Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Anolis wattsi
(Watts' anole)

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Datasheet

Anolis wattsi (Watts' anole)

Summary

  • Last modified
  • 21 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Anolis wattsi
  • Preferred Common Name
  • Watts' anole
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Reptilia
  • Summary of Invasiveness
  • A. wattsi is a small anole lizard native to Antigua and its offshore islets. There have been two known introductions, on St Lucia and Trinidad, and in both cases A. wattsi is spreading ra...

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Pictures

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PictureTitleCaptionCopyright
Watts' anole (Anolis Wattsi schwartzi) on St.Kitts. November 2012
TitleAdult
CaptionWatts' anole (Anolis Wattsi schwartzi) on St.Kitts. November 2012
Copyright©Clinton Robertson - CC BY 2.0
Watts' anole (Anolis Wattsi schwartzi) on St.Kitts. November 2012
AdultWatts' anole (Anolis Wattsi schwartzi) on St.Kitts. November 2012©Clinton Robertson - CC BY 2.0

Identity

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Preferred Scientific Name

  • Anolis wattsi Boulenger 1894

Preferred Common Name

  • Watts' anole

Other Scientific Names

  • Anolis alter Williams, 1962
  • Ctenonotus wattsi Nicholson et al., 2012

Local Common Names

  • Saint Lucia: zanndoli

Summary of Invasiveness

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A. wattsi is a small anole lizard native to Antigua and its offshore islets. There have been two known introductions, on St Lucia and Trinidad, and in both cases A. wattsi is spreading rapidly over large parts of the islands. A. wattsi is considered invasive on St Lucia (Krauss, 2010), where it is apparently replacing A. luciae, especially near man-made features (such as walls, buildings and culverts), in urban and suburban areas, and in some disturbed secondary forest areas. However, widespread adverse effects on St Lucia have not been noted so far. It is not threatened in its native Antigua.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Chordata
  •             Subphylum: Vertebrata
  •                 Class: Reptilia
  •                     Order: Sauria
  •                         Family: Iguanidae
  •                             Genus: Anolis
  •                                 Species: Anolis wattsi

Notes on Taxonomy and Nomenclature

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Lazell (1972) separated A. wattsi into four subspecies, based on their distribution on different islands or island banks. Schneider et al. (2001) recognized the four forms as separate species, with A. wattsi confined to Antigua and cays (offshore islets). The other three species are A. forresti Barbour 1923 on Barbuda, A. pogus Lazell 1972 on St Martin, and A. schwartzi Lazell 1972 on the islands of the St Kitts bank.

Of these four species, introduced populations have all apparently been identified as A. wattsi, as currently recognized, and this is the only species discussed in this datasheet. References to the other former subspecies are only noted here if they were recorded under the name A. wattsi. The proposal to place A. wattsi in the new genus Ctenotus was refuted by Poe (2013).

A. wattsi is part of the bimaculatus group of Anolis species of the northern Lesser Antilles (Stenson et al., 2004). There are now no subspecies of A. wattsi recognized.

The common name zanndoli is reported on St Lucia, but this is a general name applied to all three Anolis species on the island (Daltry, 2009).

Description

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A. wattsi is a small anole lizard, snout-vent (body excluding tail) length up to 58 mm in males and 46 mm in females. Typically a bright and very pretty anole.

Males are tan, brown or olive, with or without faint darker transverse bands. Head with rust red pigment, top of head may be suffused with blue. Underside cream to yellow, dewlap white, yellow or orange. Females brown to grey-brown, with or without yellow lateral suffusion, duller than males, with pale middorsal and flank stripes (Schwartz and Henderson, 1991). Lazell (1972) and Schwartz and Henderson (1991) gave diagnostic details of scalation.

The type specimen is BMNH 1946.8.29.12-13, belonging to the British Museum of Natural History, London.

Distribution

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It is widespread and abundant in its native Antigua and offshore islets (Daltry, 2007), where it is found on every coastal cay that supports more than herb-stage vegetation (Lazell, 1972). There have been no attempts to locate the area of origin of any of the introduced populations.

White and Hailey (2006) and Daltry (2009) provided maps of the distribution of the introduced populations of A. wattsi in Trinidad and St Lucia, respectively.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 10 Jan 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

North America

Antigua and BarbudaPresent, WidespreadNativeDaltry (2007)Common on Antigua and cays. The anole on Barbuda now recognized as a separate species, A. forresti.
Saint LuciaPresent, WidespreadIntroducedInvasiveDaltry (2009); Underwood (1959); Gorman (1976)Widespread. Displacing the native Anolis luciae; First reported: 1950s
Trinidad and TobagoPresent, LocalizedIntroducedInvasiveWhite and Hailey (2006); Boos (1996)Expanding rapidly in Central Trinidad; First reported: 1990s

History of Introduction and Spread

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There have been two introductions of A. wattsi (Kraus, 2009). A third introduction of A. wattsi, to Anguilla for research purposes (Kraus, 2009), is now assigned to A. pogus (Roughgarden et al., 1984).

In St Lucia, A. wattsi was first observed in the Botanic Gardens in Castries, in 1956 (Underwood, 1959). It was restricted to the Botanic Garden in Castries during the 1960s, when it was seen there by Lazell (1972) and Gorman (1976). Gorman (1976) did not see A. wattsi in the Botanic Gardens in 1972 and 1973, but notes that Rand and Kiester (personal communication) had observed it at another location in Castries in 1970. This is the only introduction recorded by Lever (2003).

After this slow start, A. wattsi spread rapidly in St Lucia, especially along the coastline (Daltry, 2009). Sampling in 2009 showed that it had extended its range to about half of the island, from lowland areas (urban, suburban, and disturbed seasonal forests), to an altitude of 259 m at Millet, on the edge of the rainforest. Daltry (2009) recorded A. wattsi in 14 out of 55 forest plots sampled, compared to 51 plots for the native A. luciae, accounting for 183 and 507 individuals, respectively.

A. wattsi is apparently replacing A. luciae in St Lucia, especially near man-made features (such as walls, buildings and culverts), in urban and suburban areas, and even in some disturbed secondary forest areas. There was a significant positive association (P = 0.026) between A. wattsi and the proximity of man-made features in the environment, in contrast to the native A. luciae, which showed no association (P = 0.34) (Daltry, 2009).

In Trinidad and Tobago, A. wattsi was first observed at the Caroni agricultural research station in Waterloo, Central Trinidad, in 1992 (Boos, 1996). Surveys in 2000-2004 showed it was spreading rapidly in five urban centers, over an area of about 75 square km (White and Hailey, 2006). Self-dispersal at a rate of 100 m/year and jump dispersal over distances greater than 1 km/year were both implicated in this spread. The initial introduction seems to have been on pineapple plants imported from Antigua direct into the agricultural research station (White and Hailey, 2006). A. wattsi may be displacing another introduced anole, A. aeneus, in disturbed areas, but has not entered natural habitats and there is not yet any interaction with the native A. chrysolepis (White and Hailey, 2006). A. wattsi has continued to spread through central and western Trinidad since 2004 (A Hailey, personal observations).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Saint Lucia Antigua and Barbuda 1950s Horticulture (pathway cause) Yes Gorman (1976) To Botanic Gardens in Castries
Trinidad and Tobago Antigua and Barbuda 1990s Horticulture (pathway cause) Yes White and Hailey (2006) To Caroni agricultural research station

Risk of Introduction

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There is no significant commercial trade in A. wattsi as pets, at least into or through the USA (Powell et al., 2011). There is always some risk of further introductions to the wider Caribbean region via general shipping or the horticultural trade, either from Antigua or from the introduced populations in the port of Castries on St Lucia, or on horticultural materials exported from Trinidad and Tobago.

Habitat

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Native populations occupy most terrestrial habitats, from xeric scrub to rainforest, and are frequently found in wet, shady pockets (Schwartz and Henderson, 1991). The introduced population on St Lucia has spread into disturbed forest (Daltry, 2009), but the introduced population on Trinidad is currently restricted to urban, suburban and agricultural areas (White and Hailey, 2006). Lizards are usually seen on trees, bushes, fence posts and other man-made features, up to 1 m from the ground, with females and juveniles more likely to be seen on the ground (Lazell, 1972). This species retreats into holes and crevices, rather than climbing trees as many other anoles do (Lazell, 1972).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Principal habitat Natural
Protected agriculture (e.g. glasshouse production) Present, no further details Natural
Managed forests, plantations and orchards Principal habitat Natural
Disturbed areas Principal habitat Natural
Rail / roadsides Principal habitat Natural
Urban / peri-urban areas Principal habitat Natural
Buildings Principal habitat Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Secondary/tolerated habitat Natural
Natural grasslands Present, no further details Natural
Riverbanks Secondary/tolerated habitat Natural
Rocky areas / lava flows Secondary/tolerated habitat Natural
Scrub / shrublands Secondary/tolerated habitat Natural
Littoral
Coastal areas Secondary/tolerated habitat Natural
Coastal dunes Present, no further details Natural

Biology and Ecology

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Genetics

A. wattsi (and other members of the bimaculatus group) has a karyotype with 2n = 29 in males and 2n = 30 in females, with both an acrocentric X1 chromosome and X2microchromosome (Gorman and Atkins, 1969). Females are thus X1X1X2X2 and males X1X2Y. These species show major karyotype differences to Anolis species of the roquet group (2n = 34 or 36), and A. wattsi is therefore unlikely to hybridize with other anoles on St Lucia (Giannasi et al., 1997) or Trinidad (White and Hailey, 2006).

Reproductive Biology

All female anoles lay a clutch of a single egg, often at intervals of only 1-2 weeks in productive habitats (Andrews and Rand, 1974). A. wattsi breeds year round.

Physiology and Phenology

Minimum size (snout-vent length) of mature individuals is 39 mm in males and 36 mm in females (Lazell, 1972).

Longevity

No specific information available. As a small anole, a life expectancy of 1-2 years is likely (Losos, 2009).

Activity Patterns

A. wattsi is active throughout the year, and does not hibernate or aestivate. Daily activity is from 0500-1900 h, although it is most active early (0600 h) and late (1800 h), with reduced activity during hot midday hours (Kolbe et al., 2008; Henderson and Powell, 2009). The behavioural observations of Rummel and Roughgarden (1985) on A. wattsi would now be assigned to Anolis schwartzi.

Population Size and Structure

A. wattsi is abundant all over Antigua, its native island (Lazell,1972). Population densities up to 7143/ha have been recorded in forested parts of Great Bird Island (Daltry, 2007), and overall densities of 1700/ha, 5627/ha, 4556/ha 8760/ha 4700/ha were recorded on Green Island, Great Bird Island, Maiden Island, Rabbit Island and Red Head Island, Antigua, respectively (Henderson and Powell, 2009). Densities on St Lucia are as high as 7200/ha, with an overall mean density of 332/ha (Daltry, 2009); the peak value of more than 9900/ha for A. wattsi reported in Daltry (2009) would now be assigned to Anolis schwartzi. Population density estimates for A. wattsi by Heckel and Roughgarden (1979) and McLaughlin and Roughgarden (1989) would now be assigned to Anolis pogus. Population density estimates for A. wattsi by Roughgarden et al. (1983) would now be assigned to Anolis pogus, A. schwartzi and A. forresti.

Nutrition

A. wattsi feeds on ants, spiders, crickets, cockroaches, grasshoppers and insect larvae, usually caught on the ground. It also eats soft fruits, including mangoes (Daltry, 2009).

Host Associations

The rhabditid nematode Rhabdias sp. (Dobson et al., 1992) and the oxyurid nematodes Parapharyngodon cubensis and Spauligodon anolis (Dobson et al., 1992; Goldberg et al., 1997, in Bursey et al., 2012) have been reported from A. wattsi. Dobson et al. (1992) also reported the cystacanth of an undetermined centrorhynchid acanthocephalan from A. wattsi.

The nematodes Trichospirura teixeirai and larval Abbreviata sp. (Goldberg et al., 1997, in Henderson and Powell, 2009) and an unidentified physalopterid nematode (Dobson et al., 1992, in Henderson and Powell, 2009) are also reported from A. wattsi. A higher diversity of parasites were found in individuals in forests (0.93 parasite species per host) than in xeric habitat (0.40) on Antigua (Dobson and Pacala, 1992, in Henderson and Powell, 2009). Poulin et al. (2003) noted four species of metazoan parasites in a sample of 20 A. wattsi individuals, although without giving details of taxa or location involved.

Staats and Schall (1996) examined the blood of 22 A. wattsi individuals from Antigua but found no Plasmodium parasites. Schall and Staats (1997) examined 13 male A. wattsi individuals from Antigua and reported that 62% of them were infected with helminths (four species, not named), but none with Plasmodium or haemogregarines. Reports of the malarial parasite Plasmodium azurophilum in A. wattsi (Schall, 1992) would now be assigned to Anolis pogus.

Environmental Requirements

A. wattsi is a habitat and thermal generalist. Environmental requirements may be modified by competition/predation by other lizards on Antigua. A. wattsi uses sunny habitats more frequently on islets without the lizard Ameiva griswoldi, and shady habitats more frequently on islets without Anolis leachii (Kolbe et al., 2008). Niche observations for A. wattsi by Roughgarden et al. (1983) would now be assigned to A. forresti, A. pogus and A. schwartzi.

Climate

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ClimateStatusDescriptionRemark
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Am - Tropical monsoon climate Preferred Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
10-18

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Alsophis antiguae Predator Adult Female/Adult Male/Juvenile not specific
Ameiva griswoldi Predator Adult Female/Adult Male/Eggs/Juvenile not specific
Anolis leachii Predator Adult Female/Adult Male/Juvenile not specific
Herpestes javanicus Predator Adults not specific N

Notes on Natural Enemies

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On Antigua A. wattsi is preyed upon by the racer snake Alsophis antiguae (Daltry et al., 2001) and the lizards Ameiva griswoldi and Anolis leachii (Kolbe et al., 2008). On St Lucia it is preyed on by the mongoose Herpestes javanicus (Daltry, 2009).

Means of Movement and Dispersal

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Natural Dispersal

The mean home range size is 1.8-9.5 square meters (Henderson and Powell, 2009). The rate of natural spread is about 100m/year in Trinidad (White and Hailey, 2006).

Accidental Introduction

Most likely on horticultural produce, nursery plants, building materials or general shipping. The rate of jump dispersal on such materials is greater than 1 km/year in Trinidad (White and Hailey, 2006).

Intentional Introduction

The reported introduction of A. wattsi to Anguilla for research purposes would now be assigned to A. pogus (Roughgarden et al., 1984).

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Crop productionJump dispersal at distances greater than 1 km/year in Trinidad Yes White and Hailey, 2006
HitchhikerLikely for St Lucia introduction, and jump dispersal within Trinidad Yes Yes Gorman, 1976; White and Hailey, 2006
HorticultureProbably on pineapple plants imported into Trinidad & Tobago Yes White and Hailey, 2006
Self-propelledDispersal at rate of 100 m/year in Trinidad Yes White and Hailey, 2006

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Bulk freight or cargoAdults, juveniles or eggs possible Yes
Containers and packaging - woodAdults, juveniles or eggs possible Yes
Plants or parts of plantsAdults, juveniles or eggs possible Yes White and Hailey, 2006

Environmental Impact

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Impact on Biodiversity

Replacement of or addition to other Anolis species is the only likely impact of A. wattsi introductions. A. wattsi is unlikely to invade rainforest on St Lucia, but if it does, it would reduce populations of the endemic anole A. luciae nationwide (Daltry, 2009). There was a weak negative correlation between densities of these two anole species in 100 square metre forest plots (r = −0.166); the plot with the most A. wattsi (n = 72) had no A. luciae, and plots with the highest number of A. luciae (> 30) had no A. wattsi (Daltry, 2009). Hybridisation between A. wattsi and other anoles on St Lucia or Trinidad is unlikely due to their very different karyotypes (Giannasi et al., 1997; White and Hailey, 2006).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Anolis luciaeNo detailsSaint LuciaCompetition - monopolizing resourcesDaltry, 2009

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Capable of securing and ingesting a wide range of food
  • Highly mobile locally
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
Impact outcomes
  • Conflict
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Interaction with other invasive species

Uses List

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General

  • Research model

Detection and Inspection

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A. wattsi is likely to be first noticed in a new location as an unusual anole unlike other species existing in the area. Most areas suitable for A. wattsi introduction will already have native or other introduced Anolis species. Schwartz and Henderson (1991) is the most useful initial guide to identification, but identification should be confirmed by a taxonomic specialist as there are hundreds of Anolis species potentially involved in introductions. Lizards are usually seen on trees, bushes, fence posts and other man-made features, up to 1 m from the ground, with females and juveniles more likely to be seen on the ground (Lazell, 1972). This species retreats into holes and crevices, rather than climbing trees as many other anoles do (Lazell, 1972).

Similarities to Other Species/Conditions

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There are approximately 400 species of anoles (Nicholson et al., 2012), so confirmation of new invasions of A. wattsi or a similar species requires a taxonomic specialist. Schwartz and Henderson (1991) remains the most useful guide for initial recognition of introduced anoles.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Control

The introduced mongoose (Herpestes javanicus) preys on A. wattsi in St Lucia, but apparently not enough to have a significant impact on their numbers (Daltry, 2009). A. wattsi is likely to be adversely affected by use of pesticides (Daltry, 2009).

Daltry (2009) regarded island-wide eradication of A. wattsi from St. Lucia to be neither necessary nor feasible, but suggested that removal by hand could be effective on a local scale. Her suggestions for management of the species were:

  1. Listing as Unprotected under the Wildlife Protection Act.
  2. Monitor the distribution and be vigilant for it invading mature, natural forests.
  3. Prevent this species from invading the offshore islands, especially the Maria islands, by screening all boats and baggage.
  4. Support reptile conservation in other countries by prohibiting, screening and removing anole lizards from exported cargo.

Gaps in Knowledge/Research Needs

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Most work on introduced populations of A. wattsi has focused on basic biological questions rather than invasiveness or its consequences. This state is likely to continue as economic impacts of Anolis introductions are minor or negligible.

References

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Andrews R; Rand AS, 1974. Reproductive effort in anoline lizards. Ecology, 55:1317-1327.

Barbour T, 1923. West Indian investigations of 1922. Occasional Papers of the Museum of Zoology, 132:1-9.

Boos HEA, 1996. Some recent reptilian introductions to Trinidad. Living World, 1995-1996. 17-19.

Boulenger GA, 1894. Description of a new Anolis from Antigua, West Indies. Annals and Magazine of Natural History, Series 6, 14:375-376.

Bursey CR; Goldberg SR; Telford SR Jr; Vitt LJ, 2012. Metazoan endoparasites of 13 species of Central American anoles (Sauria: Polychrotidae: Anolis) with a review of the helminth communities of Caribbean, Mexican, North American, and South American anoles. Comparative Parasitology, 79(1):75-132. http://www.bioone.org/doi/abs/10.1654/4530.1

Daltry JC, 2007. An introduction to the herpetofauna of Antigua, Barbuda and Redonda, with some conservation recommendations. Applied Herpetology, 4:97-130.

Daltry JC, 2009. The Status and Management of Saint Lucia's Forest Reptiles and Amphibians. SFA 2003/SLU/BIT-04/0711/EMF/LC., Finland: FCG Fauna & Flora, 80 pp. http://www.bananatrustslu.com/index.php?link=doccentre&project=sfa2003

Daltry JC; Bloxam Q; Cooper G; Day ML; Hartley J; Henry M; Lindsay K; Smith BE, 2001. Five years' of conserving the 'world's rarest snake', the Antiguan Racer Alsophis antiguae. Oryx, 35:119-127.

Dobson AP; Pacala SV; Roughgarden JD; Carper ER; Harris EA, 1992. The parasites of Anolis lizards in the northern Lesser Antilles. I. Patterns of distribution and abundance. Oecologia, 91(1):110-117.

Dobson AP; Pacala SW, 1992. The parasites of Anolis lizards in the northern Lesser Antilles. II. The structure of the parasite community. Oecologia, 91(1):118-125.

Garman S, 1887. On West Indian reptiles. Iguanidae. Bulletin of the Essex Institute, 19:25-50.

Giannasi N; Thorpe RS; Malhotra A, 1997. Introductions of Anolis species to the island of St. Lucia, West Indies: testing for hybrids using multivariate morphometrics. Journal of Herpetology, 31:586-589.

Goldberg SR; Bursey CR; Cheam H, 1997. Gastrointestinal helminths of 12 species of Anolis lizards (Polychrotidae) from the Lesser Antilles, West Indies. Journal of the Helminthological Society of Washington, 64:248-257.

Gorman GC, 1976. Observations on the distribution of Anolis extremus (Sauria: Iguanidae) on St Lucia, West Indies - a 'colonizing' species. Herpetologica, 32:184-188.

Gorman GC; Atkins L, 1969. The zoogeography of Lesser Antillean Anolis lizards - an analysis based upon chromosomes and lactic dehydrogenases. Bulletin of the Museum of Comparative Zoology, 138:53-80.

Heckel DG; Roughgarden J, 1979. A technique for estimating the size of lizard populations. Ecology, 60:966-975.

Henderson RW; Powell R, 2009. Natural history of West Indian reptiles and amphibians. Gainesville, Florida, USA: Florida University Press, 495 pp.

Kolbe JJ; Colbert PL; Smith BE, 2008. Niche relationships and interspecific interactions in Antiguan lizard communities. Copeia, 2008:261-272.

Kraus F, 2009. Invading nature: Springer series in invasion ecology 4. Springer, 563 pp.

Krauss U, 2010. Critical situation analysis (CSA) of invasive alien species (IAS) status and management, Saint Lucia, 2010. Critical situation analysis of invasive alien species status and management. Union, St Lucia: Forestry Department, 102 pp.

Lazell JD, 1972. The anoles (Sauria, Iguanidae) of the Lesser Antilles. Bulletin of the Museum of Comparative Zoology, 143:1-115.

Lever C, 2003. Naturalized reptiles and amphibians of the world. Oxford, UK: Oxford University Press, 318 pp.

Losos JB, 2009. Lizards in an evolutionary tree. Berkeley, California, USA: University of California Press, 512 pp.

McLaughlin JF; Roughgarden J, 1989. Avian predation on Anolis lizards in the northeastern Caribbean: an inter-island contrast. Ecology, 70:617-628.

Nicholson KE; Crother BI; Guyer C; Savage JM, 2012. It is time for a new classification of anoles (Squamata: dactyloidae)? Zootaxa, 3477:1-108.

Poe S, 2013. 1986 Redux: new genera of anoles (Squamata: Dactyloidae) are unwarranted. Zootaxa, 3626:295-299.

Poulin R; Mouillot D; George-Nascimento M, 2003. The relationship between species richness and productivity in metazoan parasite communities. Oecologia, 137(2):277-285.

Powell R; Henderson RW; Farmer MC; Breuil M; Echternacht AC; Buurt Gvan; Romagosa CM; Perry G, 2011. Introduced amphibians and reptiles in the greater Caribbean: patterns and conservation implications. In: Conservation of Caribbean island herpetofaunas, volume 1: Conservation biology and the wider Caribbean [ed. by Hailey, A. \Wilson, B. S. \Horrocks, J. A.]. Leiden, Netherlands: Brill, 63-143.

Roughgarden J; Heckel D; Fuentes ER, 1983. Coevolutionary theory and the biogeography and community structure of Anolis. In: Lizard ecology: studies of a model organism [ed. by Huey, R. B. \Pianka, E. R. \Schoener, T. W.]. Cambridge, Massachusetts, USA: Harvard University Press, 371-410.

Roughgarden J; Pacala S; Rummel J, 1984. Strong present-day competition between the Anolis lizard populations of St Maarten (Neth. Antilles). In: Evolutionary ecology [ed. by Shorrocks, B.]. Oxford, UK: Blackwell Science, 203-220.

Rummel JD; Roughgarden J, 1985. Effects of reduced perch-height separation on competition between two Anolis lizards. Ecology, 66:430-444.

Schall JJ, 1992. Parasite-mediated competition in Anolis lizards. Oecologia, 92(1):58-64.

Schall JJ; Staats CM, 1997. Parasites and the evolution of extravagant male characters: Anolis lizards on Caribbean islands as a test of the Hamilton-Zuk hypothesis. Oecologia, 111:543-548.

Schneider CJ; Losos JB; Queiroz Kde, 2001. Evolutionary relationships of the Anolis bimaculatus group from the northern lesser Antilles. Journal of Herpetology, 35:1-12.

Schwartz A; Henderson RW, 1991. Amphibians and reptiles of the West Indies. Gainesville, Florida, USA: University of Florida Press, 720 pp.

Staats CM; Schall JJ, 1996. Malarial parasites (Plasmodium) of Anolis lizards: biogeography in the Lesser Antilles. Biotropica, 28(3):388-393.

Stenson AG; Thorpe RS; Malhotra A, 2004. Evolutionary differentiation of bimaculatus group anoles based on analyses of mtDNA and microsatellite data. Molecular phylogenetics and evolution, 32:1-10.

Underwood G, 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae). III. Revisionary notes. Bulletin of the Museum of Comparative Zoology, 121:191-226.

White GL; Hailey A, 2006. The establishment of Anolis wattsi as a naturalized exotic lizard in Trinidad. Applied Herpetology, 3:11-26.

Distribution References

Boos HEA, 1996. Some recent reptilian introductions to Trinidad. In: Living World, 1995-1996, 17-19.

Daltry JC, 2007. An introduction to the herpetofauna of Antigua, Barbuda and Redonda, with some conservation recommendations. In: Applied Herpetology, 4 97-130.

Daltry JC, 2009. The Status and Management of Saint Lucia's Forest Reptiles and Amphibians. In: SFA 2003/SLU/BIT-04/0711/EMF/LC, Finland: FCG Fauna & Flora. 80 pp. http://www.bananatrustslu.com/index.php?link=doccentre&project=sfa2003

Gorman GC, 1976. Observations on the distribution of Anolis extremus (Sauria: Iguanidae) on St Lucia, West Indies - a 'colonizing' species. In: Herpetologica, 32 184-188.

Underwood G, 1959. The anoles of the Eastern Caribbean (Sauria, Iguanidae). III. Revisionary notes. In: Bulletin of the Museum of Comparative Zoology, 121 191-226.

White GL, Hailey A, 2006. The establishment of Anolis wattsi as a naturalized exotic lizard in Trinidad. In: Applied Herpetology, 3 11-26.

Links to Websites

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WebsiteURLComment
Anole annalshttp://www.anoleannals.org/
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.
The Reptile Databasehttp://reptile-database.reptarium.cz/

Contributors

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06/05/13 Original text by:

Adrian Hailey, University of the West Indies, Trinidad and Tobago

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