Natural History of West lndian Reptiles and Amphibians
Robert W. Henderson and Robert Powell
University Press of Florida
:AINESVILLE .TALLAHASSEE .TAMPA . BOCA RATON . PENSACOLA .ORLANDO ' MIAMI .JACKSONVILLE . FT. MYERS . SARASOTA Copyright 2009 by Robert W. Henderson and Robert Powell Printed in the United States of America on acid-free paper All rights reserved 14t312 111009 654321
Library of Congress Cataloging-in-Publication Data Henderson, Robert W, 1945- Natural history of West Indian reptiles and amphibians / Robert W Henderson and Robert Powell. P.cm. Includes bibliographical references and index. ISBN 978-0-8130 -339 4-5 (alk. paper) 1. Reptiles-West Indies. 2. Amphibians-West Indies. I. Powell, Robert, 1948- II. Title. QL656.5.A1H46 2009 597.909729-dc22 2009017139
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Funding to assist in publication of this book was generously provided by the following organizations:
The Falconwood Foundation through The Conservation Agency, Jamestown, Rhode Island
The Herpetologists' League. 12 Family lguanidae
Figure 15. Aggressive displays of competing male Cycluraiewlsi (lguanidae) may include gaping, lateral presentations, crest and dewlap erection, and elevation on allfour limbs. Such displays can escalate into actual combat if neither participant submits. Photograph by John F. Binns (lnternational Reptile Conservation Foundation)'
Cyclura carinata Harlan, 1824 mosaic (Iverson, 1979). Behavior: Performs headbobs to declare territories, challenge males, and for sex recognition Distribution: Two historically recognized subspecies in (Iverson, 1979); most headbob displays consist ofa series oi the Turks and Caicos Islands (C. c. carinata) and Booby alternating single and double headbobs, each display usu- Cay (east of Mayaguana Island, Bahamas; C. c. bartschi), al- ally beginning with 1-2 single headbobs or a fast "trill" of though subspecific recognition of C. c. bartschi is not war- 6-15 bobs, followed by a set of 1-6 double headbobs, fol- ranted (Stephen, 2005, 2006; Bryan el al, 2007). lowed by single headbob or two, head nods and tail raises Habitat: Xerophilic (Schwartz and Henderson, 1991); 1o- typical of other iguanas also occur (Martins and Lamont. cally abundant on smaller islands with herbaceous to shrub 1998); aggressive displays include lateral orientation. gap- stage growth (Schwartz and Carey, 1977); on Booby Cay, ing, gular expansion, arched back, crest elevation, inflated iguanas on sandy and rocky beaches, rocky areas, sparsely trunk, lateral compression, and single or double headbobs. and densely vegetated areas, and open areas, used sandy aggression also may involve tail lashing and fighting (Car- areas and holes between rocks for burrows, in bushes and penter, 1982); will take to water as an escape tactic, but onlr' (Wasilewski, abundant trees to heights of 1.8 m 1998); most as a last resort (Iverson,1979). Diet: Primarily herbivorous in rocky coppice and sandy strand habitats, sandy areas are at all ages (95.6Vo of all food items and 94.8o/o of volume: required for nesting (Gerber and Iverson, 2000). Auffenberg, 1982); feeds both terrestrially and arboreallr-. Activity: Diurnal activity bimodal during warmer months, 600/o of macroplant species (including a few toxic forms (i high midday temperatures result in reduced activity, winter eaten, with plants taken falling into three categories: activity reduced and primarily at midday, basks following Common staples of high caloric content, (2) commor. emergence and prior to termination of day's activitl', rest staples of low caloric value (roughage), (3) foods eaten of day spent feeding, interacting with other lizards, and frequently in spite of their uncommonness (plants of hig: thermoregulating, which involves shuttling in sun/shade preference levels), certain individual plants are repeatedi''
114
\- Family lguanidae . i15
browsed whereas others of the same species are ignored, populations on French, Six Hills East, Bay, and Middle cays caloric content in the form of nonstructural carbohydrates exhibit larger average adult sizes and reach maturity more is not an important factor in leaf and fruit choice, animal rapidly (1.5-2.5 years versus 6-7 years) than on source prey includes crabs (Clibanarius arrd Cardisoma), insects cays, Big Ambergris and Little Water (Gerber and Alberts, (termites, beetle larvae and adults, cicada nymph, dipteran 2005, 2006). Sex Ratio: l:1 at all age classes (Iverson, 1979). larvae), a slug, and carrion (fish, lizard, bird, rodent, and Size: Adult male SVL to 360 mm, female SVL to 292 mm canned meats, especially sardines), during the winter, re- (Schwartz and Henderson, l99l). Survivorship and Life stricted to items that are more difficult to digest (leaves), Expectancy: Survivorship positively correlated with body rvhich may pass through the digestive tract almost intact size, life expectancy 14.0 years (Iverson, 1979).TailAutot- (Iverson, 1982); cannibalism (Iverson, 1979; Aufenberg, omyz32.60/o of males and28.60/o of females on Pine Cayhad 1982); one well-developed circular valve and 4 more dis- broken tails (Iverson, 1979). tal semilunar valves partition the proximal colon (N 15), = Conservation Status: The species is listed in CITES Ap- presumably to increase digestive efficiency (Iverson, 1980); pendix I (UNEP-WCMC, 2008); 'tritically endangered" on truits of Turk's Cap Cactus (Melocactus azureus; S6nchez the IUCN Red List (Gerber, 1996a,2004b;Gerber and West Mufloz, 2008). Home Range and Territoriality: Adult Indian Iguana Specialist Group, 2000); both C. c. carinata males are territorial throughout the year (Gerber and Iver- and C. c. bartschi as "threatened" on the U.S. List of En- son, 2000); average home range 980 m2 in females, 1,260 dangered and Threatened Wildlife and Plants (U.S. Fish & m2 in males, 1,590 m'zin dominant males (Iverson, 1979). Wildlife Service, 2008) primarily due to predation by in- Parasites: Nematodes (Alaeuris travassosi, Ozolaimus troduced mammals (dogs and cats), competition with and monhystera, Paralaeuris cyclurae; Dosse, 1939); very large degradation ofhabitat by introduced herbivores (e.g., cattle oxyurid nematode (Cyrtosomum mega) populations in co- and goats), habitat alteration by human activity, and exploi- Ion, perhaps mutualistic rather than parasitic (Bowie and tation by humans; a conservation and management plan Fratz, 1974); to >15,000 presumably commensal or even exists (Burton and Bloxom, 2007a); domestic dogs and cats mutualistic nematodes in a healthy adult (Iverson,l9B2). reduced the population (estimated 5,500 animals) on pine Physiological Demands: Glucocorticoids (used to evalu- Cay almost to extirpation in 3 years following construction ate heightened physiological demands under varying con- of a hotel and tourist facility (Iverson, 1978). ditions) significantly greater in female C. c. carinata than in males, tended to be inversely related to body condition in females but not in males, in |anuary, somewhat higher Cycl u ra col I ei Gr ay, 1845 in the morning, affected by reproductive state and possibly Distribution: )amaica, Hellshire Hills (extirpated elsewhere by seasonal rainfall (MacDonald et al., 2007). Population on the island, including Goat and Little Goat islands). Density: Estimated densities along flush transects on pine Habitat: Xerophilic, tropical dry forest in low limestone Cay 1.7 -3 6.8 adults/ha ( Iverson, 197 B) ; 9 0.3 I ha in optimum habitat for juveniles, 31.1/ha for adults (Iverson, 1979);558 hills along shores and in areas between the coast and in- terior uplands, iguanas in 14.5/ha on Booby Cay, Mayaguana (Conners et in hollow trees in pastures, where congre- gated (Schwartz and Henderson, 1991); retreat a1., 2005, 2006). Population Sizes: Estimated 50,000 C. c. site fidelity ;arinata and 1,000 C. c. bartschi (Alberts, 2004); estimated is strong, with hatchlings preferring dead hollow trees 6-15 30,000 C. c. carinata and 200-300 C. c. bartschi (Hudson cm in diameter with a northwesterly aspect (Van Veen and and Alberts, 2004). Predators: Feral dogs and cats (Iver- Wilson, 2004, 2005);17 of 155 observations of headstarted son, 1978, 1979; Mitchell et al., 2002; Mitchell, 2003; Ger- iguanas released into the wild were in trees to heights of -7 m (D. S. Lewis et )er and Alberts, 2005,2006). Reproduction: Cycles are a1.,2008). with climatic 'r'nchronized cycles, courtship in early May, Behavior: Will climb, can climb a smooth fence post (C. B. :robably serial polygyny with monogyny among some Lewis, 1944); headbob displays consist ofa single headbob nales, eggs 2-glclltch in early |une, females defend bur- or three single headbobs with pauses of 0.7-1.0 sec between :ows for several days to several weeks after nesting, but are them (Martins and Lamont, 1998). Foraging and Diet: :rot territorial at other times, hatching occurs after approxi- Leaves, fruits, and flowers of a wide variety of plant spe- nately 90 days, hatchlin gs averuge79.8 mm SVL and 14.6 g, cies, supplemented occasionally by animal matter, includ- 'uvenile growth rate averaged 19.2mmSVL/yr, males reach ing snails (P. Vogel,2000a); of 25 types of plants consumed, sexual maturity at approximately 220 mm SVL, 375-475 g, 3-4 species of vines are most important (Van Veen and rge about 7 years, females 185-200 mm, 200-300 g, 6-7 Wilson, 2004,2005); foraging in tree canopies (D. S. Lewis ;ears (Iverson, 1979); individuals in recently established et al., 2008). Home Range and Territoriality: 0.59-5.82 ha 116 ' Part ll. Reptilia, Section 1. Squamata (lizards) for 6 repatriated iguanas, home ranges overlapped consid- xeric lowlands), also Isla Beata, ile-de-la-Petite GonAve, erably, probably reflecting recency of release (D. S. Lewis ile-de-1a-Tortue, ile Grande Cayemite, and possibly ile-de- et a1., 2008); injuries presumably from fighting in released la-Petite Cayemite and Isla Saona. iguanas are indicative ofterritorial behavior (R. van Veen Habitat Xerophilic to semimesophilic (Schwartz and Hen- in D. S. Lewis et a1., 2008). Movements: Farthest distance derson, 1991); dry forest with sand, coral, or eroded lime- traveled from release site (N = 6) 196-455 m (D. S. Lewis et stone substrates in coastal terraces, main-island lowlands, a1., 2008). Population Size: Estimated 150 (Alberts, 2004); and satellite islands, often using crevices as retreats or dig- estimated 100-200 (Hudson and Alberts, 2004). Preda- ging burrows in sand or fossil coral (Mertens' 1939; Carey, tors: Hatchling consumed by Epicrates subJlavus (Wilson 1975; Noble, 1923c; Ottenwalder, 2000a); basks on roads 2005, 2006); introduced predators include and Van Veen, (RP); roadkitl recorded in 1939 (Mertens, 1939); readily mongooses (Herpestes javanicus), rats, cats, dogs, and pos- climbs into trees to feed (Powell et al., 2002). sibly pigs (C. B. Lewis, 1944;P. Vogel, 2000a; Wilson et a1., 2004a,2004b); dogs used for hunting pigs are able to take Behavior: May take to water as an escape tactic (Noble, even adult iguanas (Woodley, 1980). Reproduction: Age at 1923c); individual basking in full sun on an angled tree maturity 7-8 years in captivity (R. Hudson, pers' comm. trunk 2 m above ground escaped into Opuntia thicket to j. B. Iverson); nesting activities "apparently late March when disturbed (Mertens, 1939); head "rolls" and headbobs to early May" on Goat Island in the 1930s (C' B. Lewis, used in displays (Martins and Lamont, 1998); often become 1944); females begin digging trial holes long before egg "tame" with feeding or regular exposure to humans (Ban- l*yrrg, eggs deposited in nests in two known communal bury et a1., 2000; Banbury and Ramos, 2005). Diet: Feeds nesting areas in mid-|une, with hatchlings emerging 85-87 on "harsh fruits and shore crabs]' responded to bait com- days later, females aggressively guard nests up to 2 weeks posed of mashed sweet potato soaked in rum (Eyerdam, after egg laying, using threat displays, biting, and chas- 1954); seeds of Ziziphus rignoni germinated more rapidly ing, clutch sizes average 17 eggs (16-20), hatching success after being consumed and passed by iguanas (Hartley et ranges 0-100% and appears to be related to female body al., 2000); dermatophagy (Weldon et al., 1993); a variable size (P. Vogel, 1994);in 2005, ovideposition ranged from24 number of circular (3-4) and more distal semilunar valves increase May to 20 |une for 14 nests in two known communal nest- (5-7) partition the proximal colon, presumably to ing sites (Wilson and Van Veen, 2005, 2006). Size: Adult digestive effrciency (Iverson, 1980). Longevity: In captiv- male SVL to 428 mm, female SVL to 378 mm (Schwartz rty,22years,11 months and still alive at last report in 1990 and Henderson, 1991). Survival: Eight of 20 headstarted (R. Noegel in Slavens and Slavens,2003). Parasites: Include iguanas survived 24 months or more (Wilson et a1., 2004a); ticks Amblyomma albopictum (Carey, 1975; Wiewandt, all 6 repatriated iguanas survived at least 3.5 months (D. S. 1977; Guglielmone et al., 2003), A. cruciferum (Neumann, Lewis et al., 2008). Weight Loss: Over at least 3.5 months 1901; Morel, 1967), and Ornithodoros elongatus (probably, after release, 6 headstarted iguanas lost an average of 1.69 found in container with an iguana; Kohls et al., 1965; Mo- (Atractis g/day (D. S. Lewis et al., 2008). rel, 1967; Guglielmone et a1., 2003); nematodes cruciata; Linstow, 1902). Population Size and Density: Conservation Status: The species is listed in CITES Ap- 0.03 iguanas/ha east of Pedernales in 1985 (Inch6ustegui, pendix I (UNEP-WCMC, 2008); 'tritically endangered" 2003b); 0.70 iguanas/ha on Isla Cabritos (G. Santana in in the IUCN Red List (Gibson, 1996b); 'tndangered" on Inch6ustegui, 2003b); estimated 17,000 (Alberts, 2004); es- the U.S. List of Endangered and Threatened Wildlife and timated 10,000-17,000 (Hudson and Alberts, 2004). Popu- Plants (U.S. Fish & Wildlife Service, 2008); principal causes lation Structure: )uveniles compose about one-third of the include degradation of habitat by charcoal burners and population on Isla Beata (Grupo laragua in Wiewandt and predation by introduced mongooses (Herp estes j avanicus), Garcia, 2000). Predators: Humans (primarily for food), dogs, and cats (Woodley,1980; P. Yogel,1992; Alberts,1993; feral dogs and cats, mongooses (Herpestes javanicus), and P. Vogel et al., 1996; Wilson et al., 2004a, 200ab); ongoing pigs take adults, juveniles, and eggs (Ottenwalder, 2000a); threats include proposed development of the Hellshire feral dogs (Rupp et al., 2008). Reproduction: Age at matu- Hills (Woodley,1980; P. Vogel, 2000a). rity 3-4 years (Ottenwalder, 2000a); in captivity 3-4 years (Boylan, 1984, 1985); eggs warmed by sunlight in tunnels 0.4-0.6 m long (Noble, 1923c; Haast, 1969, in captivity); ag- Cyclura cornuto Bonnaterre, 1789 gregate nesting on Isla Beata (Noble, 1923c); eggs in two Distribution: Hispaniola, widespread, but disjunct distri- clutches (in captivity at the San Diego Zoo) averaged 68.'l bution on the main island (probably once islandwide in x 46.7 mm and 79.8 g (N = 21) and 67.6 x 48.6 mm and :(LL6'1le:ra) pue zu?.{tqrs;tsu83{ 'r'3) seurnxg Ierlual (g-d t pue relnrrD (g-z) v dgepotu puB '(I = N) aputou! eql ul (suap;o alrs ,(lqeunsa;d) rorrelur dlcor aq] ul puP 'i ') vI seAIBA reunlltues € pue r?lnrrlf, t '(€ = N) otnlqclt eql Suop :(9967 'uosrell ;7161'la:r-3 pu? zuemi{rq 'r'J ul saAIP,\rBunlltuas t-zpue relnfrlf I:@2002 pu?rls Plslp s?arP serueqeg :alourout 'r '3) auo1sau11 peqruoodeuoq 'ar?q Jo ''le 1a saulH :rn4tolpun? snwryttr) sprrq8uqrory Surpnltur 'serunxg ural{uou aq} q pws Pue pue (op4datocnal sauaor7air4) suoa8r6 pou'.,tor-a}Il{M puqdoqns snu$n1) seare uI :(qOOOZ 'ddeq1 :rsu83{'c D :9OOZ'sua'\\O Jo esoql dlqrssod pue s8ullsau (uarunutaql {ror Jo '3) pue ddeu; :0002 'rlelg pue raolrng :anlqch'r sua: rale^\r?eqs s,uoqnpny se4el ,(1a.tr1e1ln)eJ olourou! 2 'J -rcq (ssuawaLpq 'te^ oaoqtrw snur6) autd aql ;o '(doue: :(qtOOz ''P la uos;all) spnpr,trpur r*a; l1a'r'r1e1ar '(q paruns uado aq1 rapun sI lellqeq parra;ard auo 'sale;lsqns {ror lo -uor aq o1 sreaddt lsou q8noqlp'(q9002 '.89002'saulH ro I{111v\ uor1e1a8a,t;o seeJ? pue 'uorlela8a'r-Io pue ddeql :e9002 'qt00z ''Ie P uosre^l :0002 'uosre^I pu?s lnoqll1}\ alnp pro^ep auolsatull rlrax 'sal{reaq .{puts :(1661 'uosrapuaH :ttlourout'r '3) de3 J?a'I uo .(lercadsa 'slsuno] tuorJ zu?,{rr{rg) cryqdosaululas 01 crpqdoray :}ellqeH -ord qsar; pue sdens alq?l salel :(0002 'uosra^l:olautout pu? 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Reptilia, Section 1' Squamata spp. and Salmonella spp'' and 4with Entero- (N 14) partition the prox- Staphylococcu.s semilunar valves in C. c. figginsi = normal rpp.lare probably commensals or part of the increase digestive efficiency ,orru'r imal colon, presumably to (C' inornata; et enteric microbial flora, respectively c' fames drink rainwater from depressions in rocks (Iverson, 1SSO); (Amblyomma dissimileon C' c' cychlura and Both source (Leaf al., 2006); ticks et a1.,1973).Genetic Diversity: 2005)' Popu- iWrt.o* C. c. Durden and Knapp' (Alligator Cay) populations of C' c' A. torrei ot figginsi; Cay) and translocated Guana Cay' Exuma and hetero- lation Density z -3.2lha(C'c'figginsion inornataexhibited low average allelic diversity Andros Carcy, 1976); <0'2 iguanas/ha on South loci known to be polymorphic Islands; zygosity for 8 microsatellite Population Size: Population Growth: Indi- (Knapp and Owens, 2008a)' in-C. iychlura (Knapp and Malone' 2003)' (Knapp' o., Gr'ru.tu Cay (Exuma Islands)' 64 individuals grew 8'33 mm SVL/yr over 3 years vidual C. c. figginsi population sizes 3'500 C' c' cychlura' rate significantly 1995); estimated total (Coenen, 1995); mean annual growth (Alberts' 2004); 1,500 C. c. 1,000 C' c' inornata (1.764 cm/yr) than in females (1'139 cm/yr; figginsi,and faster in males C' c' cychlura' 1'000-2'000 C''' fg- 1989); annual growth estimated 2,500-5,000 C. c. inornatal Iverson and Mamula' Alberts' and 400-500 C. c' figginsi (Hudson and population of C' c' inornata on Alli- ginsi, rates in an introduced from -150 in 1970 to "Per- (35'3 mm and 30'1 1OOS), C. c. inornataincreased gator Cay for both males and females Hatch- frupr'; r,:oo in 2006 (Iverson, 2oo7b)' Predation: were significantly greater th.an for the irm, respectively) striatus (Knapp and C' c' Ii.rg, .onrrr- ed by Epicrates fowleri founde. population (Knapp, 2001a); one second-year 2004; Knapp et al', 2004b' 2005) and by Cubophis pu.trl.. (Exuma Islands) had SVI' =13'7 cm Owens, figginsio., Cay apparently gape-limited Home vudii (Knapp et al., 2004b, 2005); mass 118 g (Knapp et a1'' 2004b' 2005)' ui"a Uoay = predation attemPts on iguanas (Knapp ha for females' the larg- C. vudii fult.a i" Range: :O.SS hu fo, males and 5'63 1743; Owens, 2008b); eaten by humans (e'g'' Catesby' Cyclura (Knapp and Owens' and est r"eported for free-ranging 2004b' 2005); hu- U Barbour,1904; Knapp,2005; Knapp et al'' tife History: Survival of C' c' inornata higher on 2005). (for feral pigs (on nests)' feral dogs and possi- human activity)' higher mans food), Cay than on Leaf Cay (with more et al" 1999; bly cats on Andros Island (C' c' cychlura;Knapp males (which are "bolder")' and exhibited in iemales than 2000); humans (for illegal wildlife zero' presumably Buckner and Blair' a declining trend as growth rates neared 2000b)' trade), dogs, and possibly rats (C' c' figginsi;"1upn' are near or at carrying capacity because tliese populations "guard" females on Andros (C' c' cy- captured Reproduiion: Males (Iverson et al., )OoOa) ; of 16 juvenile C' c' inornata male chasing an intruder to 150 m' males at age 12'5 years chiura),attendant and marked at age 0.5 years, 9 were alive head- engage in displays, face-offs' and lateral d rate of 95'3o/o' of 4 ofequal size (typical age at maturity), an annualize on Andros p.,rht.,g (Knapp and Owens, 2008a); copulation marked at age 1'5 years' 3 were still alive 10 oii..;ouJrrl.s in Knapp and Owens' 2008a) and rates recorded on 24 April (P. Hartman an annual rate o 97 '2o/o (highest y.u.,late., (A. Mitchell in Knapp and owens' 2008a); nest on demonstrates extreme zs Aprif for juveniles of any lizard), life table damp sand on 6 Guana Cay (C. c. figginsi) covered with very high survival throughout life' greatly de- longevity, blocked by driftwood apparently placed by a studied lizard)' extreme )une, entrance layJd maturity (longer than for any Guana Cay nest-guarding female (Knapp, 1995); nests on low reproductive output' net reproductive iteropariry and deep' with females actively de- (21 match ; long and 8-13 cm and cohtrt generation time years) -61 ."" i*. ='3.4) tunnels but not nests after ovideposi- estimated between fending incomplete the 3.2"-fold population size increase (primarily Nasu- tion (inapp, zOoOa); eggs laid in termite (Iverson ,2OO7a)'Longevity: May exceed 20 1982 and 2004 rnot"td' on Andros Island and defended Mamula' 1989); 36'7 years titermes rippertii) years in nature (Iverson and (C' cychlura; Knapp' in by females on tops of termitaria c' estimated age at first capture) fot C' c' inornata iadding and Owens, 2008a; Knapp et a1'' 1999' for C' c' 2004,2005;Knapp natu.ellverson et a1., 200ab); in captivity figginsi' coarse sand (R' 2004b,2005, 2006); two nests on Andros in months and still alive at last report in 1990 23 year's,6 gastropod (Batillaria.sp') shell frag- Mortality: Death due composed largely of Noegel in Slavens and Slavens,2003)' often used menis, both nests in iermitaria and elsewhere between tree trunks (Iverson et al" 2005); to bing wedged used again by same female)' drum (Iverson' repeatedly (6 of 12 in termitaria : indivi"dualslrapped in a 50-gallon plastic mostly ,r.rti.rg began in eaily May through mid-June' Sites: On Guana Cay (Exuma Islands)' major 2008). 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Part ll. Reptilia, Section 1. Squamata (lizards) Cyclura lewisi Grant, 1940 at least 45 species of plants in 24 families (-807o leaves and 20% fruits), particularly important are leaves of Rhynchosia Distribution: Grand Cayman Island, once islandwide. minima, Chamaecrista nictitans, and Stylosanthes hamata, whole plants of Spermacoce confusa, and seedlings of Wal- Habitat: Holes in limestone (C. B. Lewis, 1940); in sandy theria indica, all of which are weeds that thrive in disturbed and rocky scrub (Schwartz and Carey,1977); currently ex- open areas, iguanas gorge on fruits of Picrodendron bac- ploits a mosaic of natural and somewhat disturbed habitats catum, Hippomane mancinella, and Myrcianthes fragrans in response to availability of thermoregulatory sites, food, when available, these occur primarily in stands of primary substrates suitable for nesting, predation pressure, and hu- vegetation, the diet is presumably very different than it was man activity (Burton, 2000) ; captive-bred animals released prior to human settlement, exceptions to strict herbivory into the wild "preferred' modified habitats over natural include consumption of fungi, crabs, and cicadas, with the habitats and used artificial retreats more frequently than latter presumably scavenged, coprophagy and soil inges- natural retreats (Goodman et al., 2005a). tion have been observed (Burton, 2000); slugs (Veronicella Activity: Iguanas emerged earlier and retreated later in sp.; Goodman, 2003, 2007); primarily herbivorous, eating summer months than in the fall, most of the time inactive, plants, the majority of which were noncultivated, but ge- usually in states of low or medium alertness, of the small ophagy, coprophagy (including that of an agotti, Dasy- percent of time males spent active (20.3 ! 2.3o/o summer, procta punctafa), consumption of invertebrates (climbing Pseudosphinx tetrio) oc' 19.6 ! 3.Oo/o fall), they moved (8.3 ! 4.2o/o,6.4 ! 2.5o/o), for- into tree to feed on larval moths, aged (8.1 ! 4.8o/o, lO.2 r 2.2%), and engaged in social situ- curs, one iguana chased low-flying butterflies for -5 m and ations (11.4 x 2l.lo/o, 1.9 ! 3.2o/o), much inactive time spent attempted to catch them, consumes shed skin and debris basking (29.9 t 9.2o/o,23.5 ! 4.4o/o), females active (18.3 + (plastic spoon, plastic bags with food scraps, metal twist-tie, o/o, feeding 1.7 16.9 t 4. 4o/o), movin g (6.1 t l. 4o/o, 5.7 t 2. 4o/o), f o t aging with efforts to climb into trash cans), supplemental (9.0 ! 2.5o/o,8.3 t 3.4o/o), engaged in social situations (20.7 contributed little to the overall diet, but did result in ag- t15.2o/o,12.7 t20.3o/o), basking (23.5 x 4.2o/o,23.9 19.3o/o), gregate feeding of females and juveniles, but never males in general, iguanas spend little time in trees or bushes or (Goodman, 2007). Home Range: Released, head-started inside retreats during the day (Goodmary2007). Behavior: females had mean home ranges of 0.24 ha, released males Produces an "S-roll" (the nose is raised and lowered as in 0.65 ha with considerable overlap, for a density of 4-6 igua- a headbob, but is also moved in a horizontal plane during nas/ha, attributable largely to an unnatural single age class the lowering motion), although some individuals appear to in an area surrounded by unpopulated habitat (Burton, produce standard headbobs in addition to the S-rolls, the 2005a, 2005b,2006); mean home ranges of males in sum- twisting motion is not as dramatic as the full rolling mo- mer (13.567 ! 14.426 ha using minimum convex polygon tion produced by C. cornuta and C. stejnegerl, displays are and 4.559 t 3.517 ha using 95% contours for fixed kernel series of -5 S-rolls separated by pauses of -0.7 sec, a pat- use distributions) were larger than in fall (1.600 + 1.656 ha tern very similar to that of C. nubila from Cuba (Martins and 0.804 t 1.102 ha) and much larger than that of females and Lamont, 1998); adult males show little long-term fidel- in summer (0.623 ! 0.312 ha and 0.209 + 0'181ha) and fal1 ity to retreats, but females appear to occupy small, fixed (0.856 1 0.754 ha and 0.467 x 0.486 ha), with males moving territories centered on 1-2 favored retreats, the number of longer distances in summer than in fall and than females in suitable refugia may control female distribution, whereas either season (Goodman et al., 2005b). Hybridization: male density may be regulated primarily by territorial in- Cyclura nubila caymanensis will hybridize wlth C. lewisi teractions (Burton, 2000); hatchlings tongue-touched the if "brought together" (Davis, 2000; Burton, 2004c). Lon- ground as they moved away from nests, 3 of 5 climbed into gevity: In captivity, at least 54 years, estimated -69 years "murdered" trees within 20 min, suggesting greater arboreality than in (Adams, 2004). Mortality: Seven iguanas b1' adults (Goodman and Burton, 2005); extensive tongue- humans at the Queen Elizabeth II Botanic Park (Binns, touching of retreats, substrates, and feces suggests the 2008a; S. Hicks,2008). Parasites: Piroplasm Sauroplasma importance of chemosensation, supplemental feeding ap- in erlthrocl'tes of captive-reared juveniles (Alberts et a1., peared to increase incidents of aggression toward humans, 1998). Population Density: Density of head-started females iguanas responded to increasing levels of human and ve- 1.6-2.0lha (Burton, 2005a). Population Size: Estimated 30 hicular disturbance (Goodman, 2007). Diet: Plums are in nature (Alberts, 2004); estimated 100-175 total (Hudson preferred over all other fruit, mangoes are taken in season, and Alberts, 2004). Predators: Wild and domestic cats and and pawpaws are eaten by captive animals (C. B' Lewis, dogs, and humans, who perceive iguanas as a threat to their 1940); almost exclusively herbivorous, known to consume crops (Burton, 2000, 2007); Cubophis caymanus preys on eqt uaq,4{ (eppnu Sur,rnp ..p ,seqreeq olrlre lsoru :(666I la ourpeqtrS ,(lprradsa uo uoruuro) :(696l ,zl:r.lvitl)S pue opu zan8Jrpog) q 008I-0€gI pue q 00zI-0€60 ruorJ 'lepotu -reg) aarl r*olloq e ur elerueJ :(q6661 ,ourpaqrs ,uosees 1un:1 ]ppe -rq sr .(1r,rr1oe ,(urer aq1 Burrnp (epplu punoru zanBrrpo6 :qg96I ,zl.remr.l)S pue oprrreg) auolsarurl .,qtoor ,(1uo pue Sururqs sr uns erll uaq,ta dpo sa8nyar Buraeal .apmg) -3op,, pue saler ur a8n;ar s4aas :(IS6I Bur4urrp :o_r 'sr{}uotu talool aql 8ur-rnp s.,(ep dure.r ro ,(pnop uo uaas ;elerrr 4ce1d1ala1duoc ]eq1 sea.re ur earl deru :(OtOt ,uapsuleU -(1arer'1eu.rnrq :,!1,rpcy . (W rct.oprrreS),,sueunq ;o Llrp pue ;noqreg) Suotseg roy sa8pal pue Burprq roJ saf,rrrerl -rnsqe pue o8a aql o1 anp Surreaddesrp sr euen8r aql se Bur r{IM selldllard pue slpq d4cor :(7961 ,ure-ra4) srolrladuo: -lseralur pue pcrddl se IBruruB ue ]Bql alqeluaruel sr 1r,, pue aruesqe aq1 dq palelrrrte; ,(lqrssod .serrnoser Jo lerleds ,1o 'snor^qo sr Llrarlre ueunq .,firsuap ueq,!\ luasqe ro areJ 1nq -(1arre,r apr.tr e lroldxe seuen8r :(0ggI ,qre1pung) paualearqt .t\ol ]e ro ]uasqe suerunq uer{,tt .,{an8eu uorurxol seuen8r uaqrrl adetsa ,(14rrnb spurrue qrrqm olur slearlar apr,r.o:d -e3-Bueqes ap o8elgrdrquv ,plseol eqt Jo so,(ec ug :aruupunqy s{ror arer1-rt uauo dprerur;d .eqnJ uo :(1661 .uos -rapuaH pue zuemtlls) rrtqdosau o1 crtrqdo;ay :telrqeH '(sgg7'zpog) '(tVOOZ ,s8urqt 'uoyng) uorlepdod Surpaa.rq e peqsrrqelsa lou sarlrleuoru m{peot q8rq raq}o Buoue .ur sllnsar seq dlqeqo-rd raDBI er{} q8noqtp ,(stsuauow{ac .u .3) uev: sueunq qllm slleluor luanber; e'ter{rrt ,s?eJe paqrn}srp ur -.(e3 pue;g .3) ,sa.(an8eJl pelerluaruot .ueude3 uo pue (o1tqnu.a ocr6 ouand aq ol uaas slppe al11l.I uo :(666I .J elsl uo pef,npoJlur :(stsuauowf,w.,l :..salsl .ralsr5,,) rerg "le ]a ourllaqcg zan8Jrpog) puelpoo,uqnns ,(uroql 1rasap ueude3 pue ,sen8al -nuas .lsaroy puelsl ueude3 al]}1.I uo pue ocur3 pue sno11,(qdorcrtu Ielseofqns pue Ielseof o,(e3 pue (an8eue3-uueqes ap ,eurag '1sero; o,r.or8ueru .lsaJoJ o8elgrdrqc-ry EI ap snonprtaprrues pue uaar8rala snol .soarreue3 urp.re{'adrlag ue5 ap so.(e3 so1 ap o8elgrdrqt:y -1.(qdoseru q :(€002 .syaqly) puelur uer{t saurllseoc Buole ,(spue1,u.o1 'pnluarrn{ e1 ap elsl salrllales uo pue rrrax ur epr-\r luepunqe eloru sa(ur] JnoJ lnoqe seuen8r .,(eg oueuelueng -puelsr acuo,(lqeqord) puelsr urcru aq] uo uorlnqrrlsrp te :(egg61 'erera4) uorlrn'rlsuol Morrnq ur pepuadxa.(Breua lrunl -slp e r{qm 1nq 'peardsaprm areqm ,(a11qnu.zr .3) eqn3 uo Surznurunu dlqrssod ta11el aql .ra;ard seuen8r ,1uasa.rd ere rncco sarcadsqns pezruSorar yros,6po.r .rns .(puer_rnr o.rJ :uorlnqrJlsr(J pue .(pues qloq orar1m Iap o8;u1 ode3 uo :(9002'eurtuery pue sese3-saprloeg) r,l(ls o.,{e3 uo saunp [tBt'^elD \ilqnu Dlnp{) rl)eeq ur dlrrerurrd smorrnq :(qOOOt ,ourlteqls zanB;.rpo6) s{ror ro seaJJ uo punor8 aqtr e,Loqe ru g ol taqurltr }uellafxa ,arped 'saaor8ueu ur osle Iapzru)ode3 uo prerutqlnos (a8n 'slBruurBru -;arTseuun8r I'I-9.0 r{11^{) s.rorrnqJo uorleluerJo }ueurtuop Iela,I dq uorlepa,rd,(q -ard :(1661 'uosrapueH pue zgemqf,S) oqnuwrs uasrng paterrldruor uorleralle pue uorlrnrlsap tEl -lqeq o] enp,(preurrrd ,(8002 .err^res oJIIpllM .S.lr lllercadse '.se^eal ualleJ Jo ra ol punor8 ur pue .rlJer .sol g gslC .slerraleu sluuld pup aJnplr6 pauelearr{I .S.jl -laurpd 'sern-( Suorue Burppnq pue reqrunl pue pa_ra8uepugSo }sl.I .dare:)) Jo aqt uo :(eruOZ,uogng) salrd ur :(996I sasnoq alrleu punore cerg uuru.(e3 ..para8uepuq, tq.I pau H.JOI aql ,erered) ur :(SOOZ .OWCI uo salrua,rnf :(0002 uare ure]unou trlsrB{ puplur .,para8uepua,(11ecr1rrc,, -dIINn) I xrpuad ue slrq€qur uorlelndod -dv ssrtc uI polsrl sr sanads arLL :snlels uorlglresuoJ palelosr ue pue salorBueu Ielseol ,srma.I .3) ur sa8ero; :(116l 'g sa,ror8uerrrJo sar{ruBrq pue .(.ruuoc .srad.uo1;ng .l J) {red rruel $iunr] paurpur uo s{seq:(qgOOt ,epe,qsg) g;o ru slq8raq o1 -ofl II r{leqezlg ueen} ur pnprlrpur Bur8uer_aar; e tuoT seqsnq pue saerlJo seqtruerq uo qrurl).(eu .ooo3 ode3 .uaas uo paprorar se.,n (ruru SZr) TAS elerueJ lsa8.re1 aql uaaq :(086I '.zere^ly sapr,ro.reg) uorlela8a,t luatelpe ur ro r{f,Eaq a^eq ,urru slunpr^rpur ra8.re1 q8noqtp 0g9 ot TAS alptu aqi uo sJenpr^rpur ta8ml,raqp,(afiuow atoqdozrqy) saroti ultrou) te4s,'(LOOZ,ruual pue 1ue.rg) vrw L6.IAS Bull -uetrr ur apua,tn( euo ,oueso6 ode3 uo:(Zg6l,aJIIrlnS .ltrraqdec 1ap -qlteq u?etu'rurr 9t x 99 s33a ul :(E002 .uoung uI axES '11 '1) rea4 ,{14rg16 q}lm pelellosse ,(1ensn rapel pu? .arnlerodual uBupooD) raqtoul;o elBls IeuorlrJlnu ro atp u'(a4un[6) rea4 d11crr4 dq pa8pa ste{rlql poo.upreq alerlsqns ,(Zt ,f,tS 'erntsroru ot elqetnqrr11e ,(lgrssod = pue ,pos .suorlercosse lf '{sla{llr{l ruled rrrolpqs uo sser8 lue1d ruur 9'€ + 8'00I) s8urqcleq parq-aarlder ueril relletus (9 = aerql qll^\ '[rq auo]selurl papora ue;o ernsodxe urer{uou N '1AS wur v'L + €'b6) sSurn{cleq pltll{ :(qt00z .uoung) aql uo 'o3.re1 ode3 uo :(g002 te8nr)) l.ro; plseor rrrolsrq uorleqnfur ur d1,rea parp qtrlnlt puofas r.{fea ur Z seaJeq^r e spunor8 uo :(8€66I ,epe_qsg) ,saqrlnp ;o luaudolaaap dq pepa; 'ialqer^ seqtlnll ]srg ur s33a p pue puoles ur esoql --1e,(11err1eruerp suorlnqrrtsrp :(eOOOZ trtq:al : ueqtr ,ra1e1 IBlseol L L6l ra3,re1 seqrlnlr lsrg ur s33a sdep 69 eroru 9 pue ',ia:e3 pue :g96l zlreMqls tadoo3) selet llerus puu salrlar) aun{ uo s33a prel epural 91 0I puolas e ta1e1 sdep €g ororu / ur s-l\orJnq roJ alqelrns _(pepcrlred sr rerg ueu.&3 uo ruJoJ pue.,(e1q 77 uo s33a 6 prel apual auo dlpnuuelq lsau Leyl -te1d pasodxa aqlSo .s4ro.r lerluao sa8pe pasodxa qlr.tr asoql :uoglrnpordag '(9002 ,uotrng pue uerupoo3) s8u[qt1eq g71 aepruen6l I;rure1 122 ' Paft ll. Reptilia, Section l. Squamata (lizards) sun is highest (Buide, 1951); active 0730-0830 h (tempera- twisting motion is not as dramatic as the full rolling motion oC) twe = 27 on clear days, does not emerge from dens on produced by C . cornuta and C. stejnegeri, displays are series rainy days or days when cloudy early, basks until 1000 h' of -5 S-rolls separated by pauses of -0.7 sec, headbobs pro- then feeds (1000-1300 h), rests in shade before resuming duced by animals in the Puerto Rican population lasted activity after 1600 h, retires to dens by 1730 h, juveniles re- more than twice as long as those from Cuba or by C.lewisi treat just after 1200 h, almost always before 1400 h (on the (Martins and Lamont, 1998); agonistic male-male interac- Sister Isles; Grant, 1940c); active throughout most of the tions consist of "lumbering" chases, occasionally with pro- day, emerges on clear days when the temperature reaches longed pushing, hissing, and biting matches, dominant -27 'C, usually around 0830 h, with height of activity oc- males are larger, have higher testosterone levels and more curring between 1000 and 1300 h, rarely seen in the open scent glands (Alberts, 2003); dense aggregations in high after about 1430 h, resumes activity around 1600 h, not seen anthropogenic usage areas exhibited more aggressive be- afterl73O h (Cayman Brac; Carey, 1966); on Little Cayman, havior and social interactions than individuals in low usage begins to emerge -1000 h, most active during the hottest areas (C. n. nubila at the U.S. Naval Base at Guantanamo part of the day (Townson, 1980); at Guantanamo Bay, vigor- Bay; Lacy and Martins, 2003); aggressive displays include ous bursts of social activity follow emergence, relatively face-offs, lowered head, gaping, crest elevation, and head- peaceful migration to nearby areas of vegetation for mid- bobs or head shaking, aggression may involve biting, tail day feeding, ending in a second bout ofsocial interactions lashing, chasing, and fighting (Carpenter, 1982); will enter in late afternoon prior to entering nighttime refuges (Al- the sea to escape threats (Hardy, 1956); juveniles may enter berts, 2003). Behavior: Initially reacts by inflating its body water and hide among seaweeds (Thalassia testudinum;Ro' and "featuring a ferocious aspectl' but quickly learns to take driguez Schettino, 1999b); on Cayman Brac, iguanas cross food ofered by hand (Gundlach, 1880); very shy upon large open areas when disturbed instead ofrunning into a emergence from dens, but "tame" easily in captivity (on Sis- mangrove swamp (Carey, 1966); on Cayos San Felipe, seeks ter Isles; Grant, 1940c); readily climbs smooth fence posts refuge in burrows when chased by dogs trained to catch ju- (C. B. Lewis, 1944); when disturbed, rarely covers the entire tias (Capromys), but fight tenaciously (attacking dogs and distance to refuge in one dash, instead uses short spurts of men) when cornered (Varona and Garrido, 1970). Biomass: 14-18 m, during pauses "like a dog on the alertl' stretching Adult biomass -11.7 kg/ha (Cayo Sijri; Beovides-Casas and its head upward and bobbing, when cornered 'bpens its Mancina, 2006). Burrows: Burrows under bushes, esPe- mouth in threatening fashion and gives voice to a discon- cially Tournefortia gnaphaloides and Suriana maritima (C. certing hoarse noisej' readily climbs stone walls and tall n. caymanensis; C. B. Lewis, 1940); -1.2 m long and -30 cm trees, huddles together on cool nights (Buide, 1951); flees beneath surface (Grant, 1940c); may occupy crab burrows when encountered at substantial distances, retires to thick- (Neilt, 1958); an adult male resided in a small cave on Cay- ets in which they are concealed, sometimes seen in groups man Brac (Carey, 1966); on Little Cayman, lizard wedged in of 4-5 (on Cayo Largo, Cuba), elsewhere usually only a one corner of a small (10-30 cm high, <1 m long) chamber single individual at a time (L. H. Saxe in Sutcliffe, 1952); (Seidel and Franz, 1994); on Cayo Sesenta Arroba, many (7 younger individuals will bipedal when fleeing, if captured, of 19) burrows face southeast, on Cayo del Rosario, most whip tails (Z.Yogel,1965a); males responded to approaches (10 of 16) face south, mean width and height of burrows by headbobbing and broadside challenge displays, interac- 15.94L2.62cmand 10.50 + 1.34 cm, mean length and depth tions involved largest male in a population on Little Cay- 147 .64 t 59.42 cm and 40.56 t 15.89 cm, mean distance be- man chasing the largest female, which, in turn, chased tween burrows 15.04 + 8.56 m, in September, burrow occu- other females, t hatchlings were solitary (Townson, 1980); pancy 60.0%, with two iguanas in only one burrow in Oc- losers in male-male agonistic encounters flee considerable tober, occupancy 79.60/o, with 8 of 19 burrows housing two distances (-7 m), stopping at intervals to look back at the or more iguanas, with 8 iguanas in one burrow, burrow threat, eventually taking refuge in a burrow in sand or un- density on Cayo Sesenta Arroba 7.7 lha (Cubillas HernSn- der a rock, the encounter itself involves the peculiar pitch- dez and Berovides Akarez,199l); on Cayo del Rosario, 13.7 ing ofthe head, gaping, and circling the opponent, collec- burrows/ha (Berovides Alvarez,l998);also on Cayo del Ro- tively comprising ritualized behaviors that circumvent sario, mean widths and heights of burrow entrances 21.58 I actual fighting in many instances (Berovides Alvarez, 1980); 7.48 cm and 9.04 + 2.76 cm, mean distance between bur- produces an "S-roll" (the nose is raised and lowered as in a rows 18.37 r15.82 m, mean density of burrows 14.90 + 9.70 headbob, but is also moved in a horizontal plane during the burrows/ha (GonzSlez Rossell et al., 2001); mean burrou' lowering motion), although some individuals appear to entrance heights and widths 7.62 xl.6l cm and 13.33 ! 4.02 produce standard headbobs in addition to the S-rolls, the cm on Cayo Obispo and 6.05 + 2.99 cm and 14.43 t 5.59 (tol@slp sa)a,I :(666I 'oralo dol) uahof atqompb pue (t002 ''P pue'ata{r,tn Dqopfio) ondang 'suatSol' sau;u tu n ) ]a auoularlSnc :666I 'orelo do3 :v861 'zrlt) elap) aaffip - o t,(111' a ua 1s a qa s o tp r o )' a lP uzuo a a uo ut o ddry1' 1t :N pue ''ds ra.. - i:' sotopoqltulo satru :(866I ''p 1a sgaqly) lenpr.trpur -rad uoJpuapontd Jo slln{ p$e'oasu aqououo) p -read ua{el >:" qrll g'I + g'g 'deg otu?ueluenD 1e sapue,tnf Sur8ue;-aal; - o d1' o1,fiqdo ta U un qa qal Jo sra,t{og'punor ooj6 euolularT8ng :4961 oatot oqa&aua) pue'anopli '1 'aotdoc-sad oauodl 'ot:n -ro uo'ds owwo,fiEav:G,007 '.'le la 'larotr^{ :eig5l 'seran36 zaq4) wnrypunlot awwo,(1quty -aq saqwaqlol,Qg 'atoqdolloqdou,{c '2 'asonxad stu,idrl uo :(E002 "p 1a auotulart8nU:666I 'ora1g do3 :/96I 'larol { 'olnuottp arwqnog Jo sa^eal) ueude3 alU'I Paurns :6961'9961 ,(ura3 n :r0f6l 'tuerD :6€6I '<{rnllqM :e7€6I -uor aq 01 u,nou>l are s1ue1d3o sarcads OV<:(ZOO(. 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Reptilia, Section 1. Squamata (lizards) estimated 40'000-60,000 C. n. nu- contained Balantidium (ciliated protozoan) and nematode nensis (Alberts, 2004); n. caymanensis (Hudson and Alberts, eggs, "healy nematode infestations were found in the large bila and 1,000 C. (50-75/ha) on Isla Magueyes, bowel of all of the iguanas" (L. H. Saxe in Sutcliffe, 1952); 2004); estimat ed 422 igaanas (Alvarez estimated 40 C. n' cay- intestinal nematodes Atractis opeatura (P6rez Vigueras' Puerto Rico et a1.,2007); Brac (F. Burton in Goetz, 2008)' 1935b; Baru$ and Coy Otero, 1969a; Baru5 et al., 1969; Coy manensis on Cayman |. and Cayman Brac, dogs (C' Otero, 1970; Coy Otero and Baru5, 1979b; Coy Otero and Predators: On Little Cayman iguana under artificial condi- Lorenzo Hern5ndez, 1982; Coy Otero et al., 1987; Coy B. Lewis, 1940); a very young panicked in the presence of Cubophis sug- Otero, 1999); O swaldofilaria brev icaudata, Paralaeur is c y- tions fuscicauda, regularly prey on hatchling igua- clurae, Alaeuris travassosi (Barui and Coy Otero, 1969a; gesting that these snakes nas (Carey, 1966); potential predators of eggs on Cayo del Barui et a1-., 1969; Coy Otero, 1970; Coy Otero and Baru5, landcrabs, hermit crabs (Coenobita clypea- 1979b; Coy Otero and LorenzoHernlndez, 1982; Coy Otero Rosario include Grackles (Quiscalus niger), guJls et al., 1987; Coy Otero, 1999); Oswaldocruzia lenteixeirai fus), Greater Antillean (Sterna maxima), w\th (Coy Otero and Baru5, 1979b; Coy Otero and Lorenzo (Larus atricilla), and Royal Terns potentially preying on juvenile iguanas Hernindez, 1982; Coy Otero, 1999); and Ozolaimus mon- the three birds also (Berovides 1980); feral and domestic dogs (on hystera (P6rez Vigueras, 1936; Baru5 and Coy Otero, 1969a; Alvarez, cats (onjuveniles), and pigs (on eggs; Baru5 et a1., 1969; Coy Otero, l97O; Coy Otero and Baru5, adults andjuveniles), and sold as souvenirs to tourists in 1979b; Coy Otero and Loren zoHernindez, 1982; Coy Otero Estrada, 1993b); stuffed humans are not a major factor be- et al., 1987; Coy Otero, 1999); oxluroid nematodes in free- Cuba (M. eerny,l992); iguana meat or eggs is not common ranging juveniles at Guantanamo Bay (Alberts et al', 1998); cause consumption of (C. Perera, 2000); in some areas, are hunted for 500/o of large intestinal volume filled with nematodes (Ro- n. nubila; iguanas and eggs were driguez Schettino, 1999b); nematodes presumably com- food (Fong Grillo et al., 2005a);both on Cayman Brac (Carey, 1966); feral mensal or even mutualistic (Iverson, 1982); piroplasm Sau- considered a delicacy and possibly rats (C. n. caymanensis; roplasma in er1'throcytes of 7lo/o of 1993 captive hatchlings, cats, domestic dogs, feral cats and humans (in the form of road but none in 1994, and 79o/o of free-ranging juveniles and Gerber, 2000a); to be the major threats on Little Cayman >60% of cells of 26o/o of the same free-ranging lizards were mortality) appear (Goetz, Courtship probably begins infected with the haemogregarine Hepatozoon and hemo- 2008). Reproduction: from April to often in ar- coccidian Schellackia (Alberts et al., 1998). Population in April and egg deposition )une, normally occur, eggs 69'0-77'8 Density: Densities declining in many areas due to human eas where iguanas do not of eggs 67.8-75.5 mm length (Ro- activities and, in some areas, hunting, mean density 9'6 mm in length, clutch 9 maturity at 2-3 years otr iguanas/ha on Cayo del Rosario (Rodriguez Schettino, driguez Schettino, 1999b); reach Cayman Island, courtship and 1999b); on Cayo del Rosario, 3 individuals/ha (Berovides age (Perera, 2000); on Little occur in April and early May, females dig nest holes irlv ar ez, 198 0) ; 9. 9 t 0. 65 individuals/ha (6.5'12.8 iguanas/ mating 1730 and 1800 h from May to ha in different months; Gonz6lez Rossell et a1.,2001); densi- and deposit eggs between in beach sand to within m of high tide mark, ties on 11 Cuban cayos 9.3-30.0 individuals/ha, with high- early fune -9 8 and 20 eggs (C. B. Lewis, 1940); ma- est densities (25.0, 30.0, and 30.0/ha) on rocky islets and 2 females contained on Isla Magueyes, courtship be- the lowest density (9.3 iguanas/ha) on sandy Cayo del Ro- turity at 983 g body mass early April, females apparently move sario, where estimates based on numbers of burrows or gan in late March or the mating season, once a female responds distances between burrows were even lower (Berovides A1- to males during her movements to the immediate vi- var ez, t99});densities t atged 22.5-25.2 lha ot't Cayo Obispo to a maie, he restricts up 7 weeks (post-copulatorl' and 10.5-18.2/ha on Cayo Verde in 1998-2000, and was cinity of the burrow for to to leave, he circles her while vig- 14.2lhaon Cayo Sotavento in 1998 (Berovides Alvatez etal', guarding?), if she attempts several burrows initiated before one 2003); densities higher in wet than dry seasons and differed orously displaying, sandy or grassy areas, often in sides by habitat, with density in xerophltic coastal scrub (6'7 + completed, usually in in consecutive years, bur- 6.3lha) higher than in sand vegetation (3.6 ! 2.7lha) ar,d of embankments, nests reused <50 cm deep, females extremely secre- mangrove forests (2.9 x 2.9lha),with the most juveniles in rows >1 m long and defend nest vigorously, eggs 2-14 (mean the coastal scrub (Cayo Sijir; Beovides-Casas and Mancina, tive and do not 46.4 g, incubation 70-78 days, mean 2006); estimated 40 animals on Cayman Brac (C. n' cayma- 5.6) with mean mass 107 mm, major source of egg mortality is nensis) in 1993 (Gerber, 1996b); -1,800 iguanas on Little SVL of hatchlings (Christian, 1987); clutch of freshly laid eggs Cayman (Goetz,2003); -500 iguanas on7.2-ha Isla Ma- ant predation x 42 mm, hatchling SVL 84 mm, 13'4 g (l' gueyes, Puerto Rico (Garcia, 2005b, 2006b). Population (in captivity) 82 1994); on Isla Magueyes, nest depths 26-70 cm' Size: Estimated 40,000 C. n. nubila and 1,500 C. n' cayma- Schmidt, '(46) puelsl euen3 '(qOOOf '.oul]taqrS zan8ppo6) rIural{lo{aq :(985: uo sBurppnq punore Sur8ero; otr asJa pe 1ou saltua,tnf ''lp la uerlsur{3 :sal.an8eyrq u1s1 uo) t{ 00SI o} 9160 tuor: .(1a,rrle1ar sarnle.radulal .ipct ilprcadsa pue s[?.e\ >lror uo {seq IIIM :(9002 taqrag Pu? 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Surga,ruor 1e druortga ra,trol 8ur -dod ueqn3 q8noqtp '(eeeA '{ouluaqrs zan8ppoX uI BrBT -lerrpur ']eJ ssal pue yod arou peq slsau (rp ur s8urlqtteq ozrueqS'V pup our]]aqls zon8yrpog 'a:9661'elarad pue lnq']sauJo prlualod.ralem IIos o] pal?lar 1ou azls 8u[qt1eq suaqly) suorlepdod upqn3 ore se '(q966I 'l.eg) ]sl'I pag :(166I 'eluarm?'I pu? uerlsrrq3) azrs qr1n1c pue qldap zOJ xpuad ol pe1elal suolleJluaJuo) 'uIeJ Supleos reue seler lua -\f:)l aq] uI .,elqereuln^,, :(sooz'lwlM-daNn) 1 -dV SgttC ur palsrl sr saoads aql :snlols uope rasuoJ -reJlp 1e parrp slsau lnq'J. g'€€-g'0€ sa;n1e.radrua1 uearu g7i . aepruen6l ,(1gue3 126 ' Part ll. Reptilia, Section 1. Squamata (lizards) Density: 2.03 iguanas/ha Activity: Usually emerges from dens around 0700 h (mean (Alberts et al., 1998). Population in 1991, with the lower density due ambient temperatures -25 "C), moves to sunny areas and (Carey, 1975); O.36lha forage attributable to competition with intro- assumes a prostrate basking posture, after about I h (mean to degraded herbivores (Mitchell, 2000a). Popula- ambient temperatures -27 "C), begins to search for food, duced mammalian adults and 44 repatriated iguanas on stopping frequently to sample small bites of nearby plants, tion Size: 200 wild 80 in the headstart faciliry 130 on basking continues after feeding, but rarely in full sun (usu- Anegada, with another Necker (Perry and Mitchell, 2003; Ger- ally a sun-shade mosaic), during midafternoons (mean Guana, and 30 on and Gerber, 2005; Lazell,2006; Perr,v ambient temperatures -30 'C), usually remains in full or ber,2004a; Bradley estimated 400 (Alberts, 2004); estimated partial shade, after -1500 h, when temperatures drop be- et al., 2007a); (Hudson and Alberts, 2004). Population Structure: low 30'C, moves into more exposed sites, no iguanas seen 200 (87.50lo of population) outnumber juveniles (12'5Yo)' after 1800 h (Carey, 1975). Behavior: Aggressive displays Adults be monogamous, living in separate but involve stereotypical vertical headbobs (Carpenter, 1982); adults appear to (Carey, 1972,1975). Predators: juveniles engage in aggressive encounters with other ju- closely adjacent burrows (Grant, feral dogs and cats (Mitchell, 2000a, veniles, these involve headbobs and pushups and, on one Dogs 1937b); (Wing, 2001; Lazell, 2005a); Borikenophis occasion, biting, chases often followed by up to 10 min of 200b); humans Kestrels sparvarlus) take displays (Perry et al., 2003b); adults on Anegada extremely portoricensisand American lFalco juveniles Guana Island (Levering and Perry, 2003); also shy, even basking adjacent to dense sheltering vegetation, on SVL, 88-g Borikenophis portoricensis in contrast, hatchlings are quite naive (K. Bradley in Wer- on Guana, a 621-mm a hatchling iguana of 40.4 g, a mass represent- ning, 200 9). Diet Eats C o ccolob a \eav es, O p unt i a ft uJt' arrd consumed pre-meal mass (RWH, RP). Repro- whole wild nutmegs (Grant, 1937b); primarily herbivorous, irg45.9o/oof the snake's appears to be synchronized with two yearl1' but takes some animal matter, juveniles probably insectivo- duction: Cycle females with 12-16 yolked ovarian rous (Carey, 1972,1975);bulk ofdiet consists ofleaves and rainfall maxima, adult presumably due to competition fruits, composed largely offorage not taken by introduced follicles, Ioss of fecundity (Carey, 1972,1975); mating season ungulates (3Oo/o Croton discolor, which contains com- with domestic livestock females dig nests in sandy areas pounds toxic to ungulates, and >55o/o fruits of Byrs onima, on Anegada May-|une, through in which they lay up to 20 eggs Coccoloba, Eleaodendron, and Eugenia, which livestock ig- from late |une |uly (average emerge in the fall (Gerber, 2000b, nores), but animal matter (beetles, caterpillars, centipedes, 13), hatchlings and Gerber, 2005); on Guana' egg laying roaches) forms about 170 of the natural diet (Mitchell, 1999, 2004a; Bradley hatchlings emerge in September-Octo- 2000a, 2000b); native plants taken on Guana include large in |une-|uly and 2007 a) . Sex Ratio: Equal with suggested quantities of Centrosema virginiana, flowers of Tabebuia ber (Perry et al., (Carey, (M:F), with "pairs" no lon- heterophylla, seed pods of Capparis cynophallophora' and monogamy 1975); 2:l (Mitchell, 2000a, 2000b). Size: Male leaves of Stigmophyllon emarginatum and Capparis Jlex' ger clearly definable (Mitchell, 2000a), female SVL to 476 mm uosa (Mitchell, 2000b)' Growth: Adulthood at7-9 years SVL to 560 mm mean hatchling SVL on and 350-400 mm SVL (Carey, 1975); estimated 50 mm (Schwartz and Henderson, 1991); of similar length on Guana weigh increase in SVL/1T based on 5 recaptures and 47 captive- Guana 108 mm, iguanas suggesting that the latter are raised indMduals, maturity at 4-7 years, 37'8 cm SVL more than those on Anegada, (Perry al.,2O07a). Survival: 86% survival (Gerber, 2000b). Home Range: Males 116.06-985'11 m'z (N food-limited et individuals released \n 2003-2004 (Gerber = 5), females 155.47-412.31m2, aside from food, available of headstarted 37.5o/o of males and 11.8olo shelter is the most important factor for selecting a home et al., 2OO7). Tail Autotomy: had broken tails (Gerber, 2000b)' range (Carey, 1972,1975); recent population structure dif- of females on Anegada Heliothermic, exhibits behavioral ther- fers, with home ranges expanding 100-fold with a decline Thermal Biology: temperatures of 16 active animals in population size, and male home ranges averaging 6'6 ha moregulation, cloacal oC (mean 6.3 above air temperatures at (2.2-l2.3ha) and those of females 4.2 ha (2.8-5'6 ha), home ranged 1.2-9.8 "C) (CateY' 1972) . ranges broadly overlap and male centers of activity asso- the time of capture ciated with home ranges of females and those of females is listed in CITES Ap- with burrows (Mitchelt, 1999, 2000a, 2000b). Parasites: Conservation Status: The species "critically endangered" in Ticks (Amblyomma antillorum), with 20-30 (maximum pendix I (UNEP-WCMC,2008); (Mitchelt, 1996); 'endangered' on the 59) ticks per adult lizard, most frequently on the thorax the IUCN Red List Endangered and Threatened Wildlife and Plants and in the region of the femoral pores (Kohls, 1969; Carey, U.S. List of (U.S. Service, 2008); principal causes in- 1975; Keirans, 1985; Guglielmone et al., 2003); piroplasm Fish & Wildlife degradation by introduced mammalian her- Sauroplasma in erlthrocytes of captive-reared juveniles clude habitat ruoJp]\ :(sql1q)nu .r .J) suDIrV Jo ]r{BIS aql ur sdel pue seare Ileurs o,,u.1 dluo ]nq ,peleFtlef se^r eq/seuen8r ' 'f ,Qtapt 9.,; ltjttlstrr'r ) sdel erunxg uraqlnos .t.l) satr[a dlrsuap ,o[o6 3o e oqBJ leeu Bare Ilerus ,{.rarr e ur s,llo]rnc -tES pup puBIsI rope^les UBS uo rnJ)o ,rn8alsneqcul sartadsqns pazru8o Jo sraqunu uo paseq :(qEOOZ ur euelups .3 -ra: ,spuBIsJ .illuar:n: earr{l aJar{,n prueqeg :uopnqplslq :eg6g7 .rn8alsngq:u1) .9961 0002 q ey$.Idpo 1nq ur srr: -rrqe3 el{ uo eqlg pe}etur}sg :dlrsuaq uopulndo4 .(y613; E06t'la6aufa$ llailr u n pi) ''1e 1a eueyq) suatnp snnfiJalu7 pve,aazaop rapaqotafiq a '' ds wru t 1 ca qt M r g, tl@ a tq) uaq) s g,. a ya u o ds tu1as. papn i_- '(900t) .tu -ur setelosr :urouortrIry .(0g61 .uc,, ta rolD :arnleralrT luaulued Ieuopppy Ierralleq lI pupsaluJ -raa1) dcuarcga a,rrlsa8rp ,(i aseerrur o1 dlqetunsa.rd = -r '(oooz ''P te .ournby) 1a uolol Ieturxord aql uorlrlred saalea reunlrtues pu? -.r\od :€g6I saoeds ueloruedsrll pauelearr1lJo slsll Ipus Z (padola,rap IIaM tsrg eql dluo) saAIEA relnrrn 7 :(0002 .-1" ur papnllur se.u. sarcads sqt:(/002.'p 1a errery) rrlqndag 1a .(a1yeg) seuen8r ,(q passed pue perunsuot Buraq :auz uefiurruoq eql ul Ieuorleg oor891oo7 anbru4 aq] le slsxa ,(prder arorrr paleurtu;oB ruou*u sru1dut7 spaas uerSo.rd Jo :larg I{ Surpaarq aarldeo e:(nOZ,.p 1a ddn6) pelcalord '(966I 'luotuE-I pue surgery) sqoqpeaq Z_ sarras .raqlo rou sr ollrnbrrug o8el;o aror{s ulaqlnos aql Buop seuen8r Jo ; -ue uaql pue asned 3uo1 auo dq pazrtollo; -(1luanbarl .rx prllqn,{,q E -iq eere er{lJo lred roleru e :seare eruos lseal lp ur Z'0- 1o sasned qoq-relur qll^{ sqoqpeaq afurs Suro8uo sr qrrr{n ,suetunq .(s1eo8 .aplec E- Jo r$ : Lq uorlelroldxa pue ,sasned IBuorlrppe ue dq pa,no11o; ou d11enlrr,t qtlm sqc\q "3'a) sa.lo,trq-raq uerleturuetu pelnporlul qllm uoplleduol .|, -peeq serras B .uraql '1 .s8op) L- Jo r{ll.tr Suruur8aq dlpnsn uaa..r,: sasoo8uotu,s1er saJo ruJeJ uerluurrueru palnpoJl 3 -aq sasned ra8uol -ur.(q ,(la,rrssarSord qllm sqoqpeaq Jo saur: uorlepard.(uorlcnpord leorreqc pue arnllnrrr8e) sarl E 3uo1 lsrsuor s,(eldsrp :(Zg6I ,aluadre3) sqoqpuaq - r.\rtle ;o Je: uerunq ,(g uorlepe.r8ap pue uorleralle lelrqeq llns { ,(qgeq Jo -rltarr. pord,(]oerals a^Io^ur sdeldsrp aarssar88y :ror^Br{e{ -ar e se dlrruurr.rd tapp.,u.ua9g) tsl.I pe6 N)nI aql .CIr{Cl ' (1OOZ.p ur .,pa:a8uepua d1pcr1rr1, :(3OOZ -dANn) l xpuad 1a ddng :t002 rerue6 :q00(t,; li .alqo-\ -dV SEI1C ur pelsrl sr sarcads aql :snlels uorluarasuo3 lapp.ruello :166I 'uosrapuaHpve zueilqts :l€z6l -o sluaurpas aurreru q8norqt 3ur11nt seurler ou:- '(L002 .e .IAS pue spaq laer: "p la errery :3 Otf ) turu 09, ot aleuraJ d.rp 3o opeq ur suorterlua)uo) qllm ,ollrnbrrug o8el 'qr_ '(3 ooz'r) ruur ot 'IAS eleul :ez$,.(6l6t,le^no r.. s8t tlnpv eror{s -3 .3 ureqtnos aqt Suop laaer8 ;o por ,(q paralor sear. :& pue ortse) dalpeg :II = N 9.gg uearu) B sseur 6.99_9.97 tuo{ luasqe erB suap ,elqelre^e .ipoq ,(Ot ,Lu:uI }ou sr Iros aJarlM sartr-\p: lEt pue = lf Z.VZZueeru) turu 9.L€Z_Z.ilZq$ua1 ,$llol {ror pue s{unr} eerl Molloq asn IIr^r lnq pasodsr il1} lplo]'(II =N 'rrutu 7'19 uearu) tuur 0.16_r.28 145 Bunqrleq ,(so1r.rqe3 ro 'sdrunls ' 131' . aepruen6l ,(lrureg 128 ' Part ll. Reptilia, Section'1. Squamata (lizards) semilunar valves partition the et al. (2000) questioned the appropriateness ofsubsPecific single circular and 4 distal (N to increase digestive recognition, based on a lack ofhaplotypic differences, but proximal colon = 2), presumably (Iverson, Home Range and Territoriality: suggested that a final resolution should be determined by efficiency 1980). the year (Hayes, 2000a; analysis of more rapidly diverging nuclear markers, such Adult males territorial throughout 2000); restricted activity to "what as microsatellitesl distribution once more widespread on Hayes and Montanucci, territories within the vegeta- the main islands and presumably on cays currently without appeared to be rather small fixed kernel estimates of populations (Hayes,2000a, 2000b; Hayes and Montanucci, tion' (Gicca, 1980); mean 9570 males females, respectively, on 20oo). home range sizes for and Green Cay (C. r. rileyi) wete 439 and 628 m'z (S' Cyril, 2001 Habitat Xerophilic to mesophilic (Schwartz and Hender- in Hayes et al., 2004); on North Cay (C. r. nuchalis), sizes plus son, 1991); exploits varied main island communities, werc2,047 m2 for gravid females and 397 m2 for non-gravid coastal rock, sand strand and sea oats, and coastal coppice females (B. Thornton, 2000 in Hayes et al', 2004); on White plant communities on offshore cays, and blacklands cop- Cay (C. r. cristata),86% adaptive kernel estimates of home pice and mangrove communities on cays within inland ranges of 5 males and 2 females wete2,656 m'?(S. Fry' 2001 lakes, but most abundant in the vicinity of limestone out- in Hayes et al., 2004); home range sizes reflect island sizes crops and/or patches of Sea Grape (Coccoloba uvifera) and, and possibly population densities (Hayes et al., 2004). In- on some cays, patches of Buttonwoo d (Conocarpus erectus), juries: 16.0% of 75 C. r. cristata, 12.3o/o of 57,14.9o/o of 87, (C' r' presumably require sandy areas for nest construction and 20.4o/o of Sa C. r. nuchalis from Fish Cay, North Cay, a r il ey i; Hay es, 200 0a) ; coastal rock habitats with Strumpfi and a translocated population, and 33.3o/o of 3,34.5o/o of 35, dense Thatch maritima and Sea Grape, least abundant in 33.7o/o of 86,13.3o/o of 15,75.0o/o of 16,2l.lo/o of 19, and 51.67o (C. qistata; Hayes, 200b)' Palm (Thrinax moruisii) forest r. of 31 C. r.rileyifromGaulin, Goulding, Green, Guana, Lot', had missing dig- Behavior: C.yclura r. rileyi displays comprise series of up Manhead, and Pigeon cays, respectively, of 57,32,60/o of 86, and to 7 single and double headbobs, mixed, with no obvious its,42.7o/o of 75 C. r. ffistata,24,60/0 Cay, North Cay, and a pattern to the choice of single or double headbobs (Mar- 51.9o/o of 54 C. r. nuchalis from Fish of 3,22.9o/o of 35,22.60/o of tins and Lamont, 1998); somewhat arboreal, climbing into translocated population, and 07o of 22, and64.50/o of 3l C' mangroves in cormorant (Phalacrocorar) rookeries on San 84,40.0o/o of 15, 50.0% of 16,36.4o/o Green, Guana, Low, Man- Salvador Island (C. r. rileyi; Paulson, 1966); males on cays r. rileyi from Gaulin, Goulding, had injured tails, with in Acklins Bight (C. r. nuchalis) engage in jousting matches head, and Pigeon cays, respectively, more common involving open-mouthed territorial displays, chase other toe loss increasing with age and tail injuries (Carter 2004). Longev- males out of defended areas, and bear scars of bite marks on rat-infested islands and Hayes, (D. Blody in Slavens and (Hayes and Montanucci, 2000); aggressive displays involve ity: In captivity, 9 years, 2 months to being wedged be- headbobs or pushups (Carpenter, 1982);larger males guard Slavens, 2003). Mortality: Death due (Iverson 2005). Parasites: Ticks females and smaller males may engage in forced copula- tween tree trunks et al., Population tion (Hayes et al., 2004); will take to water as an escape only on Guana Cay (C. r. rileyi; Hayes, 2000a). 500 C. r. rileyi' 200 C. r. cris- tactic when harassed (Hayes etaL,2004). Commensalism: Size and Density: Estimated (Alberts, 2004); estimated Hermit crabs in all and Sphaerodactylus corticola in some tata, and 13,000 C. r. nuchalis C. r. cristata, and 15,000 C' r' iguana burrows (Gicca, 1980). Diet: Observed feeding only 500-1,000 C. r. rileyi,l50-200 (Hudson and Alberts, 2004); for C. r. rileyi, maxi' on fruits of Sea Grapes (Coccoloba uvifera; Gicca, 1980); on nuchalis Green Cay (6 iguanas/ha), maxi- Green Cay (C. r. rileyi), consumes 7 species of plants, with mum population 35 on 28 on Man Head Cay (7.Slha), maximum Silver Sea Oxeye (Borrichia arb orescens,flowers especially), mum population (2.5lha), at or near Wild Thyme (Rhachicallis americana, both flowers and population 30 on Low Cay all probably (Gicca, 300-400 C. r. nuchalis on leaves), and Opuntia stricta (a preferred item despite scar- carrying capacity 1980); (Iverson, 2007c). Population Structure: Per- city at the time of the survey) most commonly represented Bush Hill Cay (White Cay; C. r. cristata),l-l0o/o in scat samples, nonplant items included a Purple Galli- cent juveniles was l7o/o 0 ( mean 8 7o' nule (Porphyrula martinica) and two unidentified song- (mean 670, three cays ; C. r.. nu chali s), atd -20o/o Predators: "Other birds (probably scavenged) and one Bridled Tetn (Sterna six cays; C. r. rileyi; Hayes et a1.,2004). were sea gulls anaethetus) chick (likely predated), conspecific hatchlings than man, the only likely natural predator[s] (capable ofconsumingyoung) and ospreys (capable ofcon- (N = 2), a dead land crab, a grasshopper (iguanas occasion- adult iguanas)" (Gicca, 1980); Os- ally seen lunging for grasshoppers), a hermit crab, uniden- suming both young and (Pandion regularly prey on adult iguanas in tified insect material (in 22o/o of scats), and sand and soil preys haliaetus) (Sphyraena barracuda) attacked (Hayes et al., 2004); dermatophagy (Weldon et al', 1993); a Acklins Bight, a Barracuda .urtu .IAS ro auo uaq^\ efuelorl lnoqllm Surpua puu g> 3ur1se1 'uosrapueH pue zUeMqtS) ,xlil VgZ, ol alerual tlnpp -i11ensn 'pade8 sqtnou qll^{ saqr}etu Surqsnd a)eJ-ol-ereJ 'ruur 909 ol'IAS eleru l1npv:az1g'(7967 ''p 1a sadeg :7gg; Sur,rloaur pue qlSuerts;o slsai pnlrr se Sularas slq8g qlyvr ''p 1a sa.(eg q 0002 'uolurorll 'g) ragolcA ro raqtualda5 'sarrolrrral 3ur1eu rol lq8g sapru 'frlfel adecsa ue se relem ur s8ulqoleqyo aruaS.raua 'pesnal er? slroJtnq aruos lsai ot e{el deur 'suolsnrlur uetuni{ o1 d1.re1n8e-r pasodxa uaq,tr -8ns sluau8er; 1aqs33a 'azrs r{l}nlr qtl^\ pelelarroc d1a-ru euoraq ,,atuet,, uet lnq '.paqmlsp uaq.\r uap e olur 8urlea;1 -e8au ssetu 33a qtyn '(p = X) B y 17 pue (Et = ,rr) tutu z'of -ar Lqs dlpnsn :(qWA,luerg) u.frurpqlpr s,(eup 1ou sr x t'gg azrs 33a utaru 'azrs aptual r{ll.tt pe}ellosse sem osI? IIel aqt .:e{ll-rl3u}sq, tnq '.aloq Ilor e olur Suruunr dq sgearl pue (II - N'o/oy'92 pa8era e luetulsalur a.r.rlrnporda: 'azrs '(erct'trct -aU :rol^uqag lpuerltar6) Suqseq tnoqqm elerual qll.ryr patelerroc ,(la,\Ilrsod '(rt = lr '1'E uearu) g-; (}uatu ro l{llm '}sar sr eurlnor Lpp pru:ou Jo 0/og6 lnoqe azrs qrlnlr D, €e-gZ pa8uer se.rnleradrual lsau 'sreqr.upqr i -arlnbar dlpp e 1ou sr Surpaa;'uooureu? alBI pup Sururou ur seqrlnlf, peurcluof lsau euo 'sJaqrueqf a1dr11nu urpluo-a ar{l q dllrerut.rd .,s1no1oo1,, pasodxe uo $lseq 'Lep aql Leu pue uorllaJrp a8urqr ueuo sMoJJnq ' 57; aepruen6l i(lruueg 130 ' Part ll. Reptilia, Section 1' Squamata (lizards) (2.6-22.2 ha; Garcia et a1.,2007); males exhibited uni- both contenders back slowly away, occasional battles last to ha whereas females were randomly distrib- t h and may involve biting and tail lashing (Wiewandt' form dispersion, uted, mean male home range size 0.6 + 0.2 ha (0'21-0'81 ha; 1977); aggressive displays include lateral orientation' gap- polygon) and 0.09 + 0'05 ha (0'02-0'15 ing, dewlap extension, crest elevation, and headbobs or minimum convex 507o kernal algorithm), mean female home range size head rolls (Carpenter, 1982); aggressive behavior may be ha; "rolls" (one 0.09 ha (0.13-0.34 ha, minimum convex polygon) directed at humans (Tenl6n, 1997a); produces 0.28 t 0.07 + 0.04 ha (0.03-0.13 ha;5Oo/o kernal algorithm)' side of head is raised, when lowered, the opposite side is and 6 0'0o/o)' male - female overlap extensive (mean 86. 6%; 6'4 -10 raised, often repeated in long series) as well as headbobs' overlap minimal (mean 2.9o/o;0-11%), suggest- rolls occur at the beginning ofdisplays (usually) or at the male-male that males are highly territorial and that home ranges end, almost always accompanied by 1-2 series of standard ing .o.r.rpond to territories, sizes ofhome ranges suggest that headbobs, consisting of up to 6 (mean 2'O tO'42) up-and- partially responsible for low density if male car- down motions per series (Martins and Lamont' 1998)' this may be is attained (P6rez-Buitrago et al', 2007b); Blindness: Blind iguanas (Haneke, 1995); 9 of 28 animals rying capacity mean juvenile home range size 297 m'z (0-539 m2: Perez- captured in 1998-l999were blind (cataracts), also observed Buitrago and Sabat' 2007). Movements: Headstarted igua- subsequently (Hudson and Alberts, 2004; Perez-Buitrago nas moved 471-6,396 m (mean 2,844 t 2,122 m), distances et al., 2007a). Foraging and Diet: Descends 'bccasionally (K' traveled between consecutive monitoring days 56-157 m to the tillable area to feed on the young corn and cotton' (mean 76 + 33 m; Garcia et al.'Z}}7);hatchlings dispersed P. Schmidt, 1928); consumes 71 of the 400 plants species m from release site, moving on average -250 ml known to occur on Mona, food includes fruits (especially 102-5,080 (P6rez-Buitrago and Sabat, 2007); another cohort of Opuntia spp. and Hippomane mancinella),leaves' flowers' day iguanas moved 441-1,249 m (mean 633 ! 147 und ,o*. animal matter (especially seasonally abundant headstarted m; P6rez-Buitrago et a1.,2008). Nesting: Plantations of in- Pseudosphinx tetrio calerplllars and pupae), preferred Australian Pine (Casuarina equisetifulia) in nesting plants are those low in cellulose, low in aromatic com- vasive produce accumulations of leaf litter and organic mat- pounds, and easy to obtain, leaves (difficult to digest) may sites ter that iguanas cannot penetrate (Wiewandt, 1977)'Para- pass through the digestive tract almost intact, diets of iuve- sites: Ticks Amblyomma cruciferum (Bequaert, 1932; Mo- niles like those of adults, except for the size of items eaten' rel, 1967) and A. torrei (Maldonado Capriles and Medina known to eat mice in captivity (Wiewandt, 1977,1979);may (A. cruciferum) and nematodes (Wie- "grazd'(moving from food item to food item), presumably Gatd, 1977); ticks wandt, 1977); tick loa ds (Amblyomma sp.) positively corre- selecti.rg favored growth, or'gorge" (sit in one place and lated with SVL, ticks aquired quickly after release from eat everlthing within reach; RP). Growth: Estimated 52'3 (one iguana acquired 7 ticks in 4 days), loads of mm SVL during first year, but one field-captured juvenile captivity headstarted iguanas (28.2 t 25'4 ticks) comPa- grew 60 mm SVL in 0.7 years (85.7 mm/yr; Wiewandt' recaptured (av- rable to those of wild iguanas (32.2 t 22.0 ticks) of compa- iSZz); at5 months, 3 individuals had gained no weight rable size (P6rez-Buitrago et a1., 2008). Population Den- erage -1 t 2.6 g) and SVL increased 0.4 t 0'36 cm' fourth 0.33 iguanas/ha (Wiewandt, 1977), btt may have individual gained 119 g and increased SVL 4'11 cm (P6rez- sity: (Wiewandt and Garcia, 2000); 0'21 lizards/ha (J' Buitrago and Sabat,2007); growth of 9 wild iguanas 4'1-10'5 declined Moreno in P6rez-Buitrago and Sabat, 2000); 2'36 ! 3'3 mm/month (mean 6.3 + 2.2mmlmonth) and did not differ A. using line transects 16 m wide (P6rez-Buitrago significantly from growth rates of headstarted iguanas' iguanas/ha 2000); in study area ofrelatively "natural" habi- body condition (mass/SVL) declined after release to levels and Sabat, 2.7 5 iguanas/ha (P6rez-Buitrago et al', 2007b)' Popula- comparable with those of wild animals of comparable size tat Size: 2,000-4,000 individuals (Wiewandt, 197 7 ) ; l'150 ana wita adults with SVL >400 mm (P6rez-Buitrago et al'' tion (J. A. Moreno in P6rez-Buitrago and Sabat, 2000); 5,093 ! 2008). Home Range and Territoriality: Males select terri- 2,500 (Plrez-Buitrago and Sabat, 2000); estimated 1'500 tories with retreats attractive to females, females select re- (Alberts, 2004); estimated 1,500-2,000 (Hudson and Al- treats according to location and structure, favoring those 2004). Population Structure: |uveniles only 5-1070 with good drainage, ventilation, nearly horizontal resting berts, of population (Wiewandt, 1977) or 2 of 118 observations places under low ceilings, and within -lJ m of surface' if (Wiewandt and Garcia, 2000). Predators: Humans (his- iemales attempt to leave a maleb territory, he pursues her feral cats and pigs (Wiewandt, 1977l' Moteno' and tries to "herd" her back (Wiewandt, 1977 ,1979); mean torically), Rivero, 1998; Wiewandt and Garcia, 2000; Garcia et home range size for t head-started iguanas 19'8 ha (2'4- 1995; 2001; Bendon, 2004). Reproduction: Mating occurs 22.2ha;Garcia, 2OO5a,2OO6a); mean dispersal range of 8 a1., an annual 2-week period, males lie in wait for fe- radiotracked individuals (minimum convex polygon) 10'7 during (arral '(qSOOZ uo :(qZ00Z 'Inar8) seere papooM ur ]uepunqB tsoIu te13o{) ,,pe.ra8uepuq, ro (.peuatearqr,. s? atrtad uo :(qZOOZ'lnarg :8866I 1preq]S) vr Z-I Jo slq8raq palralord :(pI66I'ouaro111) ..pere8uepuq, :(886I'uosuqol 'q) lE saqsnquror{l ul l'llrerulrd salrua,Lnf 'selrs Surdaals pue .,alqerauln ,, :(V16l -Sueqre,ro saer] Bueueq qlp\ peleposse uago '.slseroJ puel Nf,nI eq1 q,.paraSuepuq, :(8002'lwlM-daNn) 1 xrpuac -,r{ol lsrou ur seurler ur 'adnolapen1 vo 'ailaqq aawodl -dv sgllc uI pelsll sr sartads eqr :snluls uolle rasuo] luepunqe Surlrods slsaroJ qllm uorlerf,osse u1 dlluanba.r; :(eL66l 'sleg(l6) saqrtrp aprsp?or uale 'alqelr?Ae sr rolem '(::oi arar{r* satrs 1e s-raqte8 :(9161 's1ag(16) staarls aql ssorle sunr 'lpue^tel A) slret ue{orq peq saprual Jo o/ot'tl pue saleu: LruelgqUeg-'tS uo :(6661 'adroql pue Brlor{lery) saarl yo Jo %o0'09 :r(uo1o1ny peJ-'(LOOZ '.teqes pue o8e.rlrng-zara6 o/ozz sumorr ur sdaays 'punor8 eql uo .(lertr 'ruaql uaamleq .,8ur eJrI Jo $laa.tt 0z lsrg Jo pua w PaleurlsE rle^l^rns '(0002 -dee1,,lq aerl ot earl ruo{ a.toru drtu learoq.re.(pBIq :(986I tolD pue 1a.no4) ruru 80S o]'IAS eleural '{rutu €€S o: 'rapulr)) su?e.rls 3o ,frunr,t aql q dla lsnlrxa uaas 'Blru 'IAS elBW :e4S '(q/002 ''1e 1a o8e-r1rng-zaq4) raqotro u: .uosrepuaH -rr'rJoC uo :(qZOOZ'pnarg :Ql,66I,u?luoJ :166I s>lea.l'r. € dluo '1roqs ,(ral uoseas Surqrleq 'o/o98-gg ssalrns puB z].rB1v\r{rs :E761'1eze1) Ierrlsarral ro IearoqrB 'padunp Surqrleq 'uDI 8'ZI alrs Surlsau e 01 apueJ r dq pa,toru a:ue: aq o1 d1a4r erour are suep.re8 se qrns slellqeq prrgrue rllm -srp rrrnrrrrxetu : (6 L6l' L L6l1pu?.ererM)'IAS tur 0'8f 'srear. palerlosse ueqm Jo seeJe frrax ur asoql searaqm'pasradsrp y9 y lynleru '(I8 = N :uxtt 6'LBZ uearu) ruru /'8I€-t'91; -er;ur aq deru spare ralslotu ur suorlepdod'* 00€ lnoq? ol qfua1p1o1'(uru g'fOt ueau) uru 6yt-916 TAS'(3 6'0: Ie^al eas ruo{ suorlelela }e s}saJoJ puBl.&\ol Jalsroru o1 sdorc ueaw) 8 9'L9-9'L€. (ZS = lf) sseur.(poq SulFlr1er{ '.rate1 s.irp -1no -,(>po.r paleuruop-snlJpJ pue lsaroJuroql rrJex uorJ l8-€8 par{r}eq aun{ 97 palrsodap s33a 9I'ralel sl.ep 19 qrrru Sur8ue,r splrqeq snorre^:(q7gg7 'pna.rg) puno;3 aql.reau ro o1 ue8aq eunl 6I palrsodap s33a 71'ruru €8 x 19 s33a'a1or: uo,tels o1.{1a4t eroru selrua n('puelsr pue 1elrqeq,{q sar.re.r. rarJ olul ro p^rr eq] le pues Suqra pue'3urqse1 Ie1 'Surtrc 1nq '(slsaro; prunq ur ru 0€ ol) patoqre d1?rtuessE :lBlIqeH '3ugnq'sa8ualpqc eprspeorq'3ur11or peaq epnpu srapru: -ur 1surc3e sasuaJep'uorlrsodap rage sdep 0I-I salrs lsau Lrl 'GZOOZ 'pnarg) anbrur1re141 uo ru 007 01 Ie^el urnlar daql uar{.tr selerual raqlo lsure8e seleruay dq pap:err": eas 'a8ur; Ieuorlelele :(866I'ore^r6) orrg o1ran4 uo punoJ eJB slsau 'seluulsrunrJll Jepun dep auo lnoqe sa.rrnt' uaaq seg I?nphrpur ue iauan&r ouon81 alrselur r{llm uoll Ieapr -ar 3u1ra.Loc pue '3url.e1 33a 'uorle-redard aloq '.pateratrlqo -ezrprrqdq ot ..]sol,, eq deu (nOZ "V 1a Inalg :eperrs?C eT are aloq eqlJo se8rlsa peaq rar.l rlllm 1l pue errel elrlad ]ou ]nq '{selrllales eluos pu? '.OrJaI-epuBJC Ip Irlun ..Suluruer_ ueql 'louunl aql olul sqrurlaroJ r{}l^{ Surqsnd pue alou 'a.r-ra1-asseg) adnolapeng pue (LOOZ 'lrnarg pue p.re1pe1q) }l e)uerlua aql olq Suntrnl dq taqrueqr 33a aqt lou lnc. anbrurlreyq uo esoqlpue'(9902'{'lB ]a{ruqlese4) peledrrga los 'sa8essed salBruaJ's33a pallaqs-Uos sazrs q]lnp uaeq a Bq o] ruadde elueleg-errery puu ' ;gg . aepruen6l ,(1rue1 132 ' Part ll. Reptilia, Section l. Squamata (lizards) Anguilla, largely restricted to thick scrub and clifs along tion, and dewlap extension, rare fights involve biting and the northern coast, often arboreal, but will readily use low tail lashing (Powell et al., 2005); on Dominica, male "tasted" vegetation or rocks, burrows in loose soil or naturally oc- iguana scat and appeared to scent-trail another individual, curring rock crevices and caves, not associated with hu- subsequently squatted by elevating its hindlegs and tail, ad- man-modified habitats (Hodge et al., 2003); most abundant pressing vent and thighs against the substrate, probably on St. Eustatius in apparently inhospitable scrub, often as- leaving scent marker immediately adjacent to the scat (|. S. sociated with White C edars (Tab eb ui a h et er o phylla), wher e Parmerlee, |r., pers. comm.). Competition: On Terre-de- threats are minimal, uses trees, shrubs, rocks, and readily Haut, apparently displaced by I. iguana (Schardt, 1998b; comes to ground, in gardens readily uses non-native plants Breuil, 2002b). Foraging and Diet: Food includes fruits of for food and shelter, will seek refuge in exotic Razor Plants Prickly Pear (Opuntia),bird eggs, and ca r riot (Lazell, 197 3) ; (Pandanus sp.), not uncommon on porches and roofs of on La D6sirade, eats leaves, flowers, and grasses, sometimes buildings where fed or at least not molested, may sleep in grazing alongside goats, a juvenile attempted to catch a trees, often on exposed branches, or in burrows (Powell et cricket (Schardt, 1998a); diet includes 28 species ofplants, al., 2005); where habitat permits, sleeps almost exclusively especially Tab eb ui a p alli da, Hipp o m an e m an cin ell a, Clero - high in trees (Breuil, 2002b). d en dr o n acule at um, C ap p ar i s Jle xuo s a, C. c y n ophallopho ra, Pisonia fragans, Guajacum oficinale, Bursera simaruba, Activity: Diurnal, sleeps in trees or rock crevices, associ- Conocarpus erecta (Barre etal.,l997); may consume flowers ated with rock piles or clifffaces, much of day spent in ther- and leaves of more than 50 species of trees and bushes in moregulation or foraging (Lazell, 1973); main activity dur- xeric coastal lowlands of Dominica, where iguanas may be ing early morning (-0830 h) and afternoon (-1500 h) on La important seed dispersal agents (Malhotra and Thorpe, D6sirade (Schardt, 1998a); on Anguilla in fune (N = 5), 1999); forages mostly in the morning, frequently associated 86.0% of time not in burrows basking on rocks, 6.57o bask- wlth Ipomea tiliacea, on which it avidly feeds, generalist ing in trees, 0.2o/o feeding,4.0% nonreproductive interac- vegetarian, diet varies by habitat and island, but may in- tions,2.0% courtship and mating, 1.8% roaming,and0.4o/o clude introduced species (e.g., ornamentals), on St.-Barth6- responding to an observer (Pasachnik et a1., 2002); some lemy, feeds on leaves (especially during the dry season, and individuals may become active at dawn or remain active favoring young leaves) and fruits (during the rainy season) until dusk, but most activity occurs from -0900 h to mid- to of Melicoccus bijugatus, Cordia dentata, Leucaena leuco- late afternoon, although some individuals may seek shelter cephala, Tabebuia pallida, Randia aculeata, consumption of during the heat ofthe day (Powell et a1.,2005). Behavior: fruits and seeds may facilitate dissemination of these spe- May bipedal and swim (Lazell, 1973); may interact with cies, some food items are toxic to birds and mammals, conspecifics at distances of l0 m, responds to disturbance drinks by licking leaves or opening mouth with head ele- by climbing higher into a tree or jumping to the ground vated during rains, may "taste' food before eating (Breuil, (Schardt, 1998a); extremely alert while basking on rocks, 2002b); on Anguilla, particularly fond of leaves, flowers, flees by running across litter-covered ground or jumping to and fruits of Chink B:ush (Bourreria succulenta) and flow- trees below the rocks (Ten16n, 1997b); quite capable ofde- ers of White Cedar (Tabebuia heterophylla; Hodge et al., fending themselves by biting and scratching with their 2003); readily consumes ornamental plants, and appears to powerful claws (Wijffels, 1997a); may congregate on fruit- exhibit a particular fondness for Hibiscus sp. on Dominica filled trees, but usually only one dominant male present (Daniells et al., 2008); on St. Eustatius, individuals may (Breuil, 2002b); on St.-Barth6lemy, iguanas easily tamed "gorgd' (consume all vegetation within reach before mov- when fed, readily enter houses, sunbathe and copulate on a ing) or "grazd' (take only a few bites before moving), fre- deck adjacent to a house (Breuil, 2003); exceedingly wary quently feeding on Tamarind (Tamarindus indica) leaves on Anguilla, relies on crlpsis, but often drops to the ground (S. P. Fogarty, unpublished data); on Dominica, female ate from elevated perches when approached at considerable dry dog food in a bowl (E. A. Daniells and N. |. V6lez Es- distances and flees quickly (Hodge et al., 2003); on St. Eu- pinet, pers. comm.). Home Range and Territoriality: On statius, readily tamed when fed, if disturbed may 'dive" Dominica, male territeries during the breeding season from arboreal or rocky perches, crashing to the ground, 1,000 m'z, female territories 2,000 m2, female ranges overlap and moving considerable.distances at high speed through those of other females and sometimes several males, sug- thick, often thorn-encrusted vegetation, headbobbing used gesting a female dominance hierarchy (Day et al., 2000; by both males and females, sometimes at substantial dis- Breuil, 2002b); males probably defend territories outside of tances, but aggressive displays, almost always between the reproductive season (Breuil, 2002b). Hurricane Ef- males, involve lateral compression, push-ups, crest erec- fects: Maybe adapted to habitats subjected to periodic hur- -png uo :(qZ00Z '.lrnerg ur /79I 'uoter8 '6) saqreaq uo pues -oe; 3ur1nur1 eq] qllm '{lalueq3 uo teql ploJ-gl lnoqe arlel ur uoilrsodap 'qc1n1c7s33a gI ol :(0002 ''p 1e deq) d1a41 alrled uo dlrsuap :(/002 ' ggg aepruen6l ,(;rLue3 134 . Part ll. Reptilia, Section l. Squamata (lizards) deloupe, mate in March and lay on beaches in May, accom- tius, maintains above ambient temperatures through the panied by males (J.-8. Du Tertre, 1667 in Breuil, 2002b); on night even when sleeping on exposed perches (RP). Dominica, females may move 460-900 m to egg-deposi- Conservation Status: Most populations remain extremely tion sites (Day et a1., 2000); on St.-Barth6lemy and Petite vulnerable to human exploitation for food, habitat altera- Terre, some females lay eggs within their home ranges, tion, introduced predators and competitors, and hybrid- (Breuil, while others move considerable distances 2000c); ization with introduc ed lguana iguana (e.g., Breuil, 2002b, nests in sand -1 m deep ending in a chamber large enough 2009; Powell, 2004e; Powell and Henderson, 2005a; Pow- nest to allow the female to turn around, in rocky areas, ell et al., 2005; Maillard and Breuil, 2007); on Chancel, an chambers less extensive (Breuil, 2002b); on ilet Chancel, additional threat is posed by tourists who trample one of suitable nesting sites are restricted and females often exca- two principal communal nesting sites, although recent vate eggs laid by other females (Bach and Schonecker, improvements (including fencing) have reduced the risk 2008); on Martinique, clutch size 20-30 (Bouton, 1630 in from tourists and sheep, also a proposal to relocate igua- ( -B. Du Tertte, 1667 Breuil, 2002b) ; on Guadeloupe, 13-25 J. nas from Chancel to ilet-ir-Ramiers may provide another in Breuil, 2002b); on Dominica and Martinique, 15-30 refuge from possible interbreeding with invasive I. iguana (Anonyme De Carpentras, 1618-1620 in Breuil, 2002b); on (Breuil, 2009); some populations (e.g., Dominica) are af- Dominica, 8-18, strongly correlated with female size, aver- forded protection under local laws (|ames, 2004); the spe- a length age weight 25 g (Day et al., 2000); to 22 eggs with cies is listed on CITES Appendix I (UNEP-WCMC,2008); (by of44 mm (Schardt, 1998a); freshly unearthed other fe- on the IUCN Red List as "vulnerable" (Breuil and Day, incuba- males) eggs 17-22 g,45 x 25 mm (Breuil, 2002b); 1996), but many individual populations meet criteria for tion 3 months (Day et a1., 2000); captive hatchling 20 g, 'tritically endangered" status (Breuil,2002; Fogarty et al., SVL77 mm (Reichling, 2000); hatchlings 17.6'18.9 g, SVL 2004; Powell, 2004a; Powell and Henderson, 2005a); Powell (Gibson, hatchling sizes vary by 80-83 mm 2001); egg and Q00ad and Powell and Henderson (2005a) recommended size is re- island and habitat (Breuil, 2002b); because clutch a reevaluation and a probable upgrade to "endangeredl' lated to female size, the smaller size of iguanas on Chancel Additional Pertinent Literature: Pasachnik et al. (2006). (compared to Petite Terre) limits recruitment into the pop- ulation (Breuil, 2009). Sex Ratio (M:F): 1:1.3 (Maillard and Breuil, 2007); 1:1 to 1:7 (Day et a1., 2000) or even l:12 on St.- l guano iguana Linnaeus,'1758 Barth6lemy (1.-C. Plassais in Breuil, 2002b). Size: Male SVL to 434 mm, female SVL to 401 mm (Breuil, 2002b); iguanas Distribution: Mainland from Mexico to Brazil, including on Chancel are smaller than those on Petite Terre (Breuil, numerous continental islands; introduced into southern oC 2009). Swimming: Male with body temperature 28 Florida, Hawaii, and Israel; in the West Indies, on Gre- jumped into water 25-26 "C, swam on the surface for 80 nada and the Grenadines, St. Vincent, Barbados, St. Lu- sec, then dove for 16 sec at a depth of 2-3 m, when emerged, cia, Martinique, Marie-Galante, Les iles des Saintes, Gua- floated perpendicular to water's surface, before climbing deloupe (Grande-Terre, Basse-Terre, and Iles de Ia Petite oC, onto a boat, within 2 min body temperature rose to 29 Terre), Montserrat, Antigua, Barbuda, Saba, St.-Martin/St. 6 minutes later dove back into the water, held his breath for Maarten, Anguilla, St. Croix (may have been introduced 28 min while immobile at 3 m at26 "C, swam on the surface by Caribs for food; Beatty in Grant, 1937b), Virgin Islands, for 6 min (M. Breuil and B. Thi6bot in Breuil, 2002b; Breuil, Puerto Rico (including Culebra), Grand Cayman, and Swan 2003). Thermal Biology: Position themselves to facilitate Islands; elevational range, sea level to -800 m; population effective thermoregulation (Lazell, 1973); mean body tem- on Barbados apparently extirpated (although some cap- peratures on La D6sirade 37.98 "C (36.53-38'77 "C) at sites tive animals remain), those on Martinique, Marie-Galante, with temperatures 30.5-31.0 "C at 0800 h, 41.0-42.2 "C at Les iles des Saintes, Guadeloupe, and St. Croix possibly midday (but in shaded sites where iguanas are most fre- introduced or of mixed origin (e.g., the historic popula- most currently quently seen -28-31 'C), -34-35 'C at 1500 h, dropping to tion on St. Croix presumably natural, but 30.3-31.0 oC in late afternoon and 28.5-29.0 "C at 1800 h, present lizards probably are descendants ofintroduced in- nighttime temperatures 24.5-25.0 "C (Schardt, 1998a); he- dividuals), and those on Antigua, Barbuda, St.-Martin/St. liothermic, males warm at exposed arboreal sleeping sites, Maarten, Anguilla, Virgin Islands, Puerto Rico, and Grand females often bask on the ground, when foraging often Cayman are introduced, although at least some individu- shuttle in and out of direct sun, highest temperature re- als on Anguilla arrived via natural over-water dispersal, oC, (Censky 1998); corded for a male digesting food 39.4 rest in shade dur- almost certainly from Guadeloupe et al., (and ing the hottest parts ofthe day (Breuil, 2002b);on St. Eusta- populations on St. Lucia, Saba, and Montserrat pos- 'puplsl rale^^ vo sttuutroqod sn\douaTtog :(0g6I .1errn3) :(qZtq.luerg) s{ror ol Surlr pue eas aq} o}ur a rp plno.e. suerunq pue (snlua,to[ salsadng) sasoo8uotu ,sBuenBI dq ualea ' on V. Rios in Joglar, 2005b); egg deposition (Grant, 1932h); on Carriacou (Grena- Gonzillezand U.S. Virgin Islands (April-May)' favored have a Terre-de-Haut at end of dry season dines), borallus grenadensis (Pendlebury' 1974); sites along beaches among clumps of vegetation' eating and their flesh is reputed to nesting "reputation as excellent (Schardt' 1998b); Ben- large rlumb.rs of hatchlings in October tasie like chicken' (Long, 1974); dogs (on St' Lucia; considerable distances to two known nesting probably mongooses (Morton' 2007); other -u:y -o,r. don, 2003) and moved 80 m' 1'57 km' and Hawks beaches on St. Lucia, 3 females than humans and domestic animals, Broad-winged nesting sites back to home ranges' 5 nests on main predators (Malhotra and 1.61 km from (Buteo platypterus) arethe on nest- (Falco St. Lucia wlthlT-2leggs (mean 19'5)' hatchlings Thorpe, 1999); presumed predators include Kestrels mid-May-mid-August (Graham and Morton' Hawks (Buteo jamaicensls)' some ing beaches spariarius), Red-tailed (14-28) in 14 nests with dogs' Zio+,zoOs);average clutch size 23 large racers (Akophis spp'), and introduced seabirds, 2OO7)' Sizet Adult SVL to 2005); on 637o hatching success (Morton, mongooses, and humans (Powell et al'' cats, rats, 1991); recent hatchlings capltred aitve' 500 mm (Scliwartz and Henderson, Grand Cayman, a Great Egr et (Ardea alba) (mean (October on Terre-de-Haut) 82-95 mm SVL 88 (T. Ebanks, in litt., 15'XII'2008); on Grand Cay- nile iguana g; Schardt, 1998b)' Thermal (T' in litt" 23'II'2009)' mm) and 13.22 g(mean 19.4 Crbophis caymanus Ebanks' thermo- Biology: Position themselves to facilitate effectlve ^un, Females mature at -3 years' later in males Reproduction: frequented by iguanas Feb- .. glrlutio., (Lazell, 197 3) ; in habitats must defend territories, males highly territorial who oC between 0930 and 1030 h' peaking to egg-laying sites in late teirperatures of 34.3 ruary-March, females migrate 1500 h and at 1215-1330 h, decreasing to 35'7 "C at female (890 g) with 17 eggs 38-45'5 x at ai.g "C Aprii-early May, (Schardt' 1998b)' in- to a minimum of 24.0 "C at night 22-28.2 mm, 14-24 g, largest eggs in distal oviducts' corre- (UNEP- cubation 90-92 days, hatchlings ]uly-early August' Conservation Status: Listed in CITES Appendix II 2002b); eggs countries' sponding to onset ofthe rainy season (Breuil' WCMC, 2008), but export quotas exist for many in old charcoal (leather luid ir, *u.-, well-drained sites on beaches' primarily for live animals (pet trade) or products hatched in native versus pits, and gardens (Breuil, 2000a, 2000c); 17 eggs goods and meat), no distinction is made for lays i+ *".t ,iurrderwood, 1962;Lotg,1974); presumably introduced or for continental versus insular populations; female 275 mm of laws eggs December-February, on Montserrat' hunting is usually prohibited, but enforcement on 24March game animal in SVL with 16 eggs (42.0-46.8 x 26'6-28'4 mm) is lax at best, the species is considered a may not ap- season (Lazell,l973);on Puerto Rico (although all data Grenada and St. Vincent, with a regulated hunting season No- ignored ply to Puerto Rican populations), reproductive and bag limits (on Grenada), which are routinely which vember-December, males establish territories from by local hunters (Powell, 2004e; Powell and Henderson' to the "greatly reduced in other males are actively expelled, courtship similar 2005a,2007); population on St' Croix depends on of man as by aggressive displays between males, copulation numbers (probably as much by the activity rarely copu- 1962); pre' females response, may last 15 min, but males that of the mongoose)" (Seaman and Randall' require high Islands late more than once/day, eggs (17-68/clutch) sumably introduced populations in the U'S' Virgin or on beaches' Code Title 12 incubation temperatures, nests in clearings are afforded special Protection under USVI on may be ugg..gut.d or dispersed, apparently depending (Platenberg, 2007). be used for several years' may be 1 m tlpe of soil, nests may populations currently as- sev- Remarks: Native West Indian urrd connected by tunnels to 20 m in length with i..p may rePresent more than one spe- may be com- signed to lguana iguana eral openings to the surface, egg deposition (e.g., 2007)' clutch sizes 12-90 (A' cies StePhen, -rrrul (Ri r..o, 1998); in Puerto Rico'