vi Contents

Contents of Volume 2

Preface 1

Karim V.D. Hodge, Robert Powell and Ellen J. CenskyConserving the herpetofauna of Anguilla

3

Jennifer C. DaltryAn introduction to the herpetofauna of Antigua, Barbuda and Redonda,with some conservation recommendations

17

Charles R. Knapp, John B. Iverson, Sandra D. Buckner andShelley V. CantConservation of amphibians and reptiles in The Bahamas

53

Angela Fields and Julia A. HorrocksThe herpetofauna of Barbados: Anthropogenic impacts and conserva-tion status

89

G. Perry and G.P. GerberConservation of amphibians and reptiles in the British Virgin Islands:Status and patterns

105

A.C. Echternacht, F.J. Burton and J.M. BlumenthalThe amphibians and reptiles of the Cayman Islands: Conservationissues in the face of invasions

129

Anita Malhotra, Roger S. Thorpe, Eric Hypolite and ArlingtonJamesA report on the status of the herpetofauna of the Commonwealth ofDominica, West Indies

149

Robert Powell and Sixto J. IncháusteguiConservation of the herpetofauna of the Dominican Republic

167

Robert PowellConservation of the herpetofauna on the Dutch Windward Islands: St.Eustatius, Saba, and St. Maarten

189

Olivier Lorvelec, Michel Pascal, Claudie Pavis and Philippe Feld-mannAmphibians and reptiles of the French West Indies: Inventory, threatsand conservation

205

Robert W. Henderson and Craig S. BergThe herpetofauna of Grenada and the Grenada Grenadines: Conserva-tion concerns

239

Contents vii

Jacques Daudin and Mark de SilvaAn annotated checklist of the amphibians and terrestrial reptiles of theGrenadines with notes on their local natural history and conservation

259

Byron S. WilsonConservation of Jamaican amphibians and reptiles

273

Michel BreuilThe terrestrial herpetofauna of Martinique: Past, present, future

311

Rafael L. Joglar, Alberto O. Álvarez, T. Mitchell Aide, DianeBarber, Patricia A. Burrowes, Miguel A. García, Abimael León-Cardona, Ana V. Longo, Néstor Pérez-Buitrago, Alberto Puente,Neftalí Rios-López and Peter J. TolsonConserving the Puerto Rican herpetofauna

339

Robert Powell and Robert W. HendersonThe St. Vincent (Lesser Antilles) herpetofauna: Conservation concerns

359

R. Graham ReynoldsStatus, conservation, and introduction of amphibians and reptiles in theTurks and Caicos Islands, British West Indies

379

Renata J. Platenberg and Ralf H. Boulon, Jr.Conservation status of reptiles and amphibians in the U.S. VirginIslands

411

Status, conservation, and introduction of amphibians andreptiles in the Turks and Caicos Islands, British WestIndies

R. Graham Reynolds

Department of Ecology and Evolutionary Biology, University of Tennessee, 569 Dabney Hall,Knoxville, TN 37996-1610, USAE-mail: rgraham@utk.edu

Abstract. The Turks and Caicos Islands, located at the southern terminus of the Bahamian Archipela-go, contain a unique native reptile assemblage consisting of 13 species, nine of which are endemicspecies or subspecies. At least four species are listed as endangered by the International Union forConservation of Nature, though most species in the region have not been evaluated. In additionto the native reptiles, at least 10 herpetofaunal species are introduced in the islands, including twoamphibians. The native fauna of the region face a plenitude of threats, including feral predators suchas cats, loss of habitat due to development, unsustainable exploitation (sea turtles), habitat degradationby semi-feral herbivores, and malicious killing. Sea turtles continue to be legally exploited for meatand their shells, and nesting activity is declining. The endemic iguana, Cyclura carinata, has beenextirpated from most major islands in the archipelago and survives on isolated islands that lackintroduced vertebrate predators. The endemic boid Epicrates c. chrysogaster has been extirpated fromseveral islands and faces additional extirpations on highly developed islands such as Providenciales.Though iguanas have received a great deal of attention from conservationists, local government, andcitizens, little is known about the biology or conservation concerns of other unique species. Additionalresearch must be implemented to inform conservation agendas, and I provide recommendations toimprove the long-term survival of this unique herpetofaunal assemblage.

Key words: Amphibians; conservation; Cyclura carinata; Epicrates chrysogaster; introduced species;reptiles; sea turtles.

Introduction

The Turks and Caicos Islands (TCI; fig. 1), located at the southern terminus ofthe Bahamian Archipelago, contain a unique endemic reptile fauna (Reynoldsand Niemiller, 2010a) that is threatened by rapidly increasing development andpopulation growth. These islands are in the midst of a huge development push toaccommodate an exponentially increasing tourist market. Tourism increased nearly

378 R.G. Reynolds

Figure 1. Map of the Turks and Caicos Islands showing the approximate extent of the banks in lightgray, representing approximately the 10 m depth line. Both banks were emergent during the last glacialmaximum, and islands on each bank were part of a single emergent land mass. The banks are separatedby the deep (>1 km) Turks Island Passage and have never been joined.

90% between 1995 and 2005, and in the rush to accommodate these visitors morethan 20 resorts opened during the same period on the island of Providenciales alone(Tsui, 2005). Providenciales has become so crowded in the last 20 years that largenew developments are moving to neighboring islands in the archipelago, includingthe formerly undeveloped island of West Caicos where a new resort has recentlybeen completed. In addition, small islands such as Big Ambergris Cay are beingbought by foreign development companies and turned into exclusive vacation areas.

It has been demonstrated that human influences are causing drastic declinesamong populations of several species of the native reptiles in this region (Iverson,1978). In addition, the number of introduced herpetofaunal species has recently be-come nearly equal to the number of native reptile species (Reynolds and Niemiller,2010a). Here I provide a comprehensive review of the herpetofauna of the TCI,which includes 13 native and 10 introduced species, as well as a discussion of con-servation concerns faced by individual species and the fauna as a whole.

The TCI (20◦59′-21◦58′ N and 70◦55′-72◦30′ W) consist of more than 200 islandsranging in size from <1 ha to >12,000 ha; plus many small rock islets distributed ontwo shallow banks, the Turks bank to the east and the Caicos bank to the west (figs 1

Amphibians and reptiles in the Turks and Caicos Islands 379

Figure 2. Names of major islands in the Turks and Caicos Archipelago.

and 2). These banks consist of platforms of marine sediments overlaid with ooliticlimestone dating back at least 150 my (Sealey, 2006), and for much of this periodthe banks likely remained near the surface due to reef building (Hedges, 2006).Islands on the banks have likely been ephemeral throughout the Pleistocene as thebanks were variously inundated and emergent (Sealey, 2006). The Turks and Caicosbanks existed as emergent islands at the height of the last (Wisconsin) glaciations17,000 years before present, though the two banks are separated by the narrowand deep Turks Island Passage and have never been joined (Lighty et al., 1982;Fairbanks, 1989; Welch et al., 2004). Since this last glacial maximum, rising sealevels have fragmented the banks into the present islands and islets (Morgan, 1989).The current islands are elevated ridges formed from wind-blown oolitic depositsof the larger glacially-emergent islands (Sealey, 2006; Ricklefs and Bermingham,2007), and the highest elevation is about 50 m above sea level. Total land area of theislands is approximately 500 km2. Vegetation consists of low coastal scrub on mostislands, interspersed with coppice on larger islands and remnant tropical dry foreston North Caicos, where most moisture occurs (Iverson, 1979; Correll and Correll,1982; Manco, 2008a). Extensive sea grass beds occur on the bank side of the majorislands of the Caicos bank, dominated by turtle grass (Thalassia testudinum) andmanatee grass (Syringodium filiforme), and provide refuge and forage for younggreen turtles (Manco, 2008a). Annual rainfall averages 750 mm on the Caicos bank,

380 R.G. Reynolds

with most rain occurring September to December. Considerable local variation inrainfall occurs, however, with the northwestern end of the TCI receiving morerainfall than the drier southwestern end (Sealey, 2006).

Human history

A valuable reference exists on the history of the TCI (Mills, 2008a), but I willprovide a brief account in the context of conservation issues. Humans have occupiedthe archipelago for at least 1300 years, beginning with the Taíno arriving fromHispaniola around 705 C.E. (Newsom and Wing, 2004; Carlson and Keegan,2006; Marvel, 2008). These peoples evolved culturally into the Lucayans, whomay have numbered as many as 10,000 in the TCI prior to European contact(Marvel, 2008). Amerindians likely had a significant impact on the native wildlifeof the TCI (Newsom and Wing, 2004; Franz and Franz, 2009). They practiced“swidden agriculture”, which involved removing native vegetation to grow foodplants, and also made extensive use of the local terrestrial fauna, including reptiles(table 1). A large species of tortoise (Chelonoidis sp.) has been found in middens onGrand Turk (“Coralie” site) (Carlson, 1999; Newsom and Wing, 2004) and MiddleCaicos (Franz and Franz, 2009); which was likely driven extinct by hunting andegg gathering. The boids Epicrates chrysogaster and Tropidophis greenwayi havebeen found in archeological sites on Grand Turk and Middle Caicos, respectively(Newsom and Wing, 2004). Anolis scriptus has been found at Middle Caicos sitesand Leiocephalus psammodromus has been found at the Grand Turk site (Newsomand Wing, 2004). Iguanas (Cyclura) and turtles (Chelonia, Caretta, and Trachemys)were also used as major food sources and are represented at sites on Grand Turk and

Table 1. Terrestrial and marine reptiles represented in archeological sites on Middle Caicos (MC-6,MC-12, and MC-32; Wing and Scudder, 1983) and Grand Turk (Coralie; Carlson, 1999). Counts aregiven as the Minimum Number of Individuals (MNI), which indicates the approximate number ofindividuals encountered at each site. Question marks indicate that individuals have been found thoughMNI for each site is unknown. After Newsom and Wing (2004).

Taxon MC-6 MC-12 MC-32 Coralie

Anolis sp. 1 0 0 0Cyclura sp. 11 2 1 1Cyclura carinata 0 0 0 386Iguanidae 0 1 2 0Leiocephalus psammodromus 0 0 0 12Epicrates chrysogaster 0 0 0 2Tropidophis sp. 0 0 1 0Trachemys sp. 0 0 1 0Chelonoidis sp. ? ? ? 18Caretta caretta 0 0 1 1Chelonia mydas 0 0 0 50Cheloniidae 4 1 0 0

Amphibians and reptiles in the Turks and Caicos Islands 381

Middle Caicos (Seidel, 1996; Lee and Ross, 2001; IUCN Iguana Specialist Group,2003; Newsom and Wing, 2004; Campbell, 2006).

After European contact, human population density dropped in the archipelago,and Ponce de Leon allegedly noted in 1513 that he did not see evidence of peopleduring a passage through the islands (Marvel, 2008). European settlement did notbegin in earnest until the early 17th century, when salt-raking became a profitablecolonial enterprise. Most of this activity was centered on the Turks bank, wherethe population was probably only a few dozen to a few hundred per island, withGrand Turk and Salt Cay having the most inhabitants. In 1781, 921 inhabitants wererecorded on the Turks bank, though most of these were seasonal migrants (Sadler,2008a). At this time land on the Caicos bank was being granted to Loyalists afterthe American War of Independence and cleared for cotton plantations. Valuablehardwoods were removed during this period, greatly altering hydrological cyclesand microclimates on the islands. By 1843 the population of the TCI was estimatedto be 2495 (Sadler, 2008b). A second wave of agriculture came in 1890, whenthousands of hectares were cleared on both banks for sisal (Agave sisalana)production. Population size probably remained below 6000 until the 1960s, whentourism sparked a wave of immigration from Jamaica and Hispaniola that continuestoday. By 1996 the population reached 15,000, doubling to 30,600 by 2005 andincreasing at roughly 3.5% per year (Jones, 2008). However, the population islikely higher than the current (2008) estimated census of 36,605, as waves ofundocumented immigrants have arrived from Haiti in recent years, including severalhundred after the catastrophic Haitian earthquake of 2010 (TCI Department ofPlanning and Statistics, 2010). Well over half of the population lives on the island ofProvidenciales, which has seen a huge increase in development since the completionof the new international airport and several resorts in the early 1980s. Tourism andbanking are the two major industries, and over 265,000 tourists arrive annually (TCIDepartment of Planning and Statistics, 2010). Many foreigners are buying homesand property in the islands, and development is occurring at a rapid pace. As wavesof tourists come to enjoy the wildlife of the islands, there is some desire amonglocals to mitigate consequences of development on wildlife; however, this is notoften accomplished in practice.

Conservation legislation and infrastructure

A National Parks committee was established in the TCI in 1969 and prepared alist of areas to be protected as Historical Sites, Sanctuaries, and Recreational Areas.These areas were surveyed and augmented by Ray and Sprunt (1971), and a NationalParks Ordinance was first enacted in 1975 to provide the framework for legal pro-tection of areas (Homer, 2000). Protection in practice was not established until thecreation of the National Parks Order in 1992, which supported four classificationsfor protected areas in the TCI, from most to least restricted: Sanctuaries, Nature Re-serves, National Parks, and Areas of Historical Interest (table 2). The Departmentof Environment and Coastal Resources (DECR) was established in 1995 to oversee

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384 R.G. Reynolds

protection of these areas, and today serves as the authority for reserves and biolog-ical resources, employing ∼180 individuals. Enforcement of environmental regula-tions is carried out by ∼38 patrolling conservation officers, largely active aroundProvidenciales, Grand Turk, and South Caicos and operating with a limited bud-get of around $50,000 US/year for gas, vehicles, and supplies (DECR, 2008). TheDECR also works with the TCI National Trust, a small non-governmental organi-zation founded in 1992 and tasked with preservation and promotion of the natural,cultural and historical heritage of the TCI (TCI National Trust, 2010). The Trust pre-serves a number of heritage sites independent of the national park system, some ofwhich are significant for local reptile populations. Wades Green Plantation on NorthCaicos protects remnants of tropical dry forest which supports large populations ofAnolis, Epicrates, Tropidophis, and Aristelliger species, as well as populations ofmost of the other native terrestrial reptile species (Reynolds and Manco, unpubl.).

The DECR has issued a series of ordinances and policies for protection ofwildlife (birds, flora) and fisheries (including marine turtles), and in 2001 theTCI Environment Charter was issued jointly by the UK and the TCI to establishguidelines and goals for environmental conservation and resource use. Priorities todate have included protection of marine fisheries, native vegetation, and protectedareas; though an endangered species bill including provisions for CITES is currentlyunder review.

The Native Herpetofauna

The terrestrial herpetofauna of the TCI is largely derived from Hispaniola, whichlies approximately 130 km to the south of the Caicos bank, a distributional patternthat follows ocean currents and hurricane tracks which might facilitate oceandispersal (Schwartz, 1967, 1968; Schwartz and Crombie, 1975; Buden, 1981;Tolson, 1987; Pregill, 1992; Franz et al., 1996; Hower and Hedges, 2003; Hedges,2006). At least six endemic reptile species occur on the islands, though no endemicreptile genera are present. Two species, Cyclura carinata and Anolis scriptus,share conspecifics on Mayaguana (Anolis) or its satellite Booby Cay (Anolis,Cyclura). Epicrates c. chrysogaster is a subspecies of the southern Bahamas boa,with other subspecies occurring on the Inaguas and the Crooked-Acklins islands.Most of the native reptilian fauna is widespread throughout the TCI, though recentextirpations have narrowed the range of some (table 3). The lizards Leiocephaluspsammodromus and Sphaerodactylus show a genetic split between banks, withsubspecies of the former occurring on each bank and widely divergent speciesof the later isolated on each bank (Reynolds et al., unpubl.). A few single-islandendemic subspecies have been described for both Leiocephalus (Schwartz, 1967)and Tropidophis (Schwartz, 1963), though it remains to be seen whether thesesubspecific epithets are confirmed by molecular analysis. A single widespreadspecies, Mabuya cf. sloanii, ranges well beyond the TCI throughout the Greaterand northern Lesser Antilles (Hedges, 2010).

Amphibians and reptiles in the Turks and Caicos Islands 385Ta

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386 R.G. Reynolds

Information contained in the species accounts is compiled from a variety ofsources, especially Schwartz and Henderson (1991) and Henderson and Powell(2009), and relies heavily on my own and others’ work in this region. In orderto provide taxonomic consistency, I have chosen to use scientific nomenclatureconsistent with other recent published works in the field of Caribbean herpetology(e.g., Henderson and Powell, 2009), with standard names largely consistent withHedges (2010). There are two islands named “Long Cay”; one on the Caicos bank0.75 km southwest of South Caicos and one on the Turks bank 2.85 km southeastof Grand Turk (fig.1); I therefore specify in parenthesis the appropriate bank.Major islands in this region are East Caicos, Grand Turk, North Caicos, MiddleCaicos, Providenciales, South Caicos, and West Caicos. Natural history informationpertinent to status and conservation of each species follows, and each accountincludes an assessment of the conservation status and main threats to the species.Data deficient indicates that little information on natural history, distribution, and/orconservation status is available for the species in the TCI.

Testudines

Cheloniidae. Caretta caretta (loggerhead turtle). Adults and juveniles are infre-quent locally, and at least one confirmed nesting event was recorded on Long Bay,Providenciales in May 2005 (Pardee, 2005; Richardson et al., 2009). This speciesis data deficient in the TCI. Chelonia mydas (green turtle). Adults and juvenilesare common in local waters. Small numbers of this species are nesting or poten-tially nesting on Big Sand Cay, Bush Cay, East Caicos, and Gibbs Cay (table 3;Godley et al., 2004; Richardson et al., 2009). Several nesting populations appear tohave been recently extirpated (table 3; Godley et al., 2004; Richardson et al., 2009).Eretmochelys imbricata (hawksbill turtle). Adults and juveniles are common in localwaters, with greater nesting frequency than green or loggerhead turtles (Richardsonet al., 2009). Small numbers of this species are nesting or potentially nesting on BigAmbergris Cay, Big Sand Cay, Bush Cay, East Caicos, Fish Cay, Middle Caicos,and North Caicos (table 3; Godley et al., 2004; Richardson et al., 2009). Severalnesting populations appear to have been recently extirpated (table 3; Godley et al.,2004; Richardson et al., 2009).

Sauria

Sphaerodactylidae. Aristelliger hechti (Caicos or Hecht’s barking gecko). Thisspecies is restricted to the Caicos bank (table 3; Bauer and Russell, 1993). Theyare common in xeric scrub on Big Ambergris Cay and in tropical dry forest onNorth Caicos. Populations possibly occur on Middle Caicos, though this species hasnot been recorded during many herpetological surveys of that island. Individualsare frequently found underneath rock piles during the day or on vegetation, rockwalls, and rocks at night; and are also associated with artificial cover (Schwartzand Crombie, 1975; Bauer and Russell, 1993). The biology of this species ispoorly understood, though the fragile skin of A. hechti has been discussed by

Amphibians and reptiles in the Turks and Caicos Islands 387

several authors (Greene, 1988; Bauer and Russell, 1992, 1993; Bauer et al., 1992).The range appears to be locally restricted within islands. This species is likely aconservation concern due to its restricted range, the presence of feral mammals onsome islands within its range, low density, and widespread introduction of the geckoHemidactylus mabouia, which is a possible competitor (Reynolds and Niemiller,2009, 2010a).

Sphaerodactylus caicosensis (Caicos banded dwarf gecko). This species is re-stricted to the Caicos bank on all major islands and some small cays and rock islandswith some vegetation (table 3). They are common in xeric habitat behind beaches aswell as in more mesic tropical dry forest on North Caicos. Individuals are frequentlyfound under rocks and litter along beaches and roads and commonly encountered onrock walls, low vegetation, and on stones on the ground at night. Many populationsare edificarian and common near human habitation and in areas with high densi-ties of feral mammals. This species does not appear to be a conservation concern.Sphaerodactylus underwoodi (Turks island dwarf gecko). This species is restrictedto the Turks bank (table 3). Individuals are edificarian and xerophillic, common un-derneath rocks and litter in areas behind beaches, and common in urban lots withsome vegetation on Grand Turk. This species does not appear to be a conservationconcern.

Iguanidae. Cyclura carinata (Turks and Caicos rock iguana). This species isdistributed on many small cays and rock islands with some vegetation on both theTurks and Caicos Banks (table 3; Welch et al., 2004). Extirpated on most of the largeislands in the TCI, this species survives on small cays that lack introduced vertebratepredators. The largest remaining populations occur on Big and Little AmbergrisCays on the southeastern edge of the Caicos bank. Individuals of this species havebeen translocated to several outlying cays to ensure their long-term survival in theevent of the introduction of cats and dogs to the Ambergris Cays (Gerber, 1996,2004, 2007).

Leiocephalidae. Leiocephalus psammodromus (Turks and Caicos curly-tailedlizard). This species occurs on most major islands larger than 0.42 km2 on theTurks and Caicos Banks (table 3; Reynolds and Fitzpatrick, unpubl.). They are likelyextirpated from some islands with a long history of European habitation and feralpopulations of vertebrate predators and herbivores, as evidenced by their presence inmiddens on Grand Turk (Newsom and Wing, 2004). Some smaller cays (<1.0 km2)

on the Caicos bank do not have populations of this species (e.g., Mangrove Cay,Middleton Cay). Six subspecies are currently recognized (Schwartz, 1967), thoughperhaps none are warranted given current molecular data (Reynolds and Fitzpatrick,unpubl.). Conspicuous where they occur, individuals occupy a variety of xerichabitats with access to both full sun and shade (Smith, 1994, 1995), as well asedificarian irrigated situations. Curly-tailed lizards are extremely susceptible topredation by feral mammals (Iverson, 1978; Smith and Iverson, 1993), and havelow clutch sizes (Smith and Iverson, 1993). More information is needed to assess

388 R.G. Reynolds

the conservation status of this species, but it is apparently extirpated from someislands while still common on many others.

Polychrotidae. Anolis scriptus scriptus (Turks and Caicos anole). This speciesoccurs on all major islands and most small cays and rocks with some vegetation(table 3). Populations in the TCI are considered a subspecies of the southernBahamas anole complex (Rand, 1962). Individuals are common in most habitattypes, including edificarian and xeric to relatively mesic zones, as well as irrigatedareas. This species does not appear to be a conservation concern.

Scincidae. Mabuya cf. sloanii (Antillean slipperyback or skink). The specificepithet sloanii is likely to change for populations in the Turks and Caicos tomore accurately reflect actual evolutionary relationships (Conn and Hedges, pers.comm.). Populations are present on all major islands and some smaller islandsand cays (table 3), though this species is data deficient in the TCI. This species isencountered infrequently, though it might be locally common. Individuals are oftenencountered in leaf litter or underneath flat rocks. This species does not appear tobe a conservation concern.

Serpentes

Boidae. Epicrates chrysogaster chrysogaster (Turks island boa). This speciesoccurs on some islands on the Caicos bank, and was recently rediscovered on GibbsCay in the Turks Cays (Reynolds and Niemiller, 2008, 2010b, unpubl.; Reynolds etal., in review). Populations of this species could possibly occur on Long Cay (Turksbank), East Cay (Buden, 1975), and West Caicos (Reynolds and Gerber, unpubl.).The original holotype is from Grand Turk, though until recently no individuals werereported from the Turks bank since the collection of the type specimen (ANSP10322; Cope, 1871) (Tolson and Henderson, 1993). Populations have apparentlybeen extirpated from Grand Turk, and possibly from Salt Cay and some of theCaicos Cays (small islands between Providenciales and North Caicos). Individualsare uncommon to rare on Providenciales and infrequent on other islands exceptfor Big Ambergris Cay, where this species is abundant (Reynolds et al., unpubl.).Diurnal refugia include large limestone rocks, logs, subterranean interstices, humandebris, and palm litter (Sheplan and Schwartz, 1974; Buden, 1975; Reynolds andGerber, unpubl.). This species is CITES Appendix II (2009) listed and is a majorconservation concern due to loss of habitat, removal for the pet trade, directpersecution, and introduced mammalian predators.

Tropidophiidae. Tropidophis greenwayi (Caicos dwarf boa). This species is re-stricted to major islands on the Caicos bank, as well as Long Cay (Caicos bank) andMiddleton Cay (Iverson, 1986). There are currently two recognized subspecies: T.g. greenwayi distributed on Big Ambergris Cay and T. g. lanthanus distributed else-where on the Caicos bank (Iverson, 1986; Schwartz and Henderson, 1991; Hedges,2002; Henderson and Powell, 2009). It is possible that the subspecific epithets will

Amphibians and reptiles in the Turks and Caicos Islands 389

collapse following further analysis (J. Iverson, in litt.). The Big Ambergris Caypopulation was thought to be extirpated, as the only known specimens (n = 2) werecollected as types by Barbour and Shreve in 1936. Reynolds et al. (2010) foundan additional individual on 20 March 2009, indicating that this subspecies mightpersist, though it is possible that this specimen represents an individual introducedduring movement of construction equipment from Providenciales. T. g. greenwayiis likely xerophilic, while T. g. lanthanus is mesophilic, frequently occurring nearsources of standing water or ground water such as old wells, ephemeral ponds, andthick forest. Individuals are occasionally found in xeric situations as well, such asrocky well-drained hillsides on Middle Caicos (Reynolds and Deal, unpubl.), andopen cactus shrubland on South Caicos (Iverson, 1986). This species is a seriousconservation concern, as it has been collected extensively for the pet trade and islikely susceptible to introduced mammalian predators (Iverson, 1986). Addition-ally, the subspecies T. g. greenwayi faces an uncertain future, as it was likely neverabundant and its range is apparently restricted to Big Ambergris Cay, which is cur-rently being heavily developed.

Typhlopidae. Typhlops cf. platycephalus (Caicos blindsnake, sensu B. Hedges,in litt.). This species is restricted to the Caicos bank, though it is data deficient.Individuals from the Caicos Islands, considered to be T. richardii in Schwartz andHenderson (1991), were reassigned to T. platycephalus by Hedges and Thomas(1991) and are currently being assigned to a unique species based on DNA sequencedata to more accurately reflect evolutionary relationships (B. Hedges, in litt.). Theseindividuals were not recognized as T. platycephalus in Henderson and Powell(2009), though this species does appear to be derived from the Puerto Rican bank(B. Hedges, in litt.). Individuals are xerophilic, found underneath rocks in sandy soilin low scrub (Big Ambergris Cay: Reynolds, 2010) and tropical dry forest (NorthCaicos), and are occasionally found on the surface at night. Conservation status ofthis species is unknown; however, it may become a conservation concern if foundto be an endemic species and if competition exists with the introduced Brahminyblind snake (Rhamphotyphlops braminus) (Reynolds and Niemiller, 2010a).

Introduced Amphibians and Reptiles

The following is a complete list of all historical and current records of introducedamphibians and reptiles in the TCI. The majority of this information comesfrom Manco (2008b) and Reynolds and Niemiller (2010a), with some additionalobservations added. Populations are considered established if juveniles and/orevidence of reproduction (i.e., hatched eggs) have been observed.

Amphibia

Eleutherodactylidae. Eleutherodactylus planirostris (greenhouse frog). Popula-tions are established on some islands on the Caicos bank (table 4), though indi-

390 R.G. Reynolds

Tabl

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Amphibians and reptiles in the Turks and Caicos Islands 391

viduals are likely to be found on additional islands. This species can be common,especially during the wet season. Individuals frequent both landscaped areas as wellas rocky areas within tropical dry forest on North Caicos; their terrestrial mode ofreproduction undoubtedly facilitates their colonization and range expansion.

Hylidae. Osteopilus septentrionalis (Cuban treefrog). Populations are establishedon some islands on the Caicos bank (table 4), though individuals are likely tobe found on additional islands. This species is abundant during rainy autumnnights on North Caicos, where individuals frequent roadways. Individuals are verycommon in uncovered wells and water tanks. This species is a documented prey itemfor Tropidophis greenwayi on Providenciales (Manco, 2006). Adult T. greenwayihave been observed to be especially common inside the walls of old stone wellswhere metamorphic O. septentrionalis emerge from the water below (Tolson andHenderson, 1993; Reynolds and Niemiller, 2010a).

Testudines

Emydidae. Trachemys scripta elegans (red-eared slider). This species is infre-quent and not established in the TCI. Individuals have been released by residentsinto artificial freshwater ponds at the golf course on the east end of Providen-ciales. Three have been removed by conservation officials and several others havebeen sighted. No hatchlings or juveniles have been observed. Trachemys stej-negeri malonei (Antillean slider). One individual was collected from Pine Cay in1975 (W. Auffenberg; UF 49,423), though others were seen in a freshwater pondon Pine Cay in December 1997 (Seidel, 1988, 1996; Lee and Ross, 2001). Thecurrent status of this population is unknown, though to my knowledge individ-uals have not been reported from Pine Cay since Lee and Ross’s last observa-tion in 1997. This population is believed to represent a human-mediated intro-duction (Seidel, 1988, 1996) from Great Inagua. However, archeological remains(shell fragments) over 700 years old have been reported from Middle Caicosand Grand Turk purportedly from the genus Trachemys (Carlson, 1994; Seidel,1996; Lee and Ross, 2001; Newsom and Wing, 2004), though Keegan and Carl-son (2008) did not consider these to represent this genus. As slider turtles werecommonly used throughout the Antilles as a food source by the Taíno (Keegan andCarlson, 2008), it is possible that repeated historic introductions to the TCI oc-curred.

Sauria

Gekkonidae. Hemidactylus mabouia (wood slave). Populations are establishedon several islands in the TCI (table 4), though individuals are likely to be foundon additional islands (Minton and Minton, 1975; Reynolds and Niemiller, 2010a).Nests are reported from Salt Cay and hatchlings have been reported from all sixislands (Reynolds and Niemiller, 2009, 2010a). The influence of this species on theendemic gecko Aristelliger hechti is unknown.

392 R.G. Reynolds

Sphaerodactylidae. Sphaerodactylus mariguanae (Mayaguana dwarf gecko orsphaero). This species was originally reported from Grand Turk (Schwartz, 1968;Schwartz and Henderson, 1991), though apparently has not been collected recently.Schwartz (1968) collected a series of 40 individuals from Cockburn Town, GrandTurk and found them to be distinct from S. underwoodi in size, color pattern, andscalation. It has been suggested that these records might have been a misidentifica-tion; however, this seems unlikely given the large numbers examined and Schwartz’scomparison with other populations of S. mariguanae from Booby Cay (Mayaguana)and Mayaguana (∼185 km NW of Grand Turk). Schwartz speculated that GrandTurk was within the natural range of this species, as several other reptiles are foundon both the Turks bank and Booby Cay (Cyclura carinata, Anolis scriptus, Mabuyasloanii) and postulated that S. mariguanae would be found on the Caicos bank (ithasn’t). If Grand Turk S. mariguanae are within the natural range of this species,it is curious that they have not been seen in recent years and that they were neverrecorded from other islands on the Turks bank. Schwartz (1968) also speculatedthat Grand Turk individuals may be an introduced population, and the most likelyscenario seems to be an introduction followed by what appears to be an extirpationof S. mariguanae on Grand Turk (Reynolds and Niemiller, 2010a).

Iguanidae. Iguana iguana (green iguana). This species has been recorded fromboth the Turks and Caicos banks, though it is not established on either. Individualslikely represent escaped or released pets, and no hatchlings or juveniles have beenreported. At least one individual was captured and removed from Providenciales,though others are occasionally seen on both Providenciales and Grand Turk.

Polychrotidae. Anolis equestris (knight anole). Adults of this species have onlybeen documented from Providenciales and it is not currently established. At leastsix individuals have been recorded from the landscaped grounds of a resort in theGrace Bay area of Providenciales and likely represent escaped or released pets. Fourindividuals were removed and placed in captivity. No hatchlings or juveniles havebeen found.

Serpentes

Colubridae. Pantherophis [Elaphe] guttatus (corn snake). Three individuals ofthis species have been removed from Grand Turk, one of which was a road-killedspecimen. One individual laid a clutch of inviable eggs after capture (Reynoldsand Niemiller 2010a). This species is likely represented by either escaped petsor individuals (possibly eggs) in live trees brought from Florida (Reynolds andNiemiller, 2010a). Two of the three specimens were found near the new cruise shipterminal, while the location of the third is unclear.

Typhlopidae. Rhamphotyphlops braminus (brahminy blind snake). This speciesis established and common on Grand Turk and Providenciales, likely having arrived

Amphibians and reptiles in the Turks and Caicos Islands 393

as a stowaway in the soil of potted plants. The influence of this species on the nativeTyphlops is unknown.

Other non-native species

Occasional pets are released or escape and survive for a period of time in the wild.A savanna monitor lizard (Varanus exanthematicus) was occasionally spotted onProvidenciales until 2004 and was a known release of a pet by a resident of thatisland (B. Riggs and B.N. Manco, in litt.). Cuban brown anoles (Anolis sagrei) arenot yet reported from the TCI, though the eventual arrival of this species seems adistinct possibility.

Species of Special Concern

Sea turtles

Four species of sea turtles occur in TCI waters; however, only three are knownto actively nest on the islands. The fourth, Dermochelys coriacea (leatherbackturtle), is occasionally seen offshore but not known to nest within the archipelago.Loggerhead turtles formerly nested in the archipelago, though nesting is nowextremely rare and sporadic (Pardee, 2005; Richardson et al., 2009). Hawksbilland green turtles continue to nest in low density on the more remote cays inthe archipelago, with hawksbill nesting occurring from at least July to Januaryand Green turtle nesting in September (Richardson et al., 2009). Though the TCIappear to support low nesting activity, the sea grass beds and fringing reefs likelyprovide regionally significant foraging areas for young green and hawksbill turtles(Richardson et al., 2009).

Harvest of green turtles remains legal in TCI waters (Richardson et al., 2006;Broderick et al., 2006), though annual take is largely unknown due to logisticalchallenges for monitoring or enforcement. Godley et al. (2004) estimated that 240-1130 individuals were harvested annually in local waters in directed fishery, plusan additional 100 individuals that were captured annually as incidental catch. Rudd(2003) estimated that the harvest was several hundred annually, whereas Richardsonet al. (2009) estimated that annual harvest of green turtles may be around 210.Juveniles are common in the extensive sea grass beds and tidal creeks on the banksides of the major Caicos Islands. Richardson et al. (2009) document fibropapillomasymptoms in a juvenile green turtle from the Caicos bank, indicating that thispopulation might be affected by this disease. This species is listed as endangered bythe IUCN (Seminoff, 2004), and current and historical take have likely significantlydecreased nesting populations in the Turks and Caicos archipelago (Richardson etal., 2009).

Harvest of hawksbill turtles also remains legal in TCI waters (Richardson etal., 2006; Broderick et al., 2006), though annual take is largely unknown dudue to logistical challenges for monitoring or enforcement. Godley et al. (2004)

394 R.G. Reynolds

estimated that between 180 and 900 individuals are harvested annually in localwaters in directed fishery, plus an additional 100 individuals captured annuallyas incidental catch. Rudd (2003) estimated that the harvest was several hundredannually. Juveniles are common on patch reefs on the Turks and Caicos banks, andexhibit a higher growth rate than other Caribbean populations (Richardson et al.,2009). Current and historical take of this species have likely significantly decreasednesting populations in the Turks and Caicos archipelago (Richardson et al., 2009).

Though turtle soup has recently been removed from menus on the main touristislands, turtles continue to be harvested for local consumption. Current fishingregulations (Fisheries Protection Ordinance, 1998) allow harvesting of one turtleper person per week, weighing >20 pounds or measuring >50.8 cm carapacelength to be legally taken in TCI waters (Broderick et al., 2006; Richardson, 2006).Under these regulations, the harvesting of eggs, nesting females, and juveniles isprohibited, though previous legislation (TCI Fisheries Protection Regulations, 1976)failed to prevent the estimated harvest of 8000-10,000 eggs per year (Fletemeyer,1984) due to lack of enforcement (Richardson et al., 2009). Eggs are still harvestedand either consumed or sold, though probably in much lower numbers than in the1970s and 1980s (Richardson et al., 2009). Penalties for illegal take of turtles haveincreased recently, with an individual receiving a maximum fine of US$7000 in2008 (P. Dickerson, pers. comm.).

Combined with the development of coastal areas, fibropapilloma, and legalharvest, sea turtles in the TCI face an uncertain future. To ensure that nestingactivity is not completely eliminated from the islands, increased surveillance ofnesting beaches is encouraged. Additional research on sea turtles is ongoing in thearchipelago (Marine Turtle Research Group, University of Exeter, TCI Departmentof Environment and Coastal Resources, and Turtles in the UK Overseas Territories),which should help to inform oversight of the turtle fishery and contribute toregulations that ensure that the ongoing take does not impact local or regionalpopulations.

Cyclura carinata

The genus Cyclura is notable for both its high level of endemism, with most ofthe 16 species restricted to one or a few islands in the Caribbean (Etheridge, 1982;Alberts, 2000; Welch et al., 2004), as well as the rapid decline of most species inthe genus due to human activities (Iverson, 1978; Alberts et al., 2004). The speciesCyclura carinata, first described by Harlan (1824), is restricted to the Turks andCaicos banks, as well as Booby Cay, a satellite of Mayaguana located approximately63 km northwest of the Caicos bank. This Booby Cay population was formerlyreferred to as the subspecies C. c. bartschi, though subsequent molecular analysishas revealed that this population is not unique enough to warrant subspecific status(Bryan et al., 2007) and was probably introduced from the Turks bank. Welch et al.(2004) found evidence for recent colonization of the Turks bank from populationsof C. carinata on the eastern Caicos bank. Iguanas will not willingly enter the

Amphibians and reptiles in the Turks and Caicos Islands 395

water, though occasionally individuals may be swept out to sea, and at least oneindividual has been found alive drifting on the Caicos bank several kilometers fromthe nearest population (Gerber, pers. comm.). Though not mentioned by Welch etal. (2004), one possible means of conveyance to the Turks bank is human-assisteddispersal. It is known that indigenous, and possibly European, people used iguanasfor food, and hence may have facilitated transport of individuals to other islands inthe archipelago (IUCN Iguana Specialist Group, 2003).

Iguanas formerly occupied most islands in the TCI, though recently they havebeen extirpated from most of the large islands and are now restricted to small caysand rock islands, a total area of only 13 km2 (Gerber and Iverson, 2000; Gerber,2004) and less than 5% of their former range (Hudson and Alberts, 2004). The majorthreats faced by iguanas include feral dogs and cats, which prey on young iguanasand destroy burrows (Iverson, 1978). Iverson (1978) documented the systematicnear extirpation of iguanas from Pine Cay on the Caicos bank in a matter of years.Iguanas have since returned to Pine Cay (Smith, 1992a) and now occur in extremelylow density on that island, though the population has never recovered and likelywill not recover until feral mammals are removed. Iguanas were also extirpatedfrom Long Cay on the Caicos bank due to predation by feral cats (Mitchell et al.,2002). A cat-eradication campaign was begun in 1999 using targeted poison-baits,and 800 iguanas were translocated from Big Ambergris Cay following removal ofthe invasive predator (Mitchell et al., 2002). The population of iguanas on LongCay has successfully established since re-introduction, and juvenile recruitment isoccurring (Reynolds, pers. obs., 2008).

Currently, the largest remaining population of iguanas occurs on Big and LittleAmbergris Cays, where an estimated 15,000-20,000 individuals occurred prior tothe start of development of Big Ambergris Cay (Gerber, 2004). Though LittleAmbergris Cay remains protected at the moment by the TCI National Trust,Big Ambergris Cay is currently being heavily developed, and iguanas there arethreatened by habitat loss due to blasting and building as well as vehicle strikes.This population has already declined, and current ongoing studies are determiningwhether individuals can be moved from areas of the island slated to be developedin the immediate future to areas that will remain undeveloped in the near future(Gerber, 1998, 2010). Iguanas have also been successfully translocated from thissource population on Big Ambergris Cay to outlying islands that are predator anddevelopment free (Alberts, 2001; Alberts and Gerber, 2004; Gerber, 2007). Thesepopulations are being monitored annually for juvenile recruitment and survival andare intended to serve as buffers against catastrophic loss of other populations suchas the one on Big Ambergris Cay. Little Ambergris Cay also remains under threatof being bought and developed by foreign companies, though it is hoped that theseentreaties will be resisted.

Iguanas in the TCI are subject to intensive monitoring and research, initiatedin the mid-1970s (Iverson) and continuing today (Glenn Gerber, San Diego ZooInstitute for Conservation Research). A small population of iguanas on Little Water

396 R.G. Reynolds

Cay (within Princess Alexandria National Park) serves as an extremely populartourist destination, and the TCI National Trust employs guides and guards tointroduce visitors to the plight of the endangered iguanas. Little Water Cay is nowconnected to Water and Pine Cays by a strip of sand, which allows feral mammalsfrom these islands to invade and potentially impact the population of iguanas onLittle Water Cay. Iguana awareness campaigns have shown great success, and anatmosphere of stewardship for this species exists in the Turks and Caicos, thoughdevelopers and pet owners often overlook the impacts of their respective activities.It is hoped that intensive management and mitigation may allow the persistence ofrock iguanas, though additional predator removal campaigns would greatly benefitthis species.

Epicrates chrysogaster chrysogaster

This endemic subspecies was first described by Cope (1871) based on an individualfrom “Turk’s Island”, which has been taken to refer to what is now Grand Turk.Since this description E. c. chrysogaster has not been recorded on Grand Turk, andwas thought to have been extirpated from the Turks bank. However, Reynolds andNiemiller (2010b) and Reynolds et al. (unpubl.) report four individuals discoveredon Gibbs Cay, which is a very small island about 1.5 km SE of Grand Turk,representing the first report of this species on the Turks bank since the originaldescription in 1871, and the only report from an island on this bank other than GrandTurk. In addition, these individuals are fixed at a unique mitochondrial haplotypethat is ∼1% divergent from Caicos bank haplotypes (Reynolds et al., unpubl.). It hasbeen suggested that given the inconsistency of map labeling of the region until the1970s (Buden, 1975), the holotype locality was in fact a different island than GrandTurk. However, it now seems more likely that this species was in fact extirpatedfrom Grand Turk and remnants of the Turks bank lineage remain on at least one ofthe Turks cays.

Genetic analysis for this species (Reynolds et al., unpubl.) has shown surprisinglyshallow overall genetic divergence among island populations of E. chrysogaster.The largest divergence (1%) in a mitochondrial gene is between haplotypes onGibbs Cay and Big Ambergris Cay, while two nuclear genes show almost nodivergence. This is in sharp contrast to other squamates in the region, whichhave been described as unique subspecies (Leiocephalus, Tropidophis) or species(Sphaerodactylus) and show large genetic divergence (Reynolds and Koneczny,unpubl.; Reynolds et al., unpubl.). This shallow genetic divergence is hypothesizedto be due to lack of historical population structure when the Turks and Caicos Bankswere each emergent during the last glaciation (Reynolds et al., unpubl.).

Turks Island boas are susceptible to predation by feral mammals (Reynolds, pers.observ.), as well as habitat loss, direct persecution, and vehicle strikes. It is likelythat this species has already been extirpated from Grand Turk and South Caicos,which both have a long history of European occupation (Mills, 2008a; Sadler,2008c). As development continues at an astounding rate in the region, it is likely

Amphibians and reptiles in the Turks and Caicos Islands 397

that more populations will be extirpated. Boas are becoming difficult to find onProvidenciales, and the future survival of that population seems bleak. The densestpopulation of boas in the TCI, and one of the densest population of Epicratesanywhere, currently occurs on Big Ambergris Cay, where a long-term monitoringeffort has recently been started (Reynolds and Gerber, unpubl.). This private islandis being heavily developed into vacation homes, and conservation officials andresearchers are working closely with developers and owners to implement effectivemitigation measures.

Epicrates chrysogaster had been a long-neglected species in terms of research andconservation assessment (likely due to a perceived low abundance), though thank-fully this has changed in the last five years. Now focused research programs includ-ing natural history, ecology, and genetic analysis have been begun (G. Reynolds andB. Fitzpatrick, University of Tennessee; G. Gerber, San Diego Zoo Institute for Con-servation Research) and are leading to more informed conservation decisions. Thepublic is also becoming aware of these snakes through the use of media outlets suchas local magazines, brochures, interpretive exhibits, and television appearances. Itis hoped that this new wave of enthusiasm will serve to ensure that sensitive popu-lations, such as those on Providenciales and Big Ambergris Cay can be protected insitu.

Discussion

The herpetofauna of the TCI consists of 23 species; 13 native (57%) and 10introduced (43%). Of the native species, two are critically endangered, two areendangered, and the rest have not been evaluated (IUCN, 2010). Reptiles in theregion are threatened by loss of habitat, feral predators, unsustainable exploitation(sea turtles), habitat degradation by semi-feral herbivores, and direct persecution(i.e., killing of snakes) (Iverson, 1978; Smith and Iverson, 1993; Gerber, 2004;Richardson et al., 2009; Reynolds et al., unpubl.). Invasive or introduced speciesalso threaten the native herpetofauna, though the impact of introduced amphibiansand reptiles is not yet known (Reynolds and Niemiller, 2010a). Many islands inthe Turks and Caicos remain uninhabitable by iguanas due to feral cats and dogs,while boas (Epicrates) appear to be extirpated from some islands such as GrandTurk and are likely declining on Providenciales (Reynolds et al., unpubl.). Curly-tailed lizards are apparently extirpated from South Caicos, Grand Turk, Salt Cay,and Cotton Cay, likely due to predation by cats (Iverson, 1978; Smith and Iverson,1993) and loss of habitat due to hundreds of years of human activities and thepersistence of semi-feral populations of donkeys, cows, goats, and horses. Theseherbivores have greatly altered the vegetation of many of the islands in the TCI.Indeed, the interiors of Salt Cay and South Caicos are dusty and sandy due to a lossof undergrowth. Sea turtle nesting is declining in the region (Richardson et al., 2009)due to harvesting of adults and loss of suitable beach nesting habitat to development.Some snake populations face a tenuous future, as some locals perceive them to be

398 R.G. Reynolds

venomous or malevolent spirits and many are either killed on sight, suffer mortalityfrom car strikes, or succumb to habitat loss or introduced predators (Reynolds etal., in review). Amazingly, a former government official granted a permit to reptiledealers to remove thousands of Tropidophis greenwayi from North Caicos for thepet trade, allegedly implying that it would be preferred if they removed all of thesnakes (Manco, pers. comm.)! Collection of iguanas and boas for the pet trade wascommon in the 1980s, and though it likely still occurs, it has probably slowed inrecent years due to increased port-of-entry surveillance in response to perceivedincreases in threats of terrorism and illegal immigration. Collection has the abilityto devastate local populations of reptiles (Webb et al., 2002), and hence enforcementmust remain strict.

One of the major obstacles to conservation of the herpetofauna of the region hasbeen a lack of knowledge about the biology, ecology, and distribution of species inthe archipelago. Thankfully, this has begun to change with increased field work andthe addition of individual species conservation programs. Turks and Caicos rockiguanas have likely been saved from extinction by the efforts of local authorities,NGOs, and conservation biologists from the U.S. and elsewhere (IUCN IguanaSpecialist Group, 2003; Gerber, 2004), initiated by a detailed study of their biologyand behavior (Iverson, 1979) and the witnessing of their near extirpation from PineCay (Iverson, 1978). Cyclura carinata is now the focus of a major awareness andconservation campaign, and has become one of the natural symbols of the TCI.

Worldwide declines in snake populations have recently been noted and representan increasing concern for conservationists (Mullin and Seigel, 2009; Reading et al.,2010). In 2007, a snake conservation program was initiated to increase knowledgeof the biology, ecology, and distribution of Epicrates chrysogaster, and to increaseawareness of the threats that this species faces (Reynolds, 2007; Reynolds and Deal,2010). An annual monitoring campaign, now in its third year, is taking place on BigAmbergris Cay (Reynolds and Gerber, unpubl.).

The future of some species of herpetofauna in the Turks and Caicos remainsuncertain, and important steps must be taken to ensure not only survival of thespecies, but also preservation of evolutionary significant units (ESUs — Ryder,1986; Waples, 1991; Dizon et al., 1992; Mortiz, 1994; Crandall et al., 2000) in theform of genetic diversity. The following strategies might contribute to alleviatingimpacts on the native herpetofauna of the region:

1. One of the greatest current threats to native herpetofauna in the Turks andCaicos is the introduction of feral predators such as cats and dogs to ecologicallynaïve island communities (Iverson, 1978). It is recommended that a great dealmore effort be exerted to prevent future introductions of cats and dogs, to spay andneuter pets or semi-feral animals, and to increase eradication campaigns on sensitiveislands. Cats can survive on islands without a permanent source of freshwater,and hence have been able to colonize many islands in the archipelago, which canhave serious consequences for local wildlife (Iverson, 1978). It is imperative thatcats be prevented from establishing on islands such as Big and Little Ambergris

Amphibians and reptiles in the Turks and Caicos Islands 399

Cays, which currently harbor critical populations of Cyclura carinata and Epicrateschrysogaster, as well as the endemic subspecies Tropidophis g. greenwayi andLeiocephalus psammodromus apocrinus. A cat recently colonized Big AmbergrisCay from a construction barge and survived for at least three months before beingcaptured and removed. Removal campaigns have been shown to restore islandsfor reintroduction of iguanas (Mitchell et al., 2002) and are becoming increasinglyeffective (Veitch and Clout, 2002).

2. Development continues at a rapid pace in the region, and mitigation of theconsequences of habitat loss must be a priority for conservation. Though BigAmbergris Cay is one of the most important refuges for herpetofauna in thearchipelago, it is currently under heavy development, with future plans indicatingthat nearly two thirds of the island will be developed. Developers engaged inthis project have thus far attempted to cooperate with conservation biologists, andproperty owners and managers seem to genuinely appreciate the need to minimizethe impact on the native wildlife. An environmental center was recently completedon the island, complete with naturalist staff, to allow residents and guests to learnabout the local flora and fauna. However, populations of Cyclura and Epicrates arelikely decreasing there due to blasting, clearing, and vehicle strikes. It is hoped thatas development slows and heavy construction trucks are replaced with golf carts,vehicle strikes will lessen, and owners will take an active role in stewardship oftheir island’s natural resources. Other developments should be required to take anactive role in mitigation, such as reducing nighttime illumination along beaches(which disorient hatchling marine turtles: Lohmann et al., 1997) and not developingecologically imperiled habitats such as the tropical dry forest on North Caicos,a large patch of which was recently clear-cut for a municipal development.

3. Several reserves, both terrestrial and marine, exist in the TCI (table 4;www.environment.tc; Carleton et al., 2006), some of which protect sensitive habitatand important reptile populations. These protected areas consist of four manage-ment units, established by the TCI National Park Ordinance (1989): National Parks,Nature Reserves, Sanctuaries, and Areas of Historic Interest (table 4). In addition,the Turks and Caicos National Trust, a local non-governmental organization, pro-tects a number of sites of historical interest on the Caicos bank. It is recommendedthat these areas be maintained in perpetuity, and that enforcement of regulationsthat protect these areas be increased. In addition, areas designated as Sanctuaries,which exist for “the avoidance of disturbance of the area by people” (Departmentof Environment and Coastal Resources, TCI, 2010), should be expanded to includeadditional important areas such as sea grass beds and fringing reefs which serve asjuvenile sea turtle rookeries. Additional recently proposed protected areas shouldincrease protection of native reptile populations (Carleton et al., 2006).

4. Introduction of non-native herpetofauna is an increasing issue in the Turksand Caicos (Reynolds and Niemiller, 2010a). Though it is not known whethercurrent introductions have negatively impacted the native herpetofauna (Reynoldsand Niemiller, 2010a), the number of recent introductions is alarming, and the

400 R.G. Reynolds

potential exists for the introduction of extremely harmful species. It has recentlybeen shown that the introduced Bufo (Rhinella) marinus (cane toad) in Jamaicais causing mortality to the endangered Epicrates subflavus (Jamaican boa) due toingestion of bufogenin toxin (Wilson et al., 2010). Importantly, Wilson et al. (2010)suggest that other island populations of Epicrates, which evolved in isolation fromthe genus Bufo and their associated bufadienolide glycosides, may be especiallysusceptible to these toxins.

5. Awareness campaigns targeted at increasing public appreciation for endemicherpetofauna have shown some success, especially for species such as the rockiguana (Cyclura carinata). Indeed, the motto for the TCI is “Beautiful by Nature”,and a sense of stewardship must be encouraged among citizens, visitors, andgovernment. Many visitors to the island of Providenciales pay for ecotourismtrips, which usually involve a stop at Little Water Cay to see the iguanas there.It is hoped that this type of interaction and the associated monetary benefits willencourage protection of endemic species (Knapp, 2004). Recent campaigns to raiseawareness of snakes in the islands face greater difficulties, including a generalfear of snakes that contributes to malicious killing. Introductions to the threatsfaced by the endemic snakes have been highlighted in magazine and newspaperarticles, brochures, interpretive exhibits at the National Environmental Centre onProvidenciales, television appearances, and use of captive animals in children’snature programs (Manco, 2006; Reynolds and Deal, 2010).

6. A great impediment to species conservation has been a lack of knowledge ofthe biology, ecology, population size, and conservation concerns for most speciesof native reptiles in the Turks and Caicos. This has recently begun to change, withlong-term intensive monitoring of Cyclura carinata (Gerber, 2004, 2010) and seaturtles (i.e., Richardson et al., 2009), and the start of a long-term research programfocused on Epicrates chrysogaster (Reynolds and Gerber, unpubl.). Additionalstudies have been conducted on Leiocephalus psammodromus and Anolis s. scriptus(Smith, 1992b, 1994, 1995; Smith and Iverson, 1993; Reynolds et al., unpubl.), andmore recently for Sphaerodactylus caicosensis and S. underwoodi (Reynolds andKoneczny, unpubl.) and Tropidophis greenwayi (Reynolds et al., 2010; unpubl.).Additional work is needed on other endemics such as Aristelliger hechti andTyphlops cf. platycephalus (Hedges, unpubl.).

The TCI are unique for their natural beauty and endemic species, both of whichare responsible for drawing a large number of economically vital tourists. However,this natural beauty is under assault from a plenitude of damaging processeswhich threaten to permanently damage or destroy the environmental uniquenessof the archipelago. Recently publicized environmental disasters, such as a largeamount of raw sewage in Turtle Cove Marina, Providenciales (Godley et al., 2004)could potentially have important economic impacts. It is hoped that the aboverecommendations will contribute to the long term preservation of the native reptilesof the TCI.

Amphibians and reptiles in the Turks and Caicos Islands 401

Acknowledgements. This work would not have been possible without the manyyears of research conducted by my fellow scientists in the TCI. I would like toacknowledge the significant contributions of other individuals working with theherpetofauna of the TCI, especially my collaborator and mentor G. Gerber. A specialthanks to the many field assistants who have helped with herpetological surveys andfield work over the years. I am also grateful to the Department of Environmentand Coastal Resources, TCI for scientific research permits and support, and to theTurks and Caicos National Trust for logistical support. My research in the TCI issupported by the following: Department of Ecology and Evolutionary Biology, TheUniversity of Tennessee Summer Research Grant Program, The Sigma Xi Grant-in-Aid of Research, The Mohamed bin Zayed Species Conservation Fund, the SanDiego Zoo Institute for Conservation Research and the Offield Family Foundation,The American Philosophical Society Lewis and Clark Fund for Exploration andResearch, The American Museum of Natural History Theodore Roosevelt MemorialFund Grant in Support of Research, and The University of Tennessee W.K. McClureScholarship for the Study of World Affairs. This manuscript was greatly improvedby discussions with G. Gerber and M. Welch and comments from A. Hailey and twoanonymous reviewers.

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