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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 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
Distribution: Two historically recognized subspecies in
the Turks and Caicos Islands (C. c. carinata) and Booby
Cay (east of Mayaguana Island, Bahamas; C. c. bartschi), al-
though subspecific recognition of C. c. bartschi is not war-
ranted (Stephen, 2005, 2006; Bryan el al, 2007).
Habitat: Xerophilic (Schwartz and Henderson, 1991); 1o-
cally abundant on smaller islands with herbaceous to shrub
stage growth (Schwartz and Carey, 1977); on Booby Cay,
iguanas on sandy and rocky beaches, rocky areas, sparsely
and densely vegetated areas, and open areas, used sandy
areas and holes between rocks for burrows, in bushes and
trees to heights of 1.8 m (Wasilewski, 1998); most abundant
in rocky coppice and sandy strand habitats, sandy areas are
required for nesting (Gerber and Iverson, 2000).
Activity: Diurnal activity bimodal during warmer months,
high midday temperatures result in reduced activity, winter
activity reduced and primarily at midday, basks following
emergence and prior to termination of day's activitl', rest
of day spent feeding, interacting with other lizards, and
thermoregulating, which involves shuttling in sun/shade
114
\-
mosaic (Iverson, 1979). Behavior: Performs headbobs to
declare territories, challenge males, and for sex recognition
(Iverson, 1979); most headbob displays consist ofa series oi
alternating single and double headbobs, each display usu-
ally beginning with 1-2 single headbobs or a fast "trill" of
6-15 bobs, followed by a set of 1-6 double headbobs, fol-
lowed by single headbob or two, head nods and tail raises
typical of other iguanas also occur (Martins and Lamont.
1998); aggressive displays include lateral orientation. gap-
ing, gular expansion, arched back, crest elevation, inflated
trunk, lateral compression, and single or double headbobs.
aggression also may involve tail lashing and fighting (Car-
penter, 1982); will take to water as an escape tactic, but onlr'
as a last resort (Iverson,1979). Diet: Primarily herbivorous
at all ages (95.6Vo of all food items and 94.8o/o of volume:
Auffenberg, 1982); feeds both terrestrially and arboreallr-.
600/o of macroplant species (including a few toxic forms
eaten, with plants taken falling into three categories: (i
Common staples of high caloric content, (2) commor.
staples of low caloric value (roughage), (3) foods eaten
frequently in spite of their uncommonness (plants of hig:
preference levels), certain individual plants are repeatedi''

browsed whereas others of the same species are ignored,
caloric content in the form of nonstructural carbohydrates
is not an important factor in leaf and fruit choice, animal
prey includes crabs (Clibanarius arrd Cardisoma), insects
(termites, beetle larvae and adults, cicada nymph, dipteran
larvae), a slug, and carrion (fish, lizard, bird, rodent, and
canned meats, especially sardines), during the winter, re-
stricted to items that are more difficult to digest (leaves),
rvhich may pass through the digestive tract almost intact
(Iverson, 1982); cannibalism (Iverson, 1979; Aufenberg,
1982); one well-developed circular valve and 4 more dis-
tal semilunar valves partition the proximal colon (N = 15),
presumably to increase digestive efficiency (Iverson, 1980);
truits of Turk's Cap Cactus (Melocactus azureus; S6nchez
Mufloz, 2008). Home Range and Territoriality: Adult
males are territorial throughout the year (Gerber and Iver-
son, 2000); average home range 980 m2 in females, 1,260
m2 in males, 1,590 m'zin dominant males (Iverson, 1979).
Parasites: Nematodes (Alaeuris travassosi, Ozolaimus
monhystera, Paralaeuris cyclurae; Dosse, 1939); very large
oxyurid nematode (Cyrtosomum mega) populations in co-
Ion, perhaps mutualistic rather than parasitic (Bowie and
Fratz, 1974); to >15,000 presumably commensal or even
mutualistic nematodes in a healthy adult (Iverson,l9B2).
Physiological Demands: Glucocorticoids (used to evalu-
ate heightened physiological demands under varying con-
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
in the morning, affected by reproductive state and possibly
by seasonal rainfall (MacDonald et al., 2007). Population
Density: Estimated densities along flush transects on pine
Cay 1.7 -3 6.8 adults/ha ( Iverson, 197 B) ; 9 0.3 I ha
habitat for juveniles, 31.1/ha for adults (Iverson, 1979);558
iguanas in 14.5/ha on Booby Cay, Mayaguana (Conners et
a1., 2005, 2006). Population Sizes: Estimated 50,000 C. c.
;arinata and 1,000 C. c. bartschi (Alberts, 2004); estimated
30,000 C. c. carinata and 200-300 C. c. bartschi (Hudson
and Alberts, 2004). Predators: Feral dogs and cats (Iver-
son, 1978, 1979; Mitchell et al., 2002; Mitchell, 2003; Ger-
)er and Alberts, 2005,2006). Reproduction: Cycles
with climatic cycles, courtship in early May,
'r'nchronized
:robably serial polygyny with monogyny among some
nales, eggs 2-glclltch in early |une, females defend bur-
:ows for several days to several weeks after nesting, but are
:rot territorial at other times, hatching occurs after approxi-
nately 90 days, hatchlin gs averuge79.8 mm SVL and 14.6 g,
'uvenile growth rate averaged 19.2mmSVL/yr, males reach
sexual maturity at approximately 220 mm SVL, 375-475 g,
rge about 7 years, females 185-200 mm, 200-300 g, 6-7
;ears (Iverson, 1979); individuals in recently established
Family lguanidae . i15
populations on French, Six Hills East, Bay, and Middle cays
exhibit larger average adult sizes and reach maturity more
rapidly (1.5-2.5 years versus 6-7 years) than on source
cays, Big Ambergris and Little Water (Gerber and Alberts,
2005, 2006). Sex Ratio: l:1 at all age classes (Iverson, 1979).
Size: Adult male SVL to 360 mm, female SVL to 292 mm
(Schwartz and Henderson, l99l). Survivorship and Life
Expectancy: Survivorship positively correlated with body
size, life expectancy 14.0 years (Iverson, 1979).TailAutot-
omyz32.60/o of males and28.60/o of females on Pine Cayhad
broken tails (Iverson, 1979).
Conservation Status: The species is listed in CITES Ap-
pendix I (UNEP-WCMC, 2008); 'tritically endangered" on
the IUCN Red List (Gerber, 1996a,2004b;Gerber and West
Indian Iguana Specialist Group, 2000); both C. c. carinata
and C. c. bartschi as "threatened" on the U.S. List of En-
dangered and Threatened Wildlife and Plants (U.S. Fish &
Wildlife Service, 2008) primarily due to predation by in-
troduced mammals (dogs and cats), competition with and
degradation ofhabitat by introduced herbivores (e.g., cattle
and goats), habitat alteration by human activity, and exploi-
tation by humans; a conservation and management plan
exists (Burton and Bloxom, 2007a); domestic dogs and cats
reduced the population (estimated 5,500 animals) on pine
Cay almost to extirpation in 3 years following construction
of a hotel and tourist facility (Iverson, 1978).
Cycl u ra col I ei Gr ay, 1845
Distribution: )amaica, Hellshire Hills (extirpated elsewhere
on the island, including Goat and Little Goat islands).
in optimum Habitat: Xerophilic, tropical dry forest in low limestone
hills along shores and in areas between the coast and in-
terior uplands, in hollow trees in pastures, where congre-
gated (Schwartz and Henderson, 1991); retreat site fidelity
is strong, with hatchlings preferring dead hollow trees 6-15
cm in diameter with a northwesterly aspect (Van Veen and
Wilson, 2004, 2005);17 of 155 observations of headstarted
iguanas released into the wild were in trees to heights of -7
m (D. S. Lewis et a1.,2008).
are
Behavior: Will climb, can climb a smooth fence post (C. B.
Lewis, 1944); headbob displays consist ofa single headbob
or three single headbobs with pauses of 0.7-1.0 sec between
them (Martins and Lamont, 1998). Foraging and Diet:
Leaves, fruits, and flowers of a wide variety of plant spe-
cies, supplemented occasionally by animal matter, includ-
ing snails (P. Vogel,2000a); of 25 types of plants consumed,
3-4 species of vines are most important (Van Veen and
Wilson, 2004,2005); foraging in tree canopies (D. S. Lewis
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-
erably, probably reflecting recency of release (D. S. Lewis
et a1., 2008); injuries presumably from fighting in released
iguanas are indicative ofterritorial behavior (R. van Veen
in D. S. Lewis et a1., 2008). Movements: Farthest distance
traveled from release site (N = 6) 196-455 m (D. S. Lewis et
a1., 2008). Population Size: Estimated 150 (Alberts, 2004);
estimated 100-200 (Hudson and Alberts, 2004). Preda-
tors: Hatchling consumed by Epicrates subJlavus (Wilson
and Van Veen, 2005, 2006); introduced predators include
mongooses (Herpestes javanicus), rats, cats, dogs, and pos-
sibly pigs (C. B. Lewis, 1944;P. Vogel, 2000a; Wilson et a1.,
2004a,2004b); dogs used for hunting pigs are able to take
even adult iguanas (Woodley, 1980). Reproduction: Age at
maturity 7-8 years in captivity (R. Hudson, pers' comm.
to j. B. Iverson); nesting activities "apparently late March
to early May" on Goat Island in the 1930s
1944); females begin digging trial holes long before egg
l*yrrg, eggs deposited in nests in two known communal
nesting areas in mid-|une, with hatchlings emerging 85-87
days later, females aggressively guard nests up to 2 weeks
after egg laying, using threat displays, biting, and chas-
ing, clutch sizes average 17 eggs (16-20), hatching success
ranges 0-100% and appears to be related to female body
size (P. Vogel, 1994);in 2005, ovideposition ranged from24
May to 20 |une for 14 nests in two known communal nest-
ing sites (Wilson and Van Veen, 2005, 2006). Size: Adult
male SVL to 428 mm, female SVL to 378 mm (Schwartz
and Henderson, 1991). Survival: Eight of 20 headstarted
iguanas survived 24 months or more (Wilson et a1., 2004a);
all 6 repatriated iguanas survived at least 3.5 months (D. S.
Lewis et al., 2008). Weight Loss: Over at least 3.5 months
after release, 6 headstarted iguanas lost an average of 1.69
g/day (D. S. Lewis et al., 2008).
Conservation Status: The species is listed in CITES Ap-
pendix I (UNEP-WCMC, 2008); 'tritically endangered"
in the IUCN Red List (Gibson, 1996b); 'tndangered" on
the U.S. List of Endangered and Threatened Wildlife and
Plants (U.S. Fish & Wildlife Service, 2008); principal causes
include degradation of habitat by charcoal burners and
predation by introduced mongooses (Herp estes j avanicus),
dogs, and cats (Woodley,1980; P. Yogel,1992; Alberts,1993;
P. Vogel et al., 1996; Wilson et al., 2004a, 200ab); ongoing
threats include proposed development of the Hellshire
Hills (Woodley,1980; P. Vogel, 2000a).
Cyclura cornuto Bonnaterre, 1789
Distribution: Hispaniola, widespread, but disjunct distri-
bution on the main island (probably once islandwide in
xeric lowlands), also Isla Beata, ile-de-la-Petite GonAve,
ile-de-1a-Tortue, ile Grande Cayemite, and possibly ile-de-
la-Petite Cayemite and Isla Saona.
(Schwartz and Hen-
Habitat Xerophilic to semimesophilic
derson, 1991); dry forest with sand, coral, or eroded lime-
stone substrates in coastal terraces, main-island lowlands,
and satellite islands, often using crevices as retreats or dig-
ging burrows in sand or fossil coral (Mertens' 1939; Carey,
1975; Noble, 1923c; Ottenwalder, 2000a); basks on roads
(RP); roadkitl recorded in 1939 (Mertens, 1939); readily
climbs into trees to feed (Powell et al., 2002).
Behavior: May take to water as an escape tactic (Noble,
1923c); individual basking in full sun on an angled tree
trunk 2 m above ground escaped into Opuntia thicket
when disturbed (Mertens, 1939); head "rolls" and headbobs
(C' B. Lewis, used in displays (Martins and Lamont, 1998); often become
"tame" with feeding or regular exposure to humans (Ban-
bury et a1., 2000; Banbury and Ramos, 2005). Diet: Feeds
on "harsh fruits and shore crabs]' responded to bait com-
posed of mashed sweet potato soaked in rum (Eyerdam,
1954); seeds of Ziziphus rignoni germinated more rapidly
after being consumed and passed by iguanas (Hartley et
al., 2000); dermatophagy (Weldon et al., 1993); a variable
number of circular (3-4) and more distal semilunar valves
(5-7) partition the proximal colon, presumably to increase
digestive effrciency (Iverson, 1980). Longevity: In captiv-
rty,22years,11 months and still alive at last report in 1990
(R. Noegel in Slavens and Slavens,2003). Parasites: Include
ticks Amblyomma albopictum (Carey, 1975; Wiewandt,
1977; Guglielmone et al., 2003), A. cruciferum (Neumann,
1901; Morel, 1967), and Ornithodoros elongatus (probably,
found in container with an iguana; Kohls et al., 1965; Mo-
rel, 1967; Guglielmone et a1., 2003); nematodes (Atractis
cruciata; Linstow, 1902). Population Size and Density:
0.03 iguanas/ha east of Pedernales in 1985 (Inch6ustegui,
2003b); 0.70 iguanas/ha on Isla Cabritos (G. Santana in
Inch6ustegui, 2003b); estimated 17,000 (Alberts, 2004); es-
timated 10,000-17,000 (Hudson and Alberts, 2004). Popu-
lation Structure: )uveniles compose about one-third of the
population on Isla Beata (Grupo laragua in Wiewandt and
Garcia, 2000). Predators: Humans (primarily for food),
feral dogs and cats, mongooses (Herpestes javanicus), and
pigs take adults, juveniles, and eggs (Ottenwalder, 2000a);
feral dogs (Rupp et al., 2008). Reproduction: Age at matu-
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-
gregate nesting on Isla Beata (Noble, 1923c); eggs in two
clutches (in captivity at the San Diego Zoo) averaged 68.'l
x 46.7 mm and 79.8 g (N = 21) and 67.6 x 48.6 mm and
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118 ' Part ll. Reptilia, Section 1' Squamata (lizards)

Staphylococcu.s spp. and Salmonella spp''

and 4with Entero-
the prox-
semilunar valves in C. c. figginsi (N = 14) partition the normal
efficiency ,orru'r rpp.lare probably commensals or part of
imal colon, presumably to increase digestive
(C' c' inornata; fames et
in rocks enteric microbial flora, respectively
(Iverson, 1SSO); drink rainwater from depressions C' c' cychlura and
Both source (Leaf al., 2006); ticks (Amblyomma dissimileon
iWrt.o* et a1.,1973).Genetic Diversity: A. torrei ot C. c. figginsi; Durden and Knapp'
2005)' Popu-
populations of C' c'
Cay) and translocated (Alligator Cay) Exuma
and hetero- lation Density -3.2lha(C'c'figginsion Guana Cay'
z
inornataexhibited low average allelic diversity South Andros
loci known to be polymorphic Islands; Carcy, 1976); <0'2 iguanas/ha on
zygosity for 8 microsatellite Size: Population
Growth: Indi- (Knapp and Owens, 2008a)' Population
in-C. iychlura (Knapp and Malone' 2003)' (Knapp'
SVL/yr over 3 years o., Gr'ru.tu Cay (Exuma Islands)' 64 individuals
vidual C. figginsi grew 8'33 mm
c.
1995); estimated total population sizes
3'500 C' c' cychlura'
(Coenen, 1995); mean annual growth rate significantly 1,000 C' c' inornata (Alberts' 2004);
(1'139 cm/yr; 1,500 C. c. figginsi,and
faster in males (1.764 cm/yr) than in females C''' fg-
1989); annual growth estimated 2,500-5,000 C' c' cychlura' 1'000-2'000
C. c. inornatal Iverson and Mamula' (Hudson and Alberts'
c' inornata on Alli- ginsi, and 400-500 C. c' figginsi
rates in an introduced population of C' from -150 in 1970 to "Per-
(35'3 mm and 30'1 1OOS), C. c. inornataincreased
gator Cay for both males and females Predation: Hatch-
greater th.an for the frupr'; r,:oo in 2006 (Iverson, 2oo7b)'
irm, respectively) were significantly (Knapp and
second-year C' c' Ii.rg, .onrrr- ed by Epicrates striatus fowleri
founde. population (Knapp, 2001a); one and by Cubophis
cm Owens, 2004; Knapp et al', 2004b' 2005)
pu.trl.. Cay (Exuma Islands) had SVI' =13'7 gape-limited
figginsio., vudii (Knapp et al., 2004b, 2005); apparently
2004b' 2005)' Home
ui"a Uoay mass = 118 g (Knapp et a1'' (Knapp
C. vudii fult.a i" predation attemPts on iguanas
Range: :O.SS hu fo, males and 5'63 ha for females' the larg-
(e'g'' Catesby' 1743;
(Knapp and Owens' and Owens, 2008b); eaten by humans
est r"eported for free-ranging Cyclura 2004b' 2005); hu-
inornata higher on U Barbour,1904; Knapp,2005; Knapp et al''
2005). tife History: Survival of C' c'
dogs and possi-
activity)' higher mans (for food), feral pigs (on nests)' feral
Cay than on Leaf Cay (with more human et al" 1999;
"bolder")' and exhibited bly cats on Andros Island (C' c' cychlura;Knapp
in iemales than males (which are (for illegal wildlife
presumably Buckner and Blair' 2000); humans
declining trend as growth rates neared zero'
a
or at carrying capacity trade), dogs, and possibly rats (C' c' figginsi;"1upn' 2000b)'
because tliese populations are near
Reproduiion: Males "guard" females on Andros (C' c' cy-
juvenile C' c' inornata captured
(Iverson et al., )OoOa) ; of 16
to 150 m' males
at age 12'5 years chiura),attendant male chasing an intruder
and marked at age 0.5 years, 9 were alive lateral head-
95'3o/o' of 4 ofequal size engage in displays, face-offs' and
(typical age at maturity), an annualize d rate of
still alive 10 p.,rht.,g(Knapp and Owens, 2008a); copulation on Andros
oii..;ouJrrl.s marked at age 1'5 years' 3 were
Owens' 2008a) and
recorded on 24 April (P. Hartman in Knapp and
y.u.,late., an annual rate o 97 '2o/o (highest rates and owens' 2008a); nest on
extreme zs Aprif (A. Mitchell in Knapp
for juveniles of any lizard), life table demonstrates with damp sand on 6
Guana Cay (C. c. figginsi) covered
longevity, very high survival throughout life'
greatly de-
driftwood apparently placed by a
for any studied lizard)' extreme )une, entrance blocked by
layJd maturity (longer than on Guana Cay
output' net reproductive nest-guarding female (Knapp, 1995); nests
iteropariry and low reproductive with females actively de-
-61 ; long and 8-13 cm deep'
."" ='3.4) and cohtrt generation time (21 years) match
i*. fending incomplete tunnels but not nests
after ovideposi-
between
the 3.2"-fold population size increase estimated laid in termite (primarily Nasu-
20 tion (inapp, zOoOa); eggs
1982 and 2004 (Iverson ,2OO7a)'Longevity: May exceed and defended
and Mamula' 1989); 36'7 years titermes rippertii) rnot"td' on Andros Island
years in nature (Iverson (C' c' cychlura; Knapp'
capture) fot C' c' inornata in by females on tops of termitaria
iadding estimated age at first 2004,2005;Knapp and Owens, 2008a; Knapp
et a1'' 1999'
C' c' figginsi'
natu.ellverson et a1., 200ab); in captivity for in coarse sand
report in 1990 (R' 2004b,2005, 2006); two nests on Andros
23 year's,6 months and still alive at last (Batillaria.sp') shell frag-
Death due composed largely of gastropod
Noegel in Slavens and Slavens,2003)' Mortality: often used
(Iverson et al" 2005); menis, both nests in iermitaria and elsewhere
to bing wedged between tree trunks used again by same female)'
repeatedly (6 of 12 in termitaria
: indivi"dualslrapped in 50-gallon plastic drum (Iverson'
a mostly
(Exuma Islands)' major ,r.rti.rg began in eaily May through mid-June'
2008). Nest Sites: On Guana Cay shrubland
fine' windblown (460/o) inopen pine followed by dry evergreen
nesting area on beach characterized by
tussocks of grass (C' (367o) closed pine habitats (18o/o)' mean canoPy cover
and
sand with numerous small palms and
Nematode Afrac- 72 x ZOVo,eggs deposited at tunnel entrances and pushed
Coenen, 1972inKnapp, 1995)' Parasites: nests
nematodes. in 4 of 6 fecal into centrai chambers with head and forelimbs'
tis opeatura(Leidy, tAgO); oxyurid sand' nest tem-
(6 plugged and covered with scattered soil or
.u-pl., bacterial isolates in all but one of 17 animals
and (29'2 !
5 with coagulase-negative p.r"ul.., (mean 32.6 I 1'5 "C) higher than ambient
w\ti Corynebacterium spp',
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120 . Part ll. Reptilia, Section 1. Squamata (lizards)
Cyclura lewisi Grant, 1940
Distribution: Grand Cayman Island, once islandwide.
Habitat: Holes in limestone (C. B. Lewis, 1940); in sandy
and rocky scrub (Schwartz and Carey,1977); currently ex-
ploits a mosaic of natural and somewhat disturbed habitats
in response to availability of thermoregulatory sites, food,
substrates suitable for nesting, predation pressure, and hu-
man activity (Burton, 2000) ; captive-bred animals released
into the wild "preferred' modified habitats over natural
habitats and used artificial retreats more frequently than
natural retreats (Goodman et al., 2005a).
Activity: Iguanas emerged earlier and retreated later in
summer months than in the fall, most of the time inactive,
usually in states of low or medium alertness, of the small
percent of time males spent active (20.3 ! 2.3o/o summer,
19.6 ! 3.Oo/o fall), they moved (8.3 ! 4.2o/o,6.4 ! 2.5o/o), for-
aged (8.1 ! 4.8o/o, lO.2 r 2.2%), and engaged in social situ-
ations (11.4 x 2l.lo/o, 1.9 ! 3.2o/o), much inactive time spent
basking (29.9 t 9.2o/o,23.5 ! 4.4o/o), females active (18.3 +
1.7 o/o, 16.9 t 4. 4o/o), movin g (6.1 t l. 4o/o, 5.7 t 2. 4o/o), f
(9.0 ! 2.5o/o,8.3 t 3.4o/o), engaged in social situations (20.7
t15.2o/o,12.7 t20.3o/o), basking (23.5 x 4.2o/o,23.9 19.3o/o),
in general, iguanas spend little time in trees or bushes or
inside retreats during the day (Goodmary2007). Behavior:
Produces an "S-roll" (the nose is raised and lowered as in
a headbob, but is also moved in a horizontal plane during
the lowering motion), although some individuals appear to
produce standard headbobs in addition to the S-rolls, the
twisting motion is not as dramatic as the full rolling mo-
tion produced by C. cornuta and C. stejnegerl, displays are
series of -5 S-rolls separated by pauses of -0.7 sec, a pat-
tern very similar to that of C. nubila from Cuba (Martins
and Lamont, 1998); adult males show little long-term fidel-
ity to retreats, but females appear to occupy small, fixed
territories centered on 1-2 favored retreats, the number of
suitable refugia may control female distribution, whereas
male density may be regulated primarily by territorial in-
teractions (Burton, 2000); hatchlings tongue-touched the
ground as they moved away from nests, 3 of 5 climbed into
trees within 20 min, suggesting greater arboreality than in
adults (Goodman and Burton, 2005); extensive tongue-
touching of retreats, substrates, and feces suggests the
importance of chemosensation, supplemental feeding ap-
peared to increase incidents of aggression toward humans,
iguanas responded to increasing levels of human and ve-
hicular disturbance (Goodman, 2007). Diet: Plums are
preferred over all other fruit, mangoes are taken in season,
and pawpaws are eaten by captive animals (C. B' Lewis,
1940); almost exclusively herbivorous, known to consume
at least 45 species of plants in 24 families (-807o leaves and
20% fruits), particularly important are leaves of Rhynchosia
minima, Chamaecrista nictitans, and Stylosanthes hamata,
whole plants of Spermacoce confusa, and seedlings of Wal-
theria indica, all of which are weeds that thrive in disturbed
open areas, iguanas gorge on fruits of Picrodendron bac-
catum, Hippomane mancinella, and Myrcianthes fragrans
when available, these occur primarily in stands of primary
vegetation, the diet is presumably very different than it was
prior to human settlement, exceptions to strict herbivory
include consumption of fungi, crabs, and cicadas, with the
latter presumably scavenged, coprophagy and soil inges-
tion have been observed (Burton, 2000); slugs (Veronicella
sp.; Goodman, 2003, 2007); primarily herbivorous, eating
plants, the majority of which were noncultivated, but ge-
ophagy, coprophagy (including that of an agotti, Dasy-
procta punctafa), consumption of invertebrates (climbing
into tree to feed on larval moths, Pseudosphinx tetrio) oc'
curs, one iguana chased low-flying butterflies for -5 m and
attempted to catch them, consumes shed skin and debris
(plastic spoon, plastic bags with food scraps, metal twist-tie,
o t aging with efforts to climb into trash cans), supplemental feeding
contributed little to the overall diet, but did result in ag-
gregate feeding of females and juveniles, but never males
(Goodman, 2007). Home Range: Released, head-started
females had mean home ranges of 0.24 ha, released males
0.65 ha with considerable overlap, for a density of 4-6 igua-
nas/ha, attributable largely to an unnatural single age class
in an area surrounded by unpopulated habitat (Burton,
2005a, 2005b,2006); mean home ranges of males in sum-
!
mer (13.567 14.426 ha using minimum convex polygon
and 4.559 t using 95% contours for fixed kernel
3.517 ha
use distributions) were larger than in fall (1.600 + 1.656 ha
and 0.804 t 1.102 ha) and much larger than that of females
in summer (0.623 ! 0.312 ha and 0.209 + 0'181ha) and fal1
(0.856 1 0.754 ha and 0.467 x 0.486 ha), with males moving
longer distances in summer than in fall and than females in
either season (Goodman et al., 2005b). Hybridization:
Cyclura nubila caymanensis will hybridize wlth C. lewisi
if "brought together" (Davis, 2000; Burton, 2004c). Lon-
gevity: In captivity, at least 54 years, estimated -69 years
(Adams, 2004). Mortality: Seven iguanas "murdered" b1'
humans at the Queen Elizabeth II Botanic Park (Binns,
2008a; S. Hicks,2008). Parasites: Piroplasm Sauroplasma
in erlthrocl'tes of captive-reared juveniles (Alberts et a1.,
1998). Population Density: Density of head-started females
1.6-2.0lha (Burton, 2005a). Population Size: Estimated 30
in nature (Alberts, 2004); estimated 100-175 total (Hudson
and Alberts, 2004). Predators: Wild and domestic cats and
dogs, and humans, who perceive iguanas as a threat to their
crops (Burton, 2000, 2007); Cubophis caymanus preys on
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122 ' Paft ll. Reptilia, Section l. Squamata (lizards)
sun is highest (Buide, 1951); active 0730-0830 h (tempera-
twe = 27 oC) on clear days, does not emerge from dens on
rainy days or days when cloudy early, basks until 1000 h'
then feeds (1000-1300 h), rests in shade before resuming
activity after 1600 h, retires to dens by 1730 h, juveniles re-
treat just after 1200 h, almost always before 1400 h (on the
Sister Isles; Grant, 1940c); active throughout most of the
day, emerges on clear days when the temperature reaches
-27 'C, usually around 0830 h, with height of activity oc-
curring between 1000 and 1300 h, rarely seen in the open
after about 1430 h, resumes activity around 1600 h, not seen
afterl73O h (Cayman Brac; Carey, 1966); on Little Cayman,
begins to emerge -1000 h, most active during the hottest
part of the day (Townson, 1980); at Guantanamo Bay, vigor-
ous bursts of social activity follow emergence, relatively
peaceful migration to nearby areas of vegetation for mid-
day feeding, ending in a second bout ofsocial interactions
in late afternoon prior to entering nighttime refuges (Al-
berts, 2003). Behavior: Initially reacts by inflating its body
and "featuring a ferocious aspectl' but quickly learns to take
food ofered by hand (Gundlach, 1880); very shy upon
emergence from dens, but "tame" easily in captivity (on Sis-
ter Isles; Grant, 1940c); readily climbs smooth fence posts
(C. B. Lewis, 1944); when disturbed, rarely covers the entire
distance to refuge in one dash, instead uses short spurts of
14-18 m, during pauses "like a dog on the alertl' stretching
its head upward and bobbing, when cornered 'bpens its
mouth in threatening fashion and gives voice to a discon-
certing hoarse noisej' readily climbs stone walls and tall
trees, huddles together on cool nights (Buide, 1951); flees
when encountered at substantial distances, retires to thick-
ets in which they are concealed, sometimes seen in groups
of 4-5 (on Cayo Largo, Cuba), elsewhere usually only a
single individual at a time (L. H. Saxe in Sutcliffe, 1952);
younger individuals will bipedal when fleeing, if captured,
whip tails (Z.Yogel,1965a); males responded to approaches
by headbobbing and broadside challenge displays, interac-
tions involved largest male in a population on Little Cay-
man chasing the largest female, which, in turn, chased
other females, t hatchlings were solitary (Townson, 1980);
losers in male-male agonistic encounters flee considerable
distances (-7 m), stopping at intervals to look back at the
threat, eventually taking refuge in a burrow in sand or un-
der a rock, the encounter itself involves the peculiar pitch-
ing ofthe head, gaping, and circling the opponent, collec-
tively comprising ritualized behaviors that circumvent
actual fighting in many instances (Berovides Alvarez, 1980);
produces an "S-roll" (the nose is raised and lowered as in a
headbob, but is also moved in a horizontal plane during the
lowering motion), although some individuals appear to
produce standard headbobs in addition to the S-rolls, the
twisting motion is not as dramatic as the full rolling motion
produced by C . cornuta and C. stejnegeri, displays are series
of -5 S-rolls separated by pauses of -0.7 sec, headbobs pro-
duced by animals in the Puerto Rican population lasted
more than twice as long as those from Cuba or by C.lewisi
(Martins and Lamont, 1998); agonistic male-male interac-
tions consist of "lumbering" chases, occasionally with pro-
longed pushing, hissing, and biting matches, dominant
males are larger, have higher testosterone levels and more
scent glands (Alberts, 2003); dense aggregations in high
anthropogenic usage areas exhibited more aggressive be-
havior and social interactions than individuals in low usage
areas (C. n. nubila at the U.S. Naval Base at Guantanamo
Bay; Lacy and Martins, 2003); aggressive displays include
face-offs, lowered head, gaping, crest elevation, and head-
bobs or head shaking, aggression may involve biting, tail
lashing, chasing, and fighting (Carpenter, 1982); will enter
the sea to escape threats (Hardy, 1956); juveniles may enter
water and hide among seaweeds (Thalassia testudinum;Ro'
driguez Schettino, 1999b); on Cayman Brac, iguanas cross
large open areas when disturbed instead ofrunning into a
mangrove swamp (Carey, 1966); on Cayos San Felipe, seeks
refuge in burrows when chased by dogs trained to catch ju-
tias (Capromys), but fight tenaciously (attacking dogs and
men) when cornered (Varona and Garrido, 1970). Biomass:
Adult biomass -11.7 kg/ha (Cayo Sijri; Beovides-Casas and
Mancina, 2006). Burrows: Burrows under bushes, esPe-
cially Tournefortia gnaphaloides and Suriana maritima (C.
n. caymanensis; C. B. Lewis, 1940); -1.2 m long and -30 cm
beneath surface (Grant, 1940c); may occupy crab burrows
(Neilt, 1958); an adult male resided in a small cave on Cay-
man Brac (Carey, 1966); on Little Cayman, lizard wedged in
one corner of a small (10-30 cm high, <1 m long) chamber
(Seidel and Franz, 1994); on Cayo Sesenta Arroba, many (7
of 19) burrows face southeast, on Cayo del Rosario, most
(10 of 16) face south, mean width and height of burrows
15.94L2.62cmand 10.50 + 1.34 cm, mean length and depth
147 .64 t t 15.89 cm, mean distance be-
59.42 cm and 40.56
+
tween burrows 15.04 8.56 m, in September, burrow occu-
pancy 60.0%, with two iguanas in only one burrow in Oc-
tober, occupancy 79.60/o, with 8 of 19 burrows housing two
or more iguanas, with 8 iguanas in one burrow, burrow
density on Cayo Sesenta Arroba 7.7 lha (Cubillas HernSn-
dez and Berovides Akarez,199l); on Cayo del Rosario, 13.7
burrows/ha (Berovides Alvarez,l998);also on Cayo del Ro-
sario, mean widths and heights of burrow entrances 21.58 I
7.48 cm and 9.04 + 2.76 cm, mean distance between bur-
rows 18.37 r15.82 m, mean density of burrows 14.90 + 9.70
burrows/ha (GonzSlez Rossell et al., 2001); mean burrou'
entrance heights and widths 7.62 xl.6l cm and 13.33 ! 4.02
cm on Cayo Obispo and 6.05 + 2.99 cm and 14.43 t 5.59
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124 ' Parl ll. Reptilia, Section 1. Squamata (lizards)
contained Balantidium (ciliated protozoan) and nematode
eggs, "healy nematode infestations were found in the large
bowel of all of the iguanas" (L. H. Saxe in Sutcliffe, 1952);
intestinal nematodes Atractis opeatura (P6rez Vigueras'
1935b; Baru$ and Coy Otero, 1969a; Baru5 et al., 1969; Coy
Otero, 1970; Coy Otero and Baru5, 1979b; Coy Otero and
Lorenzo Hern5ndez, 1982; Coy Otero et al., 1987; Coy
Otero, 1999); O swaldofilaria brev icaudata, Paralaeur is c y-
clurae, Alaeuris travassosi (Barui and Coy Otero, 1969a;
Barui et a1-., 1969; Coy Otero, 1970; Coy Otero and Baru5,
1979b; Coy Otero and LorenzoHernlndez, 1982; Coy Otero
et al., 1987; Coy Otero, 1999); Oswaldocruzia lenteixeirai
(Coy Otero and Baru5, 1979b; Coy Otero and Lorenzo
Hernindez, 1982; Coy Otero, 1999); and Ozolaimus mon-
hystera (P6rez Vigueras, 1936; Baru5 and Coy Otero, 1969a;
Baru5 et a1., 1969; Coy Otero, l97O; Coy Otero and Baru5,
1979b; Coy Otero and Loren zoHernindez, 1982; Coy Otero
et al., 1987; Coy Otero, 1999); oxluroid nematodes in free-
ranging juveniles at Guantanamo Bay (Alberts et al', 1998);
500/oof large intestinal volume filled with nematodes (Ro-
driguez Schettino, 1999b); nematodes presumably com-
mensal or even mutualistic (Iverson, 1982); piroplasm Sau-
roplasma in er1'throcytes of 7lo/o of 1993 captive hatchlings,
but none in 1994, and 79o/o of free-ranging juveniles and
>60% of cells of 26o/o of the same free-ranging lizards were
infected with the haemogregarine Hepatozoon and hemo-
coccidian Schellackia (Alberts et al., 1998). Population
Density: Densities declining in many areas due to human
activities and, in some areas, hunting, mean density 9'6
iguanas/ha on Cayo del Rosario (Rodriguez Schettino,
1999b); on Cayo del Rosario, 3 individuals/ha (Berovides
irlv ar ez, 198 0) ; 9. 9 t 0. 65 individuals/ha (6.5'12.8 iguanas/
ha in different months; Gonz6lez Rossell et a1.,2001); densi-
ties on 11 Cuban cayos 9.3-30.0 individuals/ha, with high-
est densities (25.0, 30.0, and 30.0/ha) on rocky islets and
the lowest density (9.3 iguanas/ha) on sandy Cayo del Ro-
sario, where estimates based on numbers of burrows or
distances between burrows were even lower (Berovides A1-
var ez, t99});densities t atged 22.5-25.2 lha ot't Cayo Obispo
and 10.5-18.2/ha on Cayo Verde in 1998-2000, and was
14.2lhaon Cayo Sotavento in 1998 (Berovides Alvatez etal',
2003); densities higher in wet than dry seasons and differed
by habitat, with density in xerophltic coastal scrub (6'7 +
6.3lha) higher than in sand vegetation (3.6 ! 2.7lha) ar,d
mangrove forests (2.9 x 2.9lha),with the most juveniles in
the coastal scrub (Cayo Sijir; Beovides-Casas and Mancina,
2006); estimated 40 animals on Cayman Brac (C. n' cayma-
nensis) in 1993 (Gerber, 1996b); -1,800 iguanas on Little
Cayman (Goetz,2003); -500 iguanas on7.2-ha Isla Ma-
gueyes, Puerto Rico (Garcia, 2005b, 2006b). Population
Size: Estimated 40,000 C. n. nubila and 1,500 C. n' cayma-
nensis (Alberts, 2004); estimated 40'000-60,000 C. n. nu-
bila and 1,000 C. n. caymanensis (Hudson and Alberts,
2004); estimat ed 422 igaanas (50-75/ha) on Isla Magueyes,
Puerto Rico (Alvarez et a1.,2007); estimated 40 C. n' cay-
manensis on Cayman Brac (F. |. Burton in Goetz, 2008)'
Predators: On Little Cayman and Cayman Brac, dogs (C'
B. Lewis, 1940); a very young iguana under artificial condi-
tions panicked in the presence of Cubophis fuscicauda, sug-
gesting that these snakes regularly prey on hatchling igua-
nas (Carey, 1966); potential predators of eggs on Cayo del
Rosario include landcrabs, hermit crabs (Coenobita clypea-
fus), Greater Antillean Grackles (Quiscalus niger), guJls
(Larus atricilla), and Royal Terns (Sterna maxima), w\th
the three birds also potentially preying on juvenile iguanas
(Berovides Alvarez, 1980); feral and domestic dogs (on
adults andjuveniles), cats (onjuveniles), and pigs (on eggs;
Estrada, 1993b); stuffed and sold as souvenirs to tourists in
Cuba (M. eerny,l992); humans are not a major factor be-
cause consumption of iguana meat or eggs is not common
(C. n. nubila; Perera, 2000); in some areas, are hunted for
food (Fong Grillo et al., 2005a);both iguanas and eggs were
considered a delicacy on Cayman Brac (Carey, 1966); feral
cats, domestic dogs, and possibly rats (C. n. caymanensis;
Gerber, 2000a); feral cats and humans (in the form of road
mortality) appear to be the major threats on Little Cayman
(Goetz, 2008). Reproduction: Courtship probably begins
in April and egg deposition from April to )une, often in ar-
eas where iguanas do not normally occur, eggs 69'0-77'8
mm in length, clutch of 9 eggs 67.8-75.5 mm length (Ro-
driguez Schettino, 1999b); reach maturity at 2-3 years otr
age (Perera, 2000); on Little Cayman Island, courtship and
mating occur in April and early May, females dig nest holes
and deposit eggs between 1730 and 1800 h from May to
early fune in beach sand to within -9 m of high tide mark,
2 females contained 8 and 20 eggs (C. B. Lewis, 1940); ma-
turityat 983 g body mass on Isla Magueyes, courtship be-
gan in late March or early April, females apparently move
to males during the mating season, once a female responds
to restricts her movements to the immediate vi-
a maie, he
cinity of the burrow for up to 7 weeks (post-copulatorl'
guarding?), if she attempts to leave, he circles her while vig-
orously displaying, several burrows initiated before one
completed, usually in sandy or grassy areas, often in sides
of embankments, nests reused in consecutive years, bur-
rows >1 m long and <50 cm deep, females extremely secre-
tive and do not defend nest vigorously, eggs 2-14 (mean
5.6) with mean mass 46.4 g, incubation 70-78 days, mean
SVL of hatchlings 107 mm, major source of egg mortality is
ant predation (Christian, 1987); clutch of freshly laid eggs
(in captivity) 82 x 42 mm, hatchling SVL 84 mm, 13'4 g (l'
Schmidt, 1994); on Isla Magueyes, nest depths 26-70 cm'
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126 ' Part ll. Reptilia, Section 1. Squamata (lizards)
Activity: Usually emerges from dens around 0700 h (mean
ambient temperatures -25 "C), moves to sunny areas and
assumes a prostrate basking posture, after about I h (mean
ambient temperatures -27 "C), begins to search for food,
stopping frequently to sample small bites of nearby plants,
basking continues after feeding, but rarely in full sun (usu-
ally a sun-shade mosaic), during midafternoons (mean
ambient temperatures -30 'C), usually remains in full or
partial shade, after -1500 h, when temperatures drop be-
low 30'C, moves into more exposed sites, no iguanas seen
after 1800 h (Carey, 1975). Behavior: Aggressive displays
involve stereotypical vertical headbobs (Carpenter, 1982);
juveniles engage in aggressive encounters with other ju-
veniles, these involve headbobs and pushups and, on one
occasion, biting, chases often followed by up to 10 min of
displays (Perry et al., 2003b); adults on Anegada extremely
shy, even basking adjacent to dense sheltering vegetation,
in contrast, hatchlings are quite naive (K. Bradley in Wer-
ning, 200 9). Diet Eats C o ccolob a \eav es, O p unt i a ft uJt' arrd
whole wild nutmegs (Grant, 1937b); primarily herbivorous,
but takes some animal matter, juveniles probably insectivo-
rous (Carey, 1972,1975);bulk ofdiet consists ofleaves and
fruits, composed largely offorage not taken by introduced
ungulates (3Oo/o Croton discolor, which contains com-
pounds toxic to ungulates, and >55o/o fruits of Byrs onima,
Coccoloba, Eleaodendron, and Eugenia, which livestock ig-
nores), but animal matter (beetles, caterpillars, centipedes,
roaches) forms about 170 of the natural diet (Mitchell, 1999,
2000a, 2000b); native plants taken on Guana include large
quantities of Centrosema virginiana, flowers of Tabebuia
heterophylla, seed pods of Capparis cynophallophora' and
leaves of Stigmophyllon emarginatum and Capparis Jlex'
uosa (Mitchell, 2000b)' Growth: Adulthood at7-9 years
and 350-400 mm SVL (Carey, 1975); estimated 50 mm
increase in SVL/1T based on 5 recaptures and 47 captive-
raised indMduals, maturity at 4-7 years, 37'8 cm SVL
(Gerber, 2000b). Home Range: Males 116.06-985'11 m'z (N
= 5), females 155.47-412.31m2, aside from food, available
shelter is the most important factor for selecting a home
range (Carey, 1972,1975); recent population structure dif-
fers, with home ranges expanding 100-fold with a decline
in population size, and male home ranges averaging 6'6 ha
(2.2-l2.3ha) and those of females 4.2 ha (2.8-5'6 ha), home
ranges broadly overlap and male centers of activity asso-
ciated with home ranges of females and those of females
with burrows (Mitchelt, 1999, 2000a, 2000b). Parasites:
Ticks (Amblyomma antillorum), with 20-30 (maximum
59) ticks per adult lizard, most frequently on the thorax
and in the region of the femoral pores (Kohls, 1969; Carey,
1975; Keirans, 1985; Guglielmone et al., 2003); piroplasm
Sauroplasma in erlthrocytes of captive-reared juveniles
(Alberts et al., 1998). Population Density: 2.03 iguanas/ha
(Carey, 1975); O.36lha in 1991, with the lower density due
to degraded forage attributable to competition with intro-
duced mammalian herbivores (Mitchell, 2000a). Popula-
tion Size: 200 wild adults and 44 repatriated iguanas on
Anegada, with another 80 in the headstart faciliry 130 on
Guana, and 30 on Necker (Perry and Mitchell, 2003; Ger-
ber,2004a; Bradley and Gerber, 2005; Lazell,2006; Perr,v
et al., 2007a); estimated 400 (Alberts, 2004); estimated
200 (Hudson and Alberts, 2004). Population Structure:
Adults (87.50lo of population) outnumber juveniles (12'5Yo)'
adults appear to be monogamous, living in separate but
closely adjacent burrows (Carey, 1972,1975). Predators:
Dogs (Grant, 1937b); feral dogs and cats (Mitchell, 2000a,
200b); humans (Wing, 2001; Lazell, 2005a); Borikenophis
portoricensisand American Kestrels lFalco sparvarlus) take
juveniles on Guana Island (Levering and Perry, 2003); also
on Guana, a 621-mm SVL, 88-g Borikenophis portoricensis
consumed a hatchling iguana of 40.4 g, a mass represent-
irg45.9o/oof the snake's pre-meal mass (RWH, RP). Repro-
duction: Cycle appears to be synchronized with two yearl1'
rainfall maxima, adult females with 12-16 yolked ovarian
follicles, Ioss of fecundity presumably due to competition
with domestic livestock (Carey, 1972,1975); mating season
on Anegada May-|une, females dig nests in sandy areas
from late |une through |uly in which they lay up to 20 eggs
(average 13), hatchlings emerge in the fall (Gerber, 2000b,
2004a; Bradley and Gerber, 2005); on Guana' egg laying
in |une-|uly and hatchlings emerge in September-Octo-
ber (Perry et al., 2007 a) . Sex Ratio: Equal with suggested
monogamy (Carey, 1975); 2:l (M:F), with "pairs" no lon-
ger clearly definable (Mitchell, 2000a, 2000b). Size: Male
SVL to 560 mm (Mitchell, 2000a), female SVL to 476 mm
(Schwartz and Henderson, 1991); mean hatchling SVL on
Guana 108 mm, iguanas of similar length on Guana weigh
more than those on Anegada, suggesting that the latter are
food-limited (Perry et al.,2O07a). Survival: 86% survival
of headstarted individuals released \n 2003-2004 (Gerber
et al., 2OO7). Tail Autotomy: 37.5o/o of males and 11.8olo
of females on Anegada had broken tails (Gerber, 2000b)'
Thermal Biology: Heliothermic, exhibits behavioral ther-
moregulation, cloacal temperatures of 16 active animals
oC (mean 6.3 "C) above air temperatures at
ranged 1.2-9.8
the time of capture (CateY' 1972) .
Conservation Status: The species is listed in CITES Ap-
pendix I (UNEP-WCMC,2008); "critically endangered" in
the IUCN Red List (Mitchelt, 1996); 'endangered' on the
U.S. List of Endangered and Threatened Wildlife and Plants
(U.S. Fish & Wildlife Service, 2008); principal causes in-
clude habitat degradation by introduced mammalian her-
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-dV SEI1C ur pelsrl sr sarcads aql :snlels uorluarasuo3 lapp.ruello :166I 'uosrapuaHpve zueilqts :l€z6l .alqo-\ -o
'(L002 .e
sluaurpas aurreru q8norqt 3ur11nt seurler pue spaq
laer:
:
ou
"p la errery :3 Otf ) turu 09, ot
.IAS aleuraJ
d.rp 'qr_
'(3 ooz'r) ruur s8t ot 'IAS eleul tlnpv :ez$,.(6l6t,le^no 3o opeq ur suorterlua)uo) qllm ,ollrnbrrug o8el r..
eror{s ureqtnos aqt Suop ;o por ,(q paralor sear. -3
:&
pue ortse) dalpeg :II = N .3 9.gg uearu) B laaer8
6.99_9.97 sseur tuo{ luasqe erB suap,elqelre^e
.ipoq ,(Ot ,Lu:uI }ou sr Iros aJarlM sartr-\p: lEt
pue = lf Z.VZZueeru) turu 9.L€Z_Z.ilZq$ua1
{ror pue s{unr} eerl Molloq asn IIr^r lnq ,$llol pasodsr il1}
lplo]'(II =N 'rrutu 7'19 uearu) tuur 0.16_r.28 145 Bunqrleq
'fI-8I (o8uruoq olues ur ooz ro 'sdrunls '<sqnrqs ,(so1r.rqe3 elsl uo oaoqtrw atyndotpt,: uo
Ieuorleu aql .uro6ooz w -y(3 ,lllertadsa) uorlela8aa ,(u,roqt esuap repun seruerlu:
.i1r.rr1der ur) azrs qltnp :(9002 ..p 1a ddng) raqtualdas 7
'I4
rlllm s-t{orrnq pa}e^elxa-Jles s,ra;a;d.solrrqe3 elsl uo se-(e1d 1i.
uo pa8raura Sumpreq :(SOOZ ..p 1a ddng) sBBa paqrleq
67 palsn:rua-]ps (lelI ur snorrnq .dys_ra,Lrp ]ue1d q8lq qtl,u srug <I
.lo profer e r{1r1r{
lsau auo ,(spos paurerp_lla,^{ ,daap qlln
palsaroJ .rol pue sluaurpJs aurreur dq pa.raproq slros .ie1: -lE:
speJe ,.'sopuoJ,, ur _(lalrsnlcxa lsorup) lrrqndag uef,rurruoC ,fuapa,lod pr,l'nlp dq pazrralrereqt stetrqer{ pturldo ,(i8u: jl-.'u
uratse.^.\qtnos aql ur salrs f le (eq/slseu
L.il_Z.6) peqtleq -urees qllrv\ (suorlenlrs pues pue >po,r >1ee1q Lre^ pue .qnlrs osP
stsau Jo o/ob'26-6'9g taqualda5_pnu q8notql oun{_pru anwv (qnns snloer ,.3.a) spuelpoom qnrrs urorll qll,ll patE er,i"l
Surqrleq 'aun{-pnu q8norql qore11 d1.rea Burlsau :(qOOOZ -rfosse.(1esop 'suorpnlrs prre d.ra,t ur ,trlrqdoray :lBtIqeH
':app.u,ua116) a8e
7o srea,{ p-7 1e lymleur qtrBor salerueJ 'iq
'(rlqndag uecruruoq) euoqpreg 'eco
'3 69 pue 'IAS ruru y'7g aien,re s8urlqcleq teqotro
pue ep Elnsuruad
ra./trol aql puE eqraN ap :srol
:aqualdag ur uoseas .(ure.r puocas aql alle^ .eloruedsr6 :uollnqlrlsr(l
Jo lasuo aql qllm tnq
pazruorqtuds sr Surqcleq ,s.(ep 991-96 slsel uorleqntur
'f" IE-08;o arnleradrual elqels e a^eq pue deap tuo /€B['uorqtB pue lu?unc llpou unpf) l'r!
67_ :aJ
are ter{} sreqr.uer{r 33a w (7.y 4 1.1y ueau) sBBa g1_7 de1
salpual ,(porrad.(urer,(pea aqtSo ]asuo eqt qlm pazluorql Petet
'(ptOOt .oueJoW),,paur .ura;
-u.is) aun{-,(ery ppl s33a .s1ros,(pues .aug ur salrs Bur1sa51
-teerr{},,:(enOZ.'p 1a .Lrra4) Sur.npql euen3 uo }Eqt qtr_ri
:uorlrnpo.rdag .(8002 ..p 1a ddng) s8op prag :(S002 ,.p -ra3
le '.palenlp^a eq o] luelar oo] are ro IIaM Burop aq o1 readde
uo 0{
Jdn6 :qgg67 taplB.etuello isasoo8uour .s1et .s8op) saro,r spuBIsI ur8rrl qsrlrrg oql ur aroq tesla suorlepdod paqsrl
-ruret pafnpo.rlur pue (acualsrsqns) suetunH :srolepeJd uo s
-gelsaer :(qL1oz,uoxolg pue uogng) slsrxa ueld dra,to:
' rr.)0a'suaqly -eInd
pue uospng) 000,r_000,2 pelEturlsa:(1002 -a.t serceds e:(SOOZ geqrag pue dape_rg :qOOOZ ,[aq]]ll\
'j:raqly) _erlu1
00€,I peteurpsg :ezIS uorlepdo4 .(VOOZ,.p ta epe8auy uo (s8op pue sler) sroleperd uur1eturueru parnp anp -
ser:1') sartads aqt ,(q palrqequr a.ra.tt uor8e.r eJrlua eql ul -orlur dq uorlepard pue (sorrnq ,streo8 ,aplec ..a.r)
saro.rrq Pq/ser
131' . aepruen6l ,(lrureg
128 ' Part ll. Reptilia, Section'1. Squamata (lizards)
et al. (2000) questioned the appropriateness ofsubsPecific
recognition, based on a lack ofhaplotypic differences, but
suggested that a final resolution should be determined by
analysis of more rapidly diverging nuclear markers, such
as microsatellitesl distribution once more widespread on
the main islands and presumably on cays currently without
populations (Hayes,2000a, 2000b; Hayes and Montanucci,
20oo).
Habitat Xerophilic to mesophilic (Schwartz and Hender-
son, 1991); exploits varied main island communities, plus
coastal rock, sand strand and sea oats, and coastal coppice
plant communities on offshore cays, and blacklands cop-
pice and mangrove communities on cays within inland
lakes, but most abundant in the vicinity of limestone out-
crops and/or patches of Sea Grape (Coccoloba uvifera) and,
on some cays, patches of Buttonwoo d (Conocarpus erectus),
presumably require sandy areas for nest construction (C' r'
r il ey i; Hay es, 200 0a) ; coastal rock habitats with Strumpfi a
maritima and Sea Grape, least abundant in dense Thatch
Palm (Thrinax moruisii) forest (C. r. qistata; Hayes, 200b)'
Behavior: C.yclura r. rileyi displays comprise series of up
to 7 single and double headbobs, mixed, with no obvious
pattern to the choice of single or double headbobs (Mar-
tins and Lamont, 1998); somewhat arboreal, climbing into
mangroves in cormorant (Phalacrocorar) rookeries on San
Salvador Island (C. r. rileyi; Paulson, 1966); males on cays
in Acklins Bight (C. r. nuchalis) engage in jousting matches
involving open-mouthed territorial displays, chase other
males out of defended areas, and bear scars of bite marks
(Hayes and Montanucci, 2000); aggressive displays involve
headbobs or pushups (Carpenter, 1982);larger males guard
females and smaller males may engage in forced copula-
tion (Hayes et al., 2004); will take to water as an escape
tactic when harassed (Hayes etaL,2004). Commensalism:
Hermit crabs in all and Sphaerodactylus corticola in some
iguana burrows (Gicca, 1980). Diet: Observed feeding only
on fruits of Sea Grapes (Coccoloba uvifera; Gicca, 1980); on
Green Cay (C. r. rileyi), consumes 7 species of plants, with
Silver Sea Oxeye (Borrichia arb orescens,flowers especially),
Wild Thyme (Rhachicallis americana, both flowers and
leaves), and Opuntia stricta (a preferred item despite scar-
city at the time of the survey) most commonly represented
in scat samples, nonplant items included a Purple Galli-
nule (Porphyrula martinica) and two unidentified song-
birds (probably scavenged) and one Bridled Tetn (Sterna
anaethetus) chick (likely predated), conspecific hatchlings
(N = 2), a dead land crab, a grasshopper (iguanas occasion-
ally seen lunging for grasshoppers), a hermit crab, uniden-
tified insect material (in 22o/o of scats), and sand and soil
(Hayes et al., 2004); dermatophagy (Weldon et al', 1993); a
single circular and 4 distal semilunar valves partition the
proximal colon (N = 2), presumably to increase digestive
efficiency (Iverson, 1980). Home Range and Territoriality:
Adult males territorial throughout the year (Hayes, 2000a;
Hayes and Montanucci, 2000); restricted activity to "what
appeared to be rather small territories within the vegeta-
tion' (Gicca, 1980); mean 9570 fixed kernel estimates of
home range sizes for males and females, respectively, on
Green Cay (C. r. rileyi) wete 439 and 628 m'z (S' Cyril, 2001
in Hayes et al., 2004); on North Cay (C. r. nuchalis), sizes
werc2,047 m2 for gravid females and 397 m2 for non-gravid
females (B. Thornton, 2000 in Hayes et al', 2004); on White
Cay (C. r. cristata),86% adaptive kernel estimates of home
ranges of 5 males and 2 females wete2,656 m'?(S. Fry' 2001
in Hayes et al., 2004); home range sizes reflect island sizes
and possibly population densities (Hayes et al., 2004). In-
juries: 16.0% of 75 C. r. cristata, 12.3o/o of 57,14.9o/o of 87,
and 20.4o/o of Sa C. r. nuchalis from Fish Cay, North Cay,
and a translocated population, and 33.3o/o of 3,34.5o/o of 35,
33.7o/o of 86,13.3o/o of 15,75.0o/o of 16,2l.lo/o of 19, and 51.67o
of 31 C. r.rileyifromGaulin, Goulding, Green, Guana, Lot',
Manhead, and Pigeon cays, respectively, had missing dig-
its,42.7o/o of 75 C. r. ffistata,24,60/0 of 57,32,60/o of 86, and
51.9o/o of 54 C. r. nuchalis from Fish Cay, North Cay, and a
translocated population, and 07o of 3,22.9o/o of 35,22.60/o of
84,40.0o/o of 15, 50.0% of 16,36.4o/o of 22, and64.50/o of 3l C'
r. rileyi from Gaulin, Goulding, Green, Guana, Low, Man-
head, and Pigeon cays, respectively, had injured tails, with
toe loss increasing with age and tail injuries more common
on rat-infested islands (Carter and Hayes, 2004). Longev-
ity: In captivity, 9 years, 2 months (D. Blody in Slavens and
Slavens, 2003). Mortality: Death due to being wedged be-
tween tree trunks (Iverson et al., 2005). Parasites: Ticks
only on Guana Cay (C. r. rileyi; Hayes, 2000a). Population
Size and Density: Estimated 500 C. r. rileyi' 200 C. r. cris-
tata, and 13,000 C. r. nuchalis (Alberts, 2004); estimated
500-1,000 C. r. rileyi,l50-200 C. r. cristata, and 15,000 C' r'
nuchalis (Hudson and Alberts, 2004); for C. r. rileyi, maxi'
mum population 35 on Green Cay (6 iguanas/ha), maxi-
mum population 28 on Man Head Cay (7.Slha), maximum
population 30 on Low Cay (2.5lha), all probably at or near
carrying capacity (Gicca, 1980); 300-400 C. r. nuchalis on
Bush Hill Cay (Iverson, 2007c). Population Structure: Per-
cent juveniles was l7o/o (White Cay; C. r. cristata),l-l0o/o
(mean 670, three cays ; C. r.. nu chali s), atd 0 -20o/o ( mean 8 7o'
six cays; C. r. rileyi; Hayes et a1.,2004). Predators: "Other
than man, the only likely natural predator[s] were sea gulls
(capable ofconsumingyoung) and ospreys (capable ofcon-
suming both young and adult iguanas)" (Gicca, 1980); Os-
preys (Pandion haliaetus) regularly prey on adult iguanas in
Acklins Bight, a Barracuda (Sphyraena barracuda) attacked
ro auo uaq^\ efuelorl lnoqllm Surpua puu .urtu g> 3ur1se1
-i11ensn 'pade8sqtnou qll^{ saqr}etu Surqsnd a)eJ-ol-ereJ
Sur,rloaur pue qlSuerts;o slsai pnlrr se Sularas slq8g qlyvr
'sarrolrrral 3ur1eu rol lq8g sapru 'frlfel adecsa ue se relem
ot e{el deur 'suolsnrlur uetuni{ o1 d1.re1n8e-r pasodxa uaq,tr
,,atuet,,euoraq uet lnq '.paqmlsp uaq.\r uap e olur 8urlea;1
-ar Lqs dlpnsn :(qWA,luerg) u.frurpqlpr s,(eup 1ou sr
IIel aqt .:e{ll-rl3u}sq, tnq '.aloq Ilor e olur Suruunr dq sgearl
-aU :rol^uqag '(erct'trct lpuerltar6) Suqseq tnoqqm
ro l{llm '}sar sr eurlnor Lpp pru:ou Jo 0/og6 lnoqe (}uatu
-arlnbar dlpp e 1ou sr Surpaa;'uooureu? alBI pup Sururou
ar{l q dllrerut.rd .,s1no1oo1,, pasodxe uo $lseq 'Lep aql
Surrnp pasn aqdeu pue sleerlar Ieurnlf,ou se elras (qBU *
S'0- raqueqr Furural qllm daap w ,- o1) suaq :,$r,u1ry
'(ZOOZ'teqeS pue o8erlrng-zar94) punor8 a,roqe
trt O'ZyZ'IslqBraq ]e saqrrad paleaala rar{lo ro searl ur arur}
.lpqes
Jo o/o€9 luads sarrolruat ur palpas s8ulgrleq :(0002
pue o8erlrng-zebd) slearlar se pesn arp qlrqm ,saloq{urs
,;o acuasa.rd aql rllp\ palelarrof, dla,r.rIsod lnq (slelrgeq 11e
ur luasard sgueld poo;) ,firsuap luu1d qty'r palelarrol lou
aruepunqr 'ruo8oqow aruapltuLs pue arlo!1asryba owtans
-rr3 Jo suorlelueld Lq pateldsrp uaaq seq uorlela8a,r a,n1eu
araqm puelsr ar{l Jo apls urelsa.&r arll uo .&tol 'nea1e1d aq1 uo
aterpauralur pue .ureld Ielseor er{l ul }se,lrol ...1e}lqeq.UIII,,
ur lsaq8rq eluepunqe :(966I .ore rU) ,(l1p8u ..pa1cadxaun,,
qtl-^{ soarl peulllul pue $l)or Surqurn;o alqedur slerurue
Sunod dlertadsa 1nq'leulsarral d1e:aua8 :(0002 .e;trpg
puE ]pue^\arM :6L61 'LL6I'1pue1t\arM :gZ6I ,lprwqls A
')) seaJe L4ro-r pue 'sale) 'sJrlr;o suor8ar puelur .slseof
.\ror Suop :(7961 :ra8eufa]S ur qsrp^\og) s4ro,r Euorue
:(166I 'uosrepuaH pue zlremqcg) cyqdorey :lellqeH
'aprmpuelsr Llerluassa,euol4l elsl :uoqnqlrlsr(I
9[6['alqoN pue lnoqleB ua^aufa]s unpi)
'(9697 'e*rras aJIIplt A ry qslC 'g'n) stuu14 pue aJII
.S
-plr6 pauat?erql pu? pa.re8uepuE;o ]sl.I n aql uo .,peua
-lperq],, se srlaq)nu 'r ') pue aJa$u) 't ') prre,.pa.ra8uepuq,
sE patsll sr r,{apt'r D:(qg66l .'p la .rage3) .,paraSuepua
.i11err1rr,, sr Q{apt'.r '3) ropeaps ues uo lur{} pue ,(ege1t
' 1e 1a ra1re3) .,pera8uepuq, aw (sqaqtnu 'r 'J) ]qBIg suppv
ur -ie3 qlrog pue qsrC uo esoql ,(qgOOl .sade11 pue ra1re3)
..pa:a8uepua d[e]r]rr),, s1 (a7o1s1tc't 2) tlea (,(pue5) a1rq11
uo uorlelndod aq] lnq .(eSOOt .sadeg pue rege3) ]sl.I peg
;1l aq] ul ,.para8uepua, :(8002 '3hl)M-d:INn) 1 xpuad
-\f
-Jv sgtll uI pelsll st sorcads aqr :snlels uorlelrasuoJ
'(eOOOZ'saLeg :9161 .,(a.re3 pue ztrre^tr{)S
::.'.altt 't'3) azrs dpoq uo sJureJlsuor leuorlrJlnu SurlsaSSns
'.l:rsra.\rp 1ue1d qtyur palelarroc Llelrlrsod are pue dec o1 Lur
u:or_i .ilqe:aprsuoc Lrezr. sezrs runrurxer.u pup ueeru t(I66I
57; aepruen6l i(lruueg
'uosrapueH pue zUeMqtS) ,xlil VgZ, ol .IAS alerual tlnpp
'ruur 909 ol'IAS eleru l1npv:az1g'(7967 ''p 1a sadeg :7gg;
''p 1a sa.(eg q 0002 'uolurorll 'g) ragolcA ro raqtualda5
ur s8ulqoleqyo aruaS.raua 'pesnal er? slroJtnq aruos lsai
-8ns sluau8er; 1aqs33a 'azrs r{l}nlr qtl^\ pelelarroc d1a-ru
-e8au ssetu 33a qtyn '(p = X) B y 17 pue (Et = ,rr) tutu z'of
x t'gg azrs 33a utaru 'azrs aptual r{ll.tt pe}ellosse sem osI?
pue (II N'o/oy'92 pa8era e luetulsalur a.r.rlrnporda: 'azrs
-
elerual qll.ryr patelerroc ,(la,\Ilrsod '(rt = lr '1'E uearu) g-;
azrs qrlnlr D, €e-gZ pa8uer se.rnleradrual lsau 'sreqr.upqr i
ur seqrlnlf, peurcluof lsau euo 'sJaqrueqf a1dr11nu urpluo-a
Leu pue uorllaJrp a8urqr ueuo sMoJJnq '<azrs alerual ql!-rr
dlaanrsod palelarror azrs qtllv' (uD ZZ ueau) daap rut or-H
,rro
sraqrueqr 33a pue (gt = l,I 6Zl uearu) 3uo1 uD gtz-69
smorrnq lsau 'asuaJap lseu alrlre ur pa8e8ue sepueJ fI Jo
g 'is>laam 7 o1 dn roJ sanurJuof pue uorlel?f,xa rege,{11roqs
surSaq uorlrsodapr,ro '(eq,tWZ) lellqer{ rolrelur,(ryor ur rsa
-.ttol pue '(eqgfi) lpllqeq dpues :orralur ur aterpaurelu
'(e\tZZg't) pues ]uo{qreaq ur lsaq8rq figsuap }sau .ralpl
Surlsau saletueJ re8rul qlpu 'eunl ur {aa.r\ }sel aqt Sugnp
Euruur8aq 3ur33rp $ou rlll,lr'lseu 01 uDI I ol dn a,rou ieu !
selerueJ 'aun{ dpea-dery alBI uorlelndor '3 g97 pue 115
rul 9'6I aletual arn1etu lseflptus 'snoue8l.lod Llqrssod .snou
-d8dlod ualsds Surleru 'stf)q)nu 1') roJ:(9961 ta8autar5
:r{ayt 't '3) s33a g qll^\ qr}np :(VOOZ ,'le sa.(eg :769;
}a
1a
''p sa.(eg ut I00Z 'llrtrf, 'S :eggg7 'sal.eg) sdep 76-16
-1o
seurl uorluqn)ur raUB
raqopo ro.raqrualda5 ur s8urlqtteq
;o arua8ratue '$laem Z o1 dn ro; sanurJuof pue uorlp^\sl E
-xe .reue dluoqs surSaq uorlrsodepr,to '.pesnal eJe smoJrnq
r
auos leql lsaS3ns sluau8e-ry 1aqs33a '.IAS rur g'g-Z'g prr"
_1r-r
3 9'19 salruazr,nf peqrleq ,tpuacar e '3 LV-ZV arua8rarua :r
eurl le s83a g 'ezrs qllnp qlyr pelelar.rol dlaz.rle8au sseru \
33a qtyur (rz = ru) 3 6 17 pue rnrl 9' 6Z x v'€9 s33a ueatu
a 's
'ezls alBIuaJ qllm pelellosse se.n pue (6 = N'o/oL'IZ pa8e:a-rr '-u
lueruJsalur a.r,rlcnporda.r .(s33a 91 qllm -(83 .no-I uo q)1np
auo) de3 uearg uo (S =,rf) g-tezrs qllnp '.saqllnlr Z paursl -u
-uor l.lluaredde raqtueqc 83a auo 'uorl)arrp a8ueqr uago sa
slrorrnq '(r;rrc ZZ uearu) daap url gZ-gI sraqueql 33a pue I
(9 = gg ueau) 3uo1 ruc 9II-09 suorrnq ]sau ,pareld
N'rur )
Llasolc eJe.tr slsau aJer.{.er seaJe ur uorutuoJ lsoru asua-Iap e
lseu qum 'sdep g ueql aroru roJ tueql papueJep pue lsau P!
aql fteu pauleruer €I pue esuaJep lsau a^rlre ur pa8e8ua sa:
saprual ZZlo g1'spor Sur8ueqra o qleeuaq slsau lsoru pup If
rarlrea Surlsau selerueJ ra8rel qllm .,{p{ ur >1aam lsry aqt 8rc uo
-rnp sur8aq uorlBlerxe tsau '(deyg ur {ea,n lsBI er{l Suunp
-IO
uoqelndor) dey41 ur rnro l.lqeqord Surletu pue drqsunor '; -EI
0t€ puB'IAS rul g'Iz s?,!t alBI,ueJ arnleu lsallErus aqt 'i.l2'1. Ir
' ro
r' )J :uollrnpo.rdag' (q0002'e6g6g'sadeg :sler'suoo: :eo
-re.r) sleurueur patnporlur pue (ape4 IBrurup lllllll tql :i3;
suerunq :(t002 ''p 1a sadeg :sqoq)nu'"r '3) leuueqc a.ro-d a-r!
-uerrr e ssorle Sururuvrrs euen8r llnpe ue peunsuol ptr? e$
130 ' Part ll. Reptilia, Section 1' Squamata (lizards)

ha (2.6-22.2 ha; Garcia et a1.,2007); males exhibited uni-

both contenders back slowly away, occasional battles last to
t h and may involve biting and tail lashing (Wiewandt'
1977); aggressive displays include lateral orientation' gap-
ing, dewlap extension, crest elevation, and headbobs or
head rolls (Carpenter, 1982); aggressive behavior may be
directed at humans (Tenl6n, 1997a); produces
"rolls" (one
form dispersion, whereas females were randomly distrib-
uted, mean male home range size 0.6 + 0.2 ha (0'21-0'81 ha;
minimum convex polygon) and 0.09 + 0'05 ha (0'02-0'15
ha; 507o kernal algorithm), mean female home range size
0.28 t
0.09 ha (0.13-0.34 ha, minimum convex polygon)

side of head is raised, when lowered, the opposite side is and 0.07 + 0.04 ha (0.03-0.13 ha;5Oo/o kernal algorithm)'
male - female overlap extensive (mean 86. 6%; 6'4 -10 0'0o/o)'
6
raised, often repeated in long series) as well as headbobs'
male-male overlap minimal (mean 2.9o/o;0-11%), suggest-
rolls occur at the beginning ofdisplays (usually) or at the
end, almost always accompanied by 1-2 series of standard
ing that males are highly territorial and that home ranges
.o.r.rpond to territories, ofhome ranges suggest that
sizes
headbobs, consisting of up to 6 (mean 2'O tO'42) up-and-
this may be partially responsible for low density if male car-
down motions per series (Martins and Lamont' 1998)'
Blindness: Blind iguanas (Haneke, 1995); 9 of 28 animals rying capacity is attained (P6rez-Buitrago et al', 2007b);
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
'bccasionally nas moved 471-6,396 m (mean 2,844 t 2,122 m), distances
et al., 2007a). Foraging and Diet: Descends
feed on the young corn and cotton' (K' traveled between consecutive monitoring days 56-157 m
to the tillable area to
(mean 76 + 33 m; Garcia et al.'Z}}7);hatchlings dispersed
P. Schmidt, 1928); consumes 71 of the 400 plants species
102-5,080 m from release site, moving on average -250 ml
known to occur on Mona, food includes fruits (especially
Opuntia spp. and Hippomane mancinella),leaves' flowers' day (P6rez-Buitrago and Sabat, 2007); another cohort of
headstarted iguanas moved 441-1,249 m (mean 633 ! 147
und ,o*. animal matter (especially seasonally abundant
m; P6rez-Buitrago et a1.,2008). Nesting: Plantations of in-
Pseudosphinx tetrio calerplllars and pupae), preferred
vasive Australian Pine (Casuarina equisetifulia) in nesting
plants are those low in cellulose, low in aromatic com-
sites produce accumulations of leaf litter and organic mat-
pounds, and easy to obtain, leaves (difficult to digest) may
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
"grazd'(moving from food item to food item), presumably Gatd, 1977); ticks (A. cruciferum) and nematodes (Wie-
wandt, 1977); tick loa ds (Amblyomma sp.) positively corre-
selecti.rg favored growth, or'gorge" (sit in one place and
eat everlthing within reach; RP). Growth: Estimated 52'3 lated with SVL, ticks aquired quickly after release from
mm SVL during first year, but one field-captured juvenile captivity (one iguana acquired 7 ticks in 4 days), loads of
recaptured headstarted iguanas (28.2 t 25'4 ticks) comPa-
grew 60 mm SVL in 0.7 years (85.7 mm/yr; Wiewandt'
rable to those of wild iguanas (32.2 t 22.0 ticks) of compa-
iSZz); at5 months, 3 individuals had gained no weight (av-
erage -1 t 2.6 g) and SVL increased 0.4 t 0'36 cm' fourth
rable size (P6rez-Buitrago et a1., 2008). Population Den-
individual gained 119 g and increased SVL 4'11 cm (P6rez- sity: 0.33 iguanas/ha (Wiewandt, 1977), btt may have

declined (Wiewandt and Garcia, 2000); 0'21 lizards/ha

(J'
Buitrago and Sabat,2007); growth of 9 wild iguanas 4'1-10'5
A. Moreno in P6rez-Buitrago and Sabat, 2000); 2'36 ! 3'3
mm/month (mean 6.3 + 2.2mmlmonth) and did not differ
iguanas/ha using line transects 16 m wide (P6rez-Buitrago
significantly from growth rates of headstarted iguanas' "natural" habi-
and Sabat, 2000); in study area ofrelatively
body condition (mass/SVL) declined after release to levels
tat 2.7 5 iguanas/ha (P6rez-Buitrago et al', 2007b)' Popula-
comparable with those of wild animals of comparable size
ana wita adults with SVL >400 mm (P6rez-Buitrago et al''
tion Size: 2,000-4,000 individuals (Wiewandt, 197 7 ) ; l'150
(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
berts, 2004). Population Structure: |uveniles only 5-1070
with good drainage, ventilation, nearly horizontal resting
places under low ceilings, and within m of surface' if
-lJ of population (Wiewandt, 1977) or 2 of 118 observations
to leave a maleb territory, he pursues her (Wiewandt and Garcia, 2000). Predators: Humans (his-
iemales attempt
and tries to "herd" her back (Wiewandt, 1977 ,1979); mean
torically), feral cats and pigs (Wiewandt, 1977l' Moteno'
Wiewandt and Garcia, 2000; Garcia et
home range size for t head-started iguanas 19'8 ha (2'4- 1995; Rivero, 1998;
a1., 2001; Bendon, 2004). Reproduction: Mating occurs
22.2ha;Garcia, 2OO5a,2OO6a); mean dispersal range of 8
radiotracked individuals (minimum convex polygon) 10'7 during an annual 2-week period, males lie in wait for fe-
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132 ' Part ll. Reptilia, Section l. Squamata (lizards)
Anguilla, largely restricted to thick scrub and clifs along
the northern coast, often arboreal, but will readily use low
vegetation or rocks, burrows in loose soil or naturally oc-
curring rock crevices and caves, not associated with hu-
man-modified habitats (Hodge et al., 2003); most abundant
on St. Eustatius in apparently inhospitable scrub, often as-
sociated with White C edars (Tab phylla), wher e
eb ui a h et er o
threats are minimal, uses trees, shrubs, rocks, and readily
comes to ground, in gardens readily uses non-native plants
for food and shelter, will seek refuge in exotic Razor Plants
(Pandanus sp.), not uncommon on porches and roofs of
buildings where fed or at least not molested, may sleep in
trees, often on exposed branches, or in burrows (Powell et
al., 2005); where habitat permits, sleeps almost exclusively
high in trees (Breuil, 2002b).
Activity: Diurnal, sleeps in trees or rock crevices, associ-
ated with rock piles or clifffaces, much of day spent in ther-
moregulation or foraging (Lazell, 1973); main activity dur-
ing early morning (-0830 h) and afternoon (-1500 h) on La
D6sirade (Schardt, 1998a); on Anguilla in fune (N = 5),
86.0% of time not in burrows basking on rocks, 6.57o bask-
ing in trees, 0.2o/o feeding,4.0% nonreproductive interac-
tions,2.0% courtship and mating, 1.8% roaming,and0.4o/o
responding to an observer (Pasachnik et a1., 2002); some
individuals may become active at dawn or remain active
until dusk, but most activity occurs from -0900 h to mid- to
late afternoon, although some individuals may seek shelter
during the heat ofthe day (Powell et a1.,2005). Behavior:
May bipedal and swim (Lazell, 1973); may interact with
conspecifics at distances of l0 m, responds to disturbance
by climbing higher into a tree or jumping to the ground
(Schardt, 1998a); extremely alert while basking on rocks,
flees by running across litter-covered ground or jumping to
trees below the rocks (Ten16n, 1997b); quite capable ofde-
fending themselves by biting and scratching with their
powerful claws (Wijffels, 1997a); may congregate on fruit-
filled trees, but usually only one dominant male present
(Breuil, 2002b); on St.-Barth6lemy, iguanas easily tamed
when fed, readily enter houses, sunbathe and copulate on a
deck adjacent to a house (Breuil, 2003); exceedingly wary
on Anguilla, relies on crlpsis, but often drops to the ground
from elevated perches when approached at considerable
distances and flees quickly (Hodge et al., 2003); on St. Eu-
statius, readily tamed when fed, if disturbed may 'dive"
from arboreal or rocky perches, crashing to the ground,
and moving considerable.distances at high speed through
thick, often thorn-encrusted vegetation, headbobbing used
by both males and females, sometimes at substantial dis-
tances, but aggressive displays, almost always between
males, involve lateral compression, push-ups, crest erec-
tion, and dewlap extension, rare fights involve biting and
tail lashing (Powell et al., 2005); on Dominica, male "tasted"
iguana scat and appeared to scent-trail another individual,
subsequently squatted by elevating its hindlegs and tail, ad-
pressing vent and thighs against the substrate, probably
leaving scent marker immediately adjacent to the scat (|. S.
Parmerlee, |r., pers. comm.). Competition: On Terre-de-
Haut, apparently displaced by I. iguana (Schardt, 1998b;
Breuil, 2002b). Foraging and Diet: Food includes fruits of
Prickly Pear (Opuntia),bird eggs, and ca r riot (Lazell, 197 3) ;
on La D6sirade, eats leaves, flowers, and grasses, sometimes
grazing alongside goats, a juvenile attempted to catch a
cricket (Schardt, 1998a); diet includes 28 species ofplants,
especially Tab eb ui a p alli da, Hipp o m an e m an cin ell a, Clero -
d en dr o n acule at um, C ap p ar is
Jle xuo s a, C. cy n ophallopho ra,
Pisonia fragans, Guajacum oficinale, Bursera simaruba,
Conocarpus erecta (Barre etal.,l997); may consume flowers
and leaves of more than 50 species of trees and bushes in
xeric coastal lowlands of Dominica, where iguanas may be
important seed dispersal agents (Malhotra and Thorpe,
1999); forages mostly in the morning, frequently associated
wlth Ipomea tiliacea, on which it avidly feeds, generalist
vegetarian, diet varies by habitat and island, but may in-
clude introduced species (e.g., ornamentals), on St.-Barth6-
lemy, feeds on leaves (especially during the dry season, and
favoring young leaves) and fruits (during the rainy season)
of Melicoccus bijugatus, Cordia dentata, Leucaena leuco-
cephala, Tabebuia pallida, Randia aculeata, consumption of
fruits and seeds may facilitate dissemination of these spe-
cies, some food items are toxic to birds and mammals,
drinks by licking leaves or opening mouth with head ele-
vated during rains, may "taste' food before eating (Breuil,
2002b); on Anguilla, particularly fond of leaves, flowers,
and fruits of Chink B:ush (Bourreria succulenta) and flow-
ers of White Cedar (Tabebuia heterophylla; Hodge et al.,
2003); readily consumes ornamental plants, and appears to
exhibit a particular fondness for Hibiscus sp. on Dominica
(Daniells et al., 2008); on St. Eustatius, individuals may
"gorgd' (consume all vegetation within reach before mov-
ing) or "grazd' (take only a few bites before moving), fre-
quently feeding on Tamarind (Tamarindus indica) leaves
(S. P. Fogarty, unpublished data); on Dominica, female ate
dry dog food in a bowl (E. A. Daniells and N. |. V6lez Es-
pinet, pers. comm.). Home Range and Territoriality: On
Dominica, male territeries during the breeding season
1,000 m'z, female territories 2,000 m2, female ranges overlap
those of other females and sometimes several males, sug-
gesting a female dominance hierarchy (Day et al., 2000;
Breuil, 2002b); males probably defend territories outside of
the reproductive season (Breuil, 2002b). Hurricane Ef-
fects: Maybe adapted to habitats subjected to periodic hur-
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134 . Part ll. Reptilia, Section l. Squamata (lizards)
deloupe, mate in March and lay on beaches in May, accom-
panied by males (J.-8. Du Tertre, 1667 in Breuil, 2002b); on
Dominica, females may move 460-900 m to egg-deposi-
tion sites (Day et a1., 2000); on St.-Barth6lemy and Petite
Terre, some females lay eggs within their home ranges,
while others move considerable distances (Breuil, 2000c);
nests in sand -1 m deep ending in a chamber large enough
to allow the female to turn around, in rocky areas, nest
chambers less extensive (Breuil, 2002b); on ilet Chancel,
suitable nesting sites are restricted and females often exca-
vate eggs laid by other females (Bach and Schonecker,
2008); on Martinique, clutch size 20-30 (Bouton, 1630 in
Breuil, 2002b) ; on Guadeloupe, 13-25 ( J. -B. Du Tertte, 1667
in Breuil, 2002b); on Dominica and Martinique, 15-30
(Anonyme De Carpentras, 1618-1620 in Breuil, 2002b); on
Dominica, 8-18, strongly correlated with female size, aver-
age weight 25 g (Day et al., 2000); to 22 eggs with a length
of44 mm (Schardt, 1998a); freshly unearthed (by other fe-
males) eggs 17-22 g,45 x 25 mm (Breuil, 2002b); incuba-
tion 3 months (Day et a1., 2000); captive hatchling 20 g,
SVL77 mm (Reichling, 2000); hatchlings 17.6'18.9 g, SVL
80-83 mm (Gibson, 2001); egg and hatchling sizes vary by
island and habitat (Breuil, 2002b); because clutch size is re-
lated to female size, the smaller size of iguanas on Chancel
(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.-
Barth6lemy (1.-C. Plassais in Breuil, 2002b). Size: Male SVL
to 434 mm, female SVL to 401 mm (Breuil, 2002b); iguanas
on Chancel are smaller than those on Petite Terre (Breuil,
2009). Swimming: Male with body temperature 28
oC
jumped into water 25-26 "C, swam on the surface for 80
sec, then dove for 16 sec at a depth of 2-3 m, when emerged,
floated perpendicular to water's surface, before climbing
oC,
onto a boat, within 2 min body temperature rose to 29
6 minutes later dove back into the water, held his breath for
28 min while immobile at 3 m at26 "C, swam on the surface
for 6 min (M. Breuil and B. Thi6bot in Breuil, 2002b; Breuil,
2003). Thermal Biology: Position themselves to facilitate
effective thermoregulation (Lazell, 1973); mean body tem-
peratures on La D6sirade 37.98 "C (36.53-38'77 "C) at sites
with temperatures 30.5-31.0 "C at 0800 h, 41.0-42.2 "C at
midday (but in shaded sites where iguanas are most fre-
quently seen -28-31 'C), -34-35 'C at 1500 h, dropping to
30.3-31.0 oC in late afternoon and 28.5-29.0 "C at 1800 h,
nighttime temperatures 24.5-25.0 "C (Schardt, 1998a); he-
liothermic, males warm at exposed arboreal sleeping sites,
females often bask on the ground, when foraging often
shuttle in and out of direct sun, highest temperature re-
corded for a male digesting food 39.4
oC, rest in shade dur-
ing the hottest parts ofthe day (Breuil, 2002b);on St. Eusta-
tius, maintains above ambient temperatures through the
night even when sleeping on exposed perches (RP).
Conservation Status: Most populations remain extremely
vulnerable to human exploitation for food, habitat altera-
tion, introduced predators and competitors, and hybrid-
ization with introduc ed lguana iguana (e.g., Breuil, 2002b,
2009; Powell, 2004e; Powell and Henderson, 2005a; Pow-
ell et al., 2005; Maillard and Breuil, 2007); on Chancel, an
additional threat is posed by tourists who trample one of
two principal communal nesting sites, although recent
improvements (including fencing) have reduced the risk
from tourists and sheep, also a proposal to relocate igua-
nas from Chancel to ilet-ir-Ramiers may provide another
refuge from possible interbreeding with invasive I. iguana
(Breuil, 2009); some populations (e.g., Dominica) are af-
forded protection under local laws (|ames, 2004); the spe-
cies is listed on CITES Appendix I (UNEP-WCMC,2008);
on the IUCN Red List as "vulnerable" (Breuil and Day,
1996), but many individual populations meet criteria for
'tritically endangered" status (Breuil,2002; Fogarty et al.,
2004; Powell, 2004a; Powell and Henderson, 2005a); Powell
Q00ad and Powell and Henderson (2005a) recommended
a reevaluation and a probable upgrade to "endangeredl'
Additional Pertinent Literature: Pasachnik et al. (2006).
l guano iguana Linnaeus,'1758
Distribution: Mainland from Mexico to Brazil, including
numerous continental islands; introduced into southern
Florida, Hawaii, and Israel; in the West Indies, on Gre-
nada and the Grenadines, St. Vincent, Barbados, St. Lu-
cia, Martinique, Marie-Galante, Les iles des Saintes, Gua-
deloupe (Grande-Terre, Basse-Terre, and Iles de Ia Petite
Terre), Montserrat, Antigua, Barbuda, Saba, St.-Martin/St.
Maarten, Anguilla, St. Croix (may have been introduced
by Caribs for food; Beatty in Grant, 1937b), Virgin Islands,
Puerto Rico (including Culebra), Grand Cayman, and Swan
Islands; elevational range, sea level to -800 m; population
on Barbados apparently extirpated (although some cap-
tive animals remain), those on Martinique, Marie-Galante,
Les iles des Saintes, Guadeloupe, and St. Croix possibly
introduced or of mixed origin (e.g., the historic popula-
tion on St. Croix presumably natural, but most currently
present lizards probably are descendants ofintroduced in-
dividuals), and those on Antigua, Barbuda, St.-Martin/St.
Maarten, Anguilla, Virgin Islands, Puerto Rico, and Grand
Cayman are introduced, although at least some individu-
als on Anguilla arrived via natural over-water dispersal,
almost certainly from Guadeloupe (Censky et al., 1998);
populations on St. Lucia, Saba, and Montserrat (and pos-
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saJI -raao's1a4cod puelur ur pue
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gi. aepruen6l ,(;rureg
136 ' Part ll. Reptilia, Section 1' Squamata (lizards)

on
1932h); on Carriacou (Grena- Gonzillezand V. Rios in Joglar, 2005b); egg deposition
U.S. Virgin Islands (Grant, (April-May)' favored
have a Terre-de-Haut at end of dry season
dines), borallus grenadensis (Pendlebury' 1974);
and their flesh is reputed to nesting sites along beaches among clumps of vegetation'
"reputation as excellent eating (Schardt' 1998b);
dogs (on St' Lucia; Ben- large rlumb.rs of hatchlings in October
tasie like chicken' (Long, 1974); to two known nesting
distances
don, 2003) and probably mongooses (Morton' 2007);
other -u:y -o,r. considerable
females moved 80 m' 1'57 km' and
Hawks beaches on St. Lucia, 3
than humans and domestic animals, Broad-winged home ranges' 5 nests on
(Malhotra and 1.61 km fromnesting sites back to
(Buteo platypterus) arethe main predators
(Falco St. Lucia wlthlT-2leggs (mean 19'5)' hatchlings on nest-
Thorpe, 1999); presumed predators include Kestrels and Morton'
ing beaches mid-May-mid-August (Graham
spariarius), Red-tailed Hawks (Buteo jamaicensls)'
some
14 nests with
and introduced dogs' Zio+,zoOs);average clutch size 23 (14-28) in
seabirds, large racers (Akophis spp'), to
(Powell et al'' 2005); on 637o success (Morton, 2OO7)' Sizet Adult SVL
hatching
cats, rats, mongooses, and humans recent hatchlings
aitve' 500 mm (Scliwartz and Henderson, 1991);
Grand Cayman, a Great Egr et (Ardea alba) capltred (mean 88
Grand Cay- (October on Terre-de-Haut) 82-95 mm SVL
nile iguana (T. Ebanks, in litt., 15'XII'2008); on 1998b)' Thermal
mm) and 13.22 g(mean 19.4 g; Schardt,
Crbophis caymanus (T' Ebanks' in litt" 23'II'2009)' thermo-
^un, in males Biology: Position themselves to facilitate effectlve
Reproduction: Females mature at -3 years' later by iguanas
Feb- .. glrlutio., (Lazell, 197 3) ; in habitats frequented
who must defend territories, males highly territorial oC between 0930 and 1030 h' peaking
in late teirperatures of 34.3
ruary-March, females migrate to egg-laying sites
at 1500 h and
38-45'5 x at ai.g "C at 1215-1330 h, decreasing to 35'7 "C
Aprii-early May, female (890 g) with 17 eggs (Schardt' 1998b)'
in- to a minimum of 24.0 "C at night
22-28.2 mm, 14-24 g, largest eggs in distal oviducts'
cubation 90-92 days, hatchlings ]uly-early August'
corre-
Conservation Status: Listed in CITES Appendix
II (UNEP-
(Breuil' 2002b); eggs countries'
sponding to onset ofthe rainy season WCMC, 2008), but export quotas exist for many
in old charcoal or products (leather
luid ir, *u.-, well-drained sites on beaches' primarily for live animals (pet trade)
hatched in made for native versus
pits, and gardens (Breuil, 2000a, 2000c); 17 eggs
goods and meat), no distinction is
lays populations;
i+ *".t ,iurrderwood, 1962;Lotg,1974); presumably introduced or for continental versus insular
eggs December-February, on Montserrat'
female 275 mm
hunting is usually prohibited, but enforcement
of laws
mm) on 24March animal in
SVL with 16 eggs (42.0-46.8 x 26'6-28'4 is lax at best, the species is considered a game
may not ap-
Grenada and St. Vincent, with a regulated hunting
Rico (although all data season
(Lazell,l973);on Puerto
season No- routinely ignored
ply to Puerto Rican populations), reproductive and bag limits (on Grenada), which are
which
vember-December, males establish territories from by local hunters (Powell, 2004e; Powell and Henderson'
similar to the "greatly reduced in
other males are actively expelled, courtship 2005a,2007); population on St' Croix
depends on man as by
aggressive displays between males, copulation numbers (probably as much by the activity of
rarely copu- pre'
females response, may last 15 min, but males that of the mongoose)" (Seaman and Randall' 1962);
eggs (17-68/clutch) require high populations in the U'S' Virgin Islands
late more than once/day, sumably introduced
incubation temperatures, nests in clearings or on beaches'
are afforded special Protection under USVI
Code Title 12
on
may be ugg..gut.d or dispersed, apparently depending (Platenberg, 2007).
years' may be 1 m
tlpe of soil, nests may be used for several
Remarks: Native West Indian populations currently
as-
with sev-
i..p urrd connected by tunnels to 20 m in length signed to lguana iguana may rePresent more
than one spe-
may be com-
eral openings to the surface, egg deposition
(A' cies (e.g., StePhen, 2007)'
(Ri r..o, 1998); in Puerto Rico' clutch sizes 12-90
-rrrul