

(Photographs courtesy of Mr. Joseph Burgess).

 

Cuban twig anole, (Bahamian race, Anolis angusticeps oligaspis), male. Southern Andros Island, The Bahamas.

Twig ecomorph.

(Photographs courtesy of Mr. Joseph Burgess).

 

    Among the Lesser Antillean insular banks, none has more than two anoline species. The only exception is found in the Guadaloupe bank which has several described species. However, they live in allopatry, with Anolis marmoratus in the main double island, while the others inhabit outlying smaller land masses.

 

    Anoles that live singly on their respective islands tend to be ecomorphic generalists and medium in size. This is due to the fact that in the absence of competition, and perhaps having had no time to evolve further into a plurality of species, they occupy all the available niches without a need for further variations in morphology and size. When two species are found together, they will always differ in size. This is taken to an extreme in the Lesser Antillean giant species Anolis bimaculatus, A. leachii, A. griseus, and A. richardi, some of which partially fill the crown giant ecomorph. (Though they are never as massive as, and their morphologies differ somewhat from those of the Greater Antillean giants). Their much smaller relatives in the same island, namely Anolis schwartzi, A. wattsi, A. trinitatis, and A. aeneus, resemble the "trunk ground" and "bush" anoles of the Greater Antilles.

 

    As with the Greater Antillean and Bahamian ecomorphs, these differences in size and niche occupation are explained as an evolutionary mechanism for the avoidance of competition. But one species that occupies its island all by itself breaks the rule of being medium size. This is the Marie Galant anole, Anolis ferreus, which is a giant.

 

    It should be noted that the concept of "ecomorph" could also been applied to other taxa in the Caribbean, like the rain frogs of the genus Eleutherodactylus, the geckoes of the genus Sphaerodactylus, and the boas of the genus Epicrates. Especially the first two genera have multiple species in the Greater Antilles, each of them segregated from those others sympatric with it by some difference in morphology and ecological niche. The diverse species are placed in an ecomorph class depending on whether they live in mesic or xeric forest, in caves or in bromeliads, on the ground or on trees, etc.

 

Five-striped grass anole, Anolis ophiolepis, captive specimen, male.

Grass-bush ecomorph.

(Photograph courtesy of Mr. Joseph Burgess).

 

Bahoruco long-snouted anoles (nominate race, Anolis bahorucoensis bahorucoensis).  Polo, south-western Dominican Republic, Hispaniola.

First three photographs: male.

Last photograph: female.

Its beautiful colors are actually designed as a cryptic pattern to disguise it among the vegetation.

Grass-bush ecomorph.

 

La Hotte long-snouted anole, Anolis dolichocephalus, female. Massif de La Hotte, south-western Haiti, Hispaniola.

Grass-bush ecomorph.

This species is related to A. bahorucoensis (above).

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

La Selle long-snouted anole, Anolis hendersoni, male. Near Petionville, southern Haiti, Hispaniola.

Grass-bush ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

La Hotte bush anole, Anolis monticola, males. Massif de La Hotte, south-western Haiti, Hispaniola.

Grass-bush ecomorph.

(Photographs courtesy of Mr. Eladio M. Fernandez).

 


Several Cuban anolines formerly placed in the genus Chamaeleolis have very specialized morphologies and habits.
The western bearded anole, Anolis barbatus, is one of the "twig-giants." Male and female.
Sierra de los Organos, Pinar del Rio, western Cuba.
(Photographs courtesy of Mr. Joseph Burgess).

 

Oriente bearded anole, Anolis porcus, female. Cueva de los Peces, south-western Cuba.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Cuban white-fanned anole, (nominate race, Anolis homolechis homolechis).

First photograph: Guantanamo Naval Base, Guantanamo, south-eastern Cuba. (Courtesy of Mr. Joseph Burgess).

Second photograph: La Gran Piedra, south-western Cuba. (Courtesy of Mr. Eladio M. Fernandez).
Third photograph: female. Viñales, western Cuba. (Courtesy of Mr. Joseph Burgess).

 Trunk-ground ecomorph.

Cuban blue anoles, Anolis allisoni.
First two photographs: male. Playa Larga, south-western Cuba. (Courtesy of Mr. Eladio M. Fernandez).
Last photograph: female. Hicacos Peninsula, north-western Cuba. (Photograph courtesy of Mr. Joseph Burgess).

 Trunk-crown ecomorph.

Cuban tiger anole, Anolis rubribarbus, male. Monte Iberia, Baracoa, eastern Cuba.

Trunk-ground ecomorph.

(Photograph courtesy of Dr. David Ortiz Martinez).

 

Cuban tiger anole, Anolis rubribarbus, male. El Recreo de Nibujon Natural Trail, Guantanamo, south-eastern Cuba.

Trunk-ground ecomorph.

(Photograph courtesy of Dr. Richar E. Glor).

 


Red-fanned rock anoles, Anolis mestrei.

First three photographs: male and two females. Viñales, western Cuba. (Courtesy of Mr. Joseph Burgess).
Third photograph: male. La Gran Piedra, south-western Cuba.
(Photograph courtesy of Mr. Eladio M. Fernandez).

Trunk-ground ecomorph.



Cuban eyespot anole, Anolis quadriocellifer, male. Cabo San Antonio, western Cuba.
(Photograph courtesy of Mr. Joseph Burgess).

 

    Although anoles have generally evolved on their respective islands, natural invasions do occur. The Cuban green anole, Anolis porcatus, or its ancestor, invaded The Bahamas, and there speciated into others, like A. smaragdinus and A. fairchildi. It also invaded further north, were it evolved into the North American green anole, A. carolinensis. So, in this case, while the different species evolved in each island, the ecomorph did not, but rather evolved in only one island and then spread to others.


North American green anoles, Anolis carolinensis.

First photograph, male. Brevard County, Florida, south-eastern United States. (Courtesy of Mr. Joseph Burgess).
Last two photographs: females. Fort Polk, Louisiana, southern United States.
This species of temperate areas is actually Neotropical in origin, one of the very few reptiles which ancestor invaded North America from the Antilles.

It has a common parent population with the modern Anolis porcatus of Cuba.

Trunk-crown ecomorph

 

    An alternative scenario, at least as far as the Bahamian populations are concerned, assumes that A. porcatus was already present in the Bahamian region when it was connected, at one time, with Cuba. Once the islands fragmented, the populations present there from the start diverged from one another. The end-product of this process is called "vicariance", a process of speciation which occurs as a result of the separation and subsequent isolation of portions of an original population.

 

    Either one of the above cases occurred also, for example, with the Hispaniolan and Puerto Rican crown giant anoles. The Puerto Rican bank giants, Anolis cuvieri and A. roosevelti, are closely related to the three Hispaniolan bank species, A. baleatus, A. barahonae and A. ricordii. As with the A. porcatus group of species, the way in which the Puerto Rican crown giant species got there - whether by invasion and further speciation, or through vicariance and further speciation - is not known for certain.

 

    If you wish to learn more about anoline ecomorphs, as well as about many Caribbean amphibians and reptiles, please visit Dr. S. Blair Hedges excellent web site: Caribherp.

 


Dominican giant anole (subspecies Anolis baleatus scelestus) male. National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.
This is a crown giant anoline related the Puerto Rican giant (further below).
I photographed this individual about 15 minutes apart to show its metachrosis from brown to green.

Among crown giants, species with long legs, like this one, are able to jump easily among tree branches.

In the second photograph, notice how anoles have a lateral groove on each side of the snout. Their eyes can

move independently (in this individual, one eye is pointing to the front, while the other is looking upwards).

However, when the animal gets ready to pounce on its prey, both eyes look to the front along the grooves, granting

the animal a degree of binocular vision which allows it to measure distance accurately.



Anolis baleatus litorisilva, female. Parque Nacional del Este, eastern Diominican Republic, Hispaniola.
Photographs taken two minutes apart.

 


Anolis baleatus scelestus, female, in cryptic position. National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.

It presses itself against the stem, and flattens its body laterally, in order to appear less conspicuous.

 


Anolis baleatus scelestus, female. National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.

 


Anolis baleatus scelestus, juvenile. National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.

(Photograph courtesy or Mr. Pedro Genaro Rodriguez).

 

Bahoruco giant anole, (western race, Anolis barahonae mulitus), male. Bahoruco National Park, south-western Dominican Republic, Hispaniola.

Crown giant ecomorph.

 


Caymans blue-fanned anoles, (eastern Grand Cayman race, Anolis conspersus lewisi), two males and two females. Queen Elizabeth II Botanic Park, central Grand Cayman.

As is typical with species that are the only ones on their islands, this is an ecomorphic generalist.

Trunk-crown ecomorph.
(Photographs courtesy of Mr. Marcos J. Rodriguez).

 

Opal-bellied anole, Anolis opalinus, male and female.

First photograph: Blue Mountains, east-central Jamaica.

Second photograph: Port Antonio, north-eastern Jamaica.

Trunk-crown ecomorph.

 

Green-banded anole, Anolis fowleri, male. Le Vega, central Dominican Republic, Hispaniola.

(Photograph courtesy of Dr. Richard E. Glor).

 

    Some anoles that belong on certain ecomorphs are so partial to their niches that they are practically never found outside of such. At the other extreme, some species, especially those that are the only ones found in their island, are generalists that don't fall within any given ecomorph, and are found almost in any place where there is some vegetation.

 


Northern orange armpit anole, Anolis aliniger, male. Central Mountain Range, Dominican Republic, Hispaniola.
Twig anoles are thus called for their habit of perching on small branches and twigs of the underbrush and canopies of forested areas.

Trunk-crown ecomorph.

 

    Anoles are the main diurnal insectivores in the Antilles. However, many will eat vegetable matter on occasion, mostly fruit. Additionally, some will be partial carnivores, especially the giant species: they will eat smaller lizards, as well as small frogs, mammals, and birds.

 


Anoles are voracious animals, sometimes devouring even other anoles. Here, a female

Puerto Rican crested anole, Anolis cristatellus cristatellus swallows a juvenile Anolis krugi.

El Rosario, south-western Puerto Rico.

Trunk-ground ecomorph.

 

A male Anolis coelestinus coelestinus feasts on an arboreal cricket. Near Polo, Bahoruco Mountains, south-western Dominican Republic, Hispaniola. Trunk-crown ecomorph.



A male Anolis cristatellus cristatellus devours a blindsnake, Typhlops hypomethes. Trujillo Alto, north-eastern Puerto Rico.

Perhaps it did bite more than it could chew.

Trunk-ground ecomorph.
(Photographs courtesy of Mr. Alfredo Colon Archilla).



Anoles will occasionally feed on vegetable matter. This female Saint Lucian anole (Anolis luciae), eats pieces of coconut and yam at a bird-feeder.
Soufriere Botanical garden, south-western Saint Lucia, Lesser Antilles.
Habitat generalist.

 

    Anoles possess a very keen eyesight. Their eyes are capable of moving and focusing independently from one another. Moreover, the sides of the snout between the "canthi rostralis" (the raised lateral edges on top of the snout, which many amphibians and reptiles have) and their upper lips are concave. So when an anole directs both eyes to the front, it achieves a certain degree of binocular vision. With such, and their quick reflexes, they can accurately judge the distance to prey, and quickly capture their next meal. It is a fascinating spectacle to watch any giant anole slowly crawl and remain motionless by turns, as it stalks an individual of a smaller - and frequently more agile - species. When it comes within striking distance, the lighting-fast attack belies its seemingly slow and cumbersome demeanor.

 

    Due to their divergent evolution within a given island, as many as eleven species can occur in sympatry in the same general area. The following six species are found together in the mesic and hydric forests of Puerto Rico.

 

Yellow-chinned anole, Anolis gundlachi, female. Toro Negro State Forest, central Puerto Rico. Trunk-ground ecomorph.


Puerto Rican giant anoles, Anolis cuvieri, males.
First photograph: Guajataca State Forest, north-western Puerto Rico.
Second photograph: Florida, central Puerto Rico.
Third photoraph: Toa Baja, north-eastern Puerto Rico.
This elegant lizard exhibits two color morphs. The more common one, shown above, varies from a yellowish green to a deep emerald,

sometimes with black spots. The males frequently have a blue or purple tinge to their heads, and this color often continues along the spine.

Afraid of the camera, the individual on the left (shedding its skin) opens its eyes wide to appear more menacing.

In fact, all anoles are shy and harmless animals, though giant species can deliver quite a bite if handled carelessly.

Crown giant ecomorph.

Anolis cuvieri, male and female. Florida, central Puerto Rico.

This is the other, rarer morph: greenish gray, usually with diffuse, darker spots and bands.

Although they are rather slow animals when moving about, members of this and other crown giant species are

known to perch near flowers and capture hummingbirds that approach to drink nectar.

Also, with its long legs, this is a crown giant "jumper", like its close relatives, the Hispaniolan giants.

 

Anolis cuvieri, females.
First photograph: Guajataca State Forest, north-western Puerto Rico.
Second photograph: Toa Baja, north-eastern Puerto Rico.


A male Anolis cuvieri sleeps at night on a thin branch.

This habit seems to be a defensive behavior against boas and other nocturnal predators. Any movement on the flimsy perch will wake them up and alert them to danger.

Florida, central Puerto Rico.

 


Neonate Anolis cuvieri are vastly different from adults. They mimic dry leaves and twigs, and choose low perches.

This color behavior seem to be suited to escape predators, including adults of the same species.

Ciales, central Puerto Rico.


Emerald anole, Anolis evermanni. First photograph: male. Carite State Forest, east-central Puerto Rico.
Second photograph: female. El Yunque National Forest, north-eastern Puerto Rico.
Trunk-crown ecomorph.



Puerto Rican spotted anole, Anolis stratulus, male. Guajataca, north-western Puerto Rico.
Named for the black markings on its back, more noticeable on males.
This is a species with an excellent camouflage against its habitual background

of tree-bark, and inhabits the canopies of mesic forests by the thousands per acre.
Trunk-crown ecomorph.

 

Puerto Rican spotted anole, Anolis stratulus, female. Guana Island, British Virgin Islands.
Trunk-crown ecomorph.



Anolis krugi, female. Camuy Caverns Park, Camuy, north-western Puerto Rico.
Grass-bush ecomorph.


Puerto Rican twig anoles, Anolis occultus.

First two photographs: male. Mata de Platano Field Station, Arecibo, northern Puerto Rico.

Third photograph: female (asleep at night). Barranquitas, central Puerto Rico.
Last photograph: female. Toro Negro State Forest, central Puerto Rico.

This smallest of Puerto Rican anolines mimics its substrate in both shape and color, and flattens itself against twigs when approached.

Its short, prehensile tail helps it keep its balance as it slowly maneuvers in search of food and mates.

Its belly is concave, a feature which allows it to "fuse" with its thin perch, effectively disappearing from sight in an instant.

Twig ecomorph.


    Some of the diverse species of dwarf twig anoles, in particular, can be so similar that it takes a whole lot of experience to tell them apart. Just consider the last species shown above together with the next three. All have the same basic morphology adapted to balance themselves carefully on, well... twigs. Personally, they are my favorite kind of anoline after the crown giant species.

 

Cordillera Central twig anole, Anolis insolitus, female.

Ebano Verde Nature Reserve, Central Mountain Range, Dominican Republic, Hispaniola.

Twig ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Bahoruco twig anole, Anolis sheplani, female.

Near Polo, Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

Twig anoles, like this one move slowly and fluidly, more like a chameleon than a typical anole.

Twig ecomorph.

 

Neiba twig anole, Anolis placidus, juvenile. Neiba Mountains, south-central Dominican Republic, Hispaniola.
Twig ecomoph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

    Occupying a similar niche, twig giants (only two species) are considerably larger, though still small by absolute standards. The reason for their larger size seems to be the absence of some competition in their niche. Green twig anoles are indeed that color, and have somewhat longer legs, making them quicker in movements than other twig anoles.

 


Jamaican twig anoles, Anolis valencienni. Windsor, north-central Jamaica.

First five photographs: males. Last photograph: female.

Twig ecomorph.



Female Jamaican rwig anoles seeking to lay their eggs in a communal nest site inside a hollow tree.
Hellshire Hills, southern-eastern Jamaica.
(Photograph courtesy of Mr. Joseph Burgess).


Hispaniolan green twig anoles, Anolis singularis. Polo, Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

First two photographs: male. Last photograph: female.

Twig ecomorph.

 

    Most anole species are strongly territorial. Both sexes, but especially males, will defend their territories from members of the same species, and even from others similar to their own. When a territorial exhibition (dewlap extensions, push-ups, etc.) fails to convince a rival that it is not welcome, a fight may ensue. After some mutual pushing and biting, the defeated antagonist will retreat quickly. However, one or both individuals might loose teeth and receive wounds on the head, neck, and limbs, which might later become infected, causing the death of the animal. Thus, although such fierce battles are seldom, if ever, directly fatal, both combatants avoid as much as possible to prolong them unnecessarily. As I stated above, when physical violence does ensue many species become vocal in their fright or anger.

 

Male yellow-chinned anoles, Anolis gundlachi, fight for territory.

After a series of push-ups and dewlap and tongue extensions failed to settle the dispute, the two individuals came to blows.

Carite State Forest, east-central Puerto Rico.

Trunk-ground ecomorph.


Even female anoles may exhibit strong territorial responses. This female Anolis gundlachi threatens the camera as I approach it.

Carite State Forest, east-central Puerto Rico.
Trunk-ground ecomorph.

 

Some anoles have brightly-colored irides. They can open their eyes wide to add to the aggressive display.

These are yellow-chinned anoles, Anolis gundlachi, males.
First photograph: Guajataca State Forest, north-western Puerto Rico.

Second photograph: El Yunque National Forest, north-eastern Puerto Rico.

Trunk-ground ecomorph.

A male Puerto Rican emerald anole, Anolis evermanni, raises its crests and flattens its body laterally to bluff me away.

Guilarte State Forest, west-central Puerto Rico.
Trunk-crown ecomorph.

 

An angry male olive bush anole, Anolis krugi, sticks its tongue at me and threatens with its dewlap.

El Yunque National Forest, north-eastern Puerto Rico.

Grass-bush ecomorph.

Sometimes anoles hurt each other seriously during a fight.

The first photograph shows a female Anolis krugi that had its snout torn apart by a rival. It is now condemned to a slow death by starvation.

Camuy Caverns Park, Camuy, north-western Puerto Rico.
Grass-bush ecomorph.
In the second photograph a female Anolis cristatellus cristatellus seems to have managed to survive, and it is even gravid.
Carite State Forest, east Central Puerto Rico.
Trunk-ground ecomorph.


    Like most lizards, anoles reproduce by means of eggs. Females hide their eggs (usually laid one at a time) underground or below rocks and debris. In the lowlands, reproduction may take place at any time, although it is normally tied to the onset of the summer and winter rainy seasons. In the cooler highlands, it generally takes place only during summer.

 


Two pairs of Anolis krugi copulate. El Yunque National Forest, north-eastern Puerto Rico.

As with all lizards and snakes, males have two hemipenises, but insert only one at a time.
Grass-bush ecomorph.

 

    Neonate anoles may be colored in a way strikingly different from adults. Also, they choose lower perches than their cannibalistic parents, in order to avoid ending up as their next meal.

 

    Many anoles that live on mountainous areas exhibit the behavioral trait of choosing exposed perches to sleep at night. This habit results from the need to raise their body temperatures as fast as possible in the morning, once the sun comes up. Froim such perches, they can receive the first rays of warm sunlight.

 

    Among Antillean vertebrates, some species of anoles also attain very high densities in some areas. In Puerto Rico, Anolis stratulus and Anolis pulchellus can be found by the tens of thousands per acre in their respective habitats. In the Saint Croix's island bank, Anolis acutus can be found in similar numbers.

 


Puerto Rican bush anole, Anolis pulchellus, females. Florida, central Puerto Rico.

Grass-bush ecomorph.

Saint Croix anoles, Anolis acutus.

First photograph: male, Creque Dam, north-western Saint Croix.
Second photograph: female, Barrenspot, central Saint Croix, United States Virgin Islands.

This is the only anole in its island bank, and is an ecomorphic generalist.

 

    Some anole species that belong to the same ecomorph do live on the same island. However, the usually segregate from one another and avoid competition through variations in their ecological niches. In Puerto Rico Anolis krugi, A. pulchellus (both shown above) and A. poncensis (shown below) are closely related bush anoles. However, they live in dense mesic forests, open savanna, and in xeric forest and semi-desert, respectively. Thus, they are seldom found in the same general area, and almost never in the same microhabitat. The same general phenomenon of niche segregation can be seen in all the Greater Antilles and, to a lesser degree, in The Bahamas.

 


Ponce small-fanned anole, Anolis poncensis. First two photographs: males. Guanica State Forest, south-western Puerto Rico.

Third photograph: female. Camp Santiago, Salinas, south-eastern Puerto Rico.

The species has a merely vestigial dewlap, totally covered in large keeled scales, hence its common name.

Grass-bush ecomorph.

 

    A number of anole species and their eggs have been shipped accidentally as stowaways in cargo sent among islands or regions, and have invaded areas where they are not native. In southern Florida, United States, several Caribbean species have found a home in both the natural sub-tropical vegetation and residential areas. Likewise, the North American green anole, Anolis carolinensis, one of the few temperate zone species, is now firmly established in places as far as Barcelona, Spain.

 

    Unfortunately, some pet dealers have created confusion among inexperienced hobbyists by inventing names for anoles whose identity or origins they ignore. Pet-trade names like "Cuban anole" (at times applied to Anolis equestris, below) mean next to nothing, since there are dozens of species of "Cuban" anoles, to begin with. Other names, like Anolis "barbadensis" (perhaps referring to the species endemic to Barbados, shown further down) are figments of someone's imagination.

 


Cuban giant anole, Anolis equestris, female. Captive individual from a population introduced in Miami, Florida.

Crown giants with short legs, like this species, are crawlers.


Western giant anole (nominate race, Anolis luteogularis luteogularis), female. Sierra del Rosario, western Cuba.
This is another member of the equestris group of Cuban giant anoles.
Crown giant ecomorph.

(Photograph courtesy of Mr. Joseph Burgess).



Western giant anole (extreme western race, Anolis luteogularis jaumei), juvenile. Guanahacabibes Peninsula, western Cuba.
(Photograph courtesy of Mr. Joseph Burgess).

 

Green-blotched giant anoles, (Guantanamo race, Anolis smallwoodi palardis), female and mating pair.

Guantanamo Naval Base, Guantanamo, south-eastern Cuba.

Crown giant ecomorph.

(Photograph courtesy of Mr. Joseph Burgess).

 

Baracoa giant anole, Anolis baracoae. Rio Yurumi, south-eastern Cuba.

Crown giant ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).


Hispaniolan northern green anoles, (southern Santo Domingan race Anolis chlorocyanus cyanostictus) male and female.
National Botanical Garden, Santo Domingo, southern Dominican Republic, Hispaniola.

Trunk-crown ecomorph.

 

Hispaniolan northern green anoles (nominate race, Anolis chlorocyanus chlorocyanus) female.

Central Mountain Range Dominican Republic, Hispaniola.
This species is also one of several introduced into Florida, United States.
Trunk-crown ecomorph.

 

Jamaican turquoise anoles, Anolis grahami, five males and a female.

First photograph: Anchovy, north-western Jamaica.

Next three photographs: Windsor, north-central Jamaica.
Fourth photograph: Hellshire Hills, south-eastern Jamaica. (Courtesy of Mr. Joseph Burgess).

Last photograph, Port Antonio, north-eastern Jamaica.

Trunk-crown ecomorph.

 


Bahamian green anoles, (Grand Bahama bank race, Anolis smaragdinus smaragdinus), males.
First photograph: southern Andros Island, The Bahamas.
Second photograph: Eleuthera, The Bahamas.

Trunk-crown ecomorph.

(Photographs courtesy of Mr. Joseph Burgess).

 

Hispaniolan southern green anole (nominate race Anolis coelestinus coelestinus), male and female. Pedernales, south-western Dominican Republic, Hispaniola.

Trunk-crown ecomorph.

 

Hispaniolan montane bush anole, Anolis etheridgei, females. Ebano Verde Nature Reserve, Central Mountain Range, Dominican Republic, Hispaniola.

Habitat generalist.

(Photographs courtesy of Mr. Eladio M. Fernandez).

 

Central pallid anole, Anolis centralis, female. Rio Yurumi, south-eastern Cuba.

Grass-bush ecomorph.

(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Brown red-bellied anole, Anolis koopmani, male. Near Les Cayes, south-western Haiti, Hispaniola.

Grass-bush ecomorph.

(Photographs courtesy of Mr. Eladio M. Fernandez).

 

Barahonan grass anoles, Anolis alumina, male and female. Bahoruco National Park, south-western Dominican Republic, Hispaniola.

Grass-bush ecomorph.

 

    Lesser Antillean anoles fall into two natural groups. The ones of the subgenus Ctenonotus range from the northern Lesser Antilles south to Dominica. Those of the subgenus Dactyloa are found from Martinique south to northern South America.


Saint Christopher's Bank tree anoles, Anolis bimaculatus, male and female. Oranjestad, western Saint Eustatius, Lesser Antilles.


Saint Christopher's Bank tree anoles, Anolis bimaculatus, male and female. Oualie Beach, north-western Nevis, Lesser Antilles.


Barbudan Bank tree anole, Anolis leachii, displaying male and female. From a population introduced in Charlotte Amalie, southern Saint Thomas, United States Virgin Islands.
One of the Lesser Antillean giant anoles, this species is endemic to the Antigua-Barbuda island bank.
 
Habitat generalist.



Anguillan Bank tree anoles, Anolis gingivinus, two males and a female.
First two photographs: near Marigot, north-western Saint Martin, Lesser Antilles.
Third photograph: southern Siint Maartin, Lesser Antilles.

Habitat generalist.

 


Saban anole, Anolis sabanus, three males and a female.
First, second and third photographs: slopes of Mount Scenery, central Saba, Lesser Antilles.

Third photograph: Windwardside, central Saba, Lesser Antilles.

Habitat generalist.

 

Dominican anoles (north-western race, Anolis oculatus cabritensis), two males and a female. Cabrits National Park, north-western Dominica, Lesser Antilles.

Habitat generalist.

Above and below: Dominican anoles (highland race, Anolis oculatus montanus) males. Morne Trois Pitons National Park, south-central Dominica, Lesser Antilles.

Habitat generalist.
(Third photograph courtesy of Mr. Mel Jose Rivera).


Guadeloupan anole (nominate race, Anolis marmoratus marmoratus), male. Eastern Basse Terre, Guadeloupe, Lesser Antilles.
(Photograph courtesy of Mr. Joseph Burgess).



Anolis marmoratus alliaceous, male. central Basse Terre, Guadeloupe, Lesser Antilles.
(Photograph courtesy of Mr. Joseph Burgess).



Anolis marmoratus girafus, male. Western Basse Terre, Guadeloupe, Lesser Antilles.
(Photograph courtesy of Mr. Joseph Burgess).


Guadeloupan anole, Anolis marmoratus speciosus, captive male.

(Photograph courtesy of Mr. Joseph Burgess).

 

Grenadian tree anoles, Anolis richardi, males.

First two photographs: Grand Anse, south-western Grenada, Lesser Antilles.

Last photograph: Mount Hartman Bay, south-western Grenada, Lesser Antilles.

This is another Lesser Antillean species that is habitat generalist. However, it approaches the Greater Antillean crown giants in its ecology.


Grenadian tree anoles, Anolis richardi, females.

First photograph: Mount Hartman Bay, south-western Grenada, Lesser Antilles.

Second photograph: Grand Anse, south-western Grenada, Lesser Antilles.
Habitat generalist.

 

Saint Vincent's tree anoles, Anolis griseus, two males and a female.

First photograph: Kingstown Botanical Garden, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.

Last two photographs: Layou Valley, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.

Habitat generalist, but could be considered a crown giant anoline.

 


Saint Lucian anoles (Anolis luciae), male and female. Millet Nature Reserve, central Saint Lucia, Lesser Antilles.
  Habitat generalist.



Martinique's anole (nominate race, Anolis roquet roquet, male. near Fort-du France, southern Martinique, Lesser Antilles.

Habitat generalist.
(Photograph courtesy or Mr. Joseph Burgess).


Anolis roquet zebrilus, male. North-western Martinique, Lesser Antilles.
(Photograph courtesy of Mr. Joseph Burgess).

Grenadian bush anoles, Anolis aeneus, male and female. Saint Georges, south-western Grenada, Lesser Antilles.

Habitat generalist.

 

Another color morph, of the Grenadian bush anole, male and female. Grand Anse, south-western Grenada, Lesser Antilles.
Habitat generalist.



Grenadian bush anoles, Anolis aeneus, male and female. Union, Saint Vincent and the Grenadines, Lesser Antilles.

Habitat generalist.
(Photographs courtesy of Mr. Mel Jose Rivera).

 

Saint Vincent's bush anole, Anolis trinitatis, two males and female. Kingstown, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.

Habitat generalist.



Barbadian anoles, Anolis extremus, male and female. From a population introduced into Castries, north-western Saint Lucia, Lesser Antilles.
  Habitat generalist.



Barbudan bush anole, Anolis wattsi, two males and a female.
From a population introduced into Castries, north-western Saint Lucia, Lesser Antilles.
  Habitat generalist, althought it is similar to Greater Antillean trunk-ground groups.



Anguillan Bank bush anoles, Anolis pogus, male and female. Near Marigot, north-western Saint Martin.
  Habitat generalist, approaching trunk-ground ecomorph.



Statian Bank bush anole, Anolis schwartzi, male and two females.
First two photographs, Oualie Beach, north-western Nevis, Lesser Antilles.
  Third photograph: The Quill National Park, southern Saint Eustatius, Lesser Antilles.
Habitat generalist, approaching trunk-ground ecomorph.


   Some anole species range widely, often almost throughout every habitat in an island bank. In fact, a few species have actually benefited enormously from man-made forest clearings and urban development. People who have seen the high numbers of Puerto Rican crested anoles and Hispaniolan stout anoles that can be found in some areas may find it difficult to appreciate that the species was probably quite rare in pre-Columbian times. The only reason why today they are almost omnipresent all over their islands is, indirectly, man himself.



Puerto Rican crested anole, Anolis cristatellus cristatellus, male. Florida, central Puerto Rico.
Trunk-ground ecomorph.



A male Anolis cristatellus cristatellus contorts its body to get rid of an ant (out of view) that repeatedly bit its tail.
Guanica State Forest, south-western Puerto Rico.

 


Puerto Rican crested anole, (eastern Puerto Rican Bank race, Anolis cristatellus wileyeae
), male, and female.

First photograph: Guana Island, British Virgin Islands.

Second photograph: Mount Sage, west-central Tortola, British Virgin Islands.
Trunk-ground ecomorph.



Derived from a rather recent invasion of The Bahamas from distant Puerto Rico, the southern Bahamian anole,
(nominate race Anolis scriptus scriptus) is very similar to its parent species, Anolis cristatellus (higher up the page).

Trunk-ground ecomorph.
(Photograph courtesy of Mr. Joseph Burgess).

 

Southern Bahamian anoles, (Turks and Caicos race, Anolis scriptus mariguanae). Male and female.

First photograph: Six Hill Cay, Turks and Caicos.

Second photograph: Big Ambergris Cay, Turks and Caicos.

(Photographs courtesy of Mr. Joseph Burgess).



Southern Bahamian anole, (Great Inaguan race, Anolis scriptus leucopheus), male and female.

Great Inagua, The Bahamas.
(Photographs courtesy of Mr Joseph Burgess).



Hispaniolan stout anoles (nominate race, Anolis cybotes cybotes), three males and a female. First three photographs: Santo Domingo, southern Dominican Republic, Hispaniola.

Last photograph: San Rafael, south-western Dominican Republic, Hispaniola.
This and related species are also called "large-headed anoles" because of the proportionally enormous heads of old males.

This is one of the most common and widespread species in its home island.
Trunk-ground ecomorph.

 

Black-throated stout anole, Anolis armouri, female. Bahoruco National Park, south-western Dominican Republic, Hispaniola.
Trunk-ground ecomorph.

 


Red-fanned stout anole, Anolis marcanoi, male and female. San José de Ocoa, south-central Dominican Republic, Hispaniola.
Trunk-ground ecomorph.

 

Barahona stout anole (mainland race, Anolis longitibialis specuum), two males and two females. Jaragua National Park, south-western Dominican Republic, Hispaniola.
Trunk-ground ecomorph.

 

Bahoruco stout anole, Anolis strahmi, male. Bahoruco Mountains, south-western Dominican Republic, Hispaniola.

Trunk-ground ecomorph.
(Photograph courtesy of Mr. Eladio M. Fernandez).

 

Cuban brown anoles, Anolis sagrei, two males and a female.

From a population introduced into Kingstown, south-western Saint Vincent, Saint Vincent and the Grenadines, Lesser Antilles.
Trunk-ground ecomorph.



Spanish flag anole, Anolis allogus, male. Sierra del Rosario, western Cuba.
(Photograph courtesy of Mr. Joseph Burgess).



 
    Other anoles have very restricted natural ranges and are very susceptible to the alteration of their habitat. The Virgin Islands' giant anole, Anolis roosevelti, has not been seen with certainty for decades, and may have totally disappeared from its natural range of the islands of Culebra and Vieques, and the northern Virgin Islands. Others, like Cook's pallid anole, Anolis cooki, of mainland Puerto Rico, have apparently always endured intense natural competitive pressure from other species, but such is now exacerbated by modification and destruction of habitat within their limited geographical distributions.

 

    Some species that are very partial to a specific habitat are, of course, endangered by environmental destruction but may recover easily when their habitat is allowed to replenish. At the beginning of the XX century, when Puerto Rico was all but totally deforested, the Puerto Rican giant anole was feared extinct, since it had not been observed for many years. The species needs closed-canopy mesic forests in order to survive, and such had been by then cut down to make way for sugar cane and tobacco. However, with the collapse of agriculture as the main economic force in the island and the subsequent growth of its secondary forests, the lizard made a remarkable comeback and now it is again quite common in many areas of Puerto Rico.

 

    This example only proves that allowing ecosystems to subsist, even if only partially, may ensure the continued existence of many of the biological treasures in the Caribbean and elsewhere. Sadly, now one can only dream of all the species of anoles and other taxa that have gone extinct in places like Haiti, most of which is today denuded of natural vegetation, and where even the few national parks lack effective protection and are under intense agricultural pressure.

 

Carrot Rock's anoles, Anolis ernestwilliamsi, male and females. Captive individuals from Carrot Rock, British Virgin Islands.
This lizard is endemic to the tiny islet that gives it its name, and has one of the most
restricted natural distributions among vertebrates. It is closely related to Anolis cristatellus (above).
Trunk-ground ecomorph.

 


Guanica pallid anole, Anolis cooki, two males and and two females. Guanica State Forest, south-western Puerto Rico.
An inhabitant of a few square kilometers of coastal desert and littoral xeric forest in the south-western region of that island.
It is very similar to Anolis cristatellus cristatellus, with which it lives in sympatry. It can, however, be distinguished
color wise by the vertically-arranged, tiny yellow spots on its flanks, by the more sharply defined dorsal and ventral colors and, mostly, by its squinty eyes with white lids.
The similarity is still such that it causes very strong competition between them. This conflict Cook's anole seems
to be loosing, but only because its limited habitat has been seriously degraded by man, which turns competition against it.

Like many vertebrates of xeric regions, this species is  very pale in color.
Trunk-ground ecomorph.

 

Monan anoles, Anolis monensis, male and female. Mona Island.

This species is closely related to Anolis cooki (above) and, like it, it is a xerophile.

Trunk-ground ecomorph.

(Photograph courtesy of Mr. Joseph Burgess).

 

Another xerophile with light color: the pallid stout anole (nominate race, Anolis whitemani whitemani) male. Azua, southern Dominican Republic, Hispaniola.

Trunk-ground ecomorph.

 

Yet another xerophile species: the Hispaniolan desert grass anole Anolis olssoni, subspecies extentus, female. Azua, south-western Dominican Republic, Hispaniola.

Grass-bush ecomorph.



Jamaican Gray anoles (nominate race, Anolis lineatopus lineatopus), male and female. Hellshire Hills, south-eastern Jamaica.
(Photographs courtesy of Mr. Joseph Burgess).



Jamaican gray anoles (northern race, Anolis lineatopus merope), male and female. Windsor, north-central Jamaica.
Trunk-ground ecomorph.



Jamaican gray anoles (eastern race, Anolis lineatopus neckeri), male and female. Treasure Beach, south-western Jamaica.



Blue Mountains anole, Anolis reconditus, male. Blue Mountains, east-central Jamaica.
Ecomorphic generalist.



"Cliff" anolines are mostly restricted to Cuba.

Anolis bartschi, male, female and juvenile.  Sierra de los Organos, western Cuba.
The species inhabits limestone cliffs.
(Photographs courtesy of Mr. Joseph Burgess).

 

    Altogether, anoles are so abundant in the West Indies that they form an enormously important component of their ecosystems. Their immense numbers are not matched by those of the species that inhabit the American continents. One who is familiar with their omnipresence in the Antilles can be baffled by their relative scarcity in many places of Central and South America. In those other regions as many species as in the Caribbean islands can be found, yet one might only see two or three individual lizards in one day. By comparison, in some lowland forests and grasslands of the Antilles, two or three dozens of anoles can be seen in a matter of minutes. This phenomenon can be at least partially explained by the fact that in the continents, ecological communities are much more complex. Anoles there might be subjected to far more pressure from organisms not found in the islands, which compete or prey upon them, thus reducing their numbers and life spans.