iguana volume 10, number 3 september 2003 us$6 · diverse cadre of species. the long-term (20+...

Adult male St. Lucian Iguana (Iguana iguana) (story on p. 71). Photograph by Matt Morton, Durrell Wildlife Conservation Trust.

IGUANAJournal of the International Iguana Society www.IguanaSociety.org

Vo l u m e 1 0 , N u m b e r 3 S e p t e m b e r 2 0 0 3 U S $ 6 . 0 0

In This Issue...FEATURESA Backward Glance at Iguana ExploitationHenry S. Fitch and Robert W. Henderson . . . . . . . . . . . . . . . 63

Exploitation of Reptiles in the West Indies:A Long HistoryRobert Powell . . . . . . . . . . . . . . . . . . . . . 67

The St. Lucian Green Iguana — A Special Case?John S. Bendon . . . . . . . . . . . . . . . . . . . . 71

Black Iguanas: Name and SystematicsGunther Köhler, AJ Gutman, and Robert Powell . . . . . . . . . . . . . . . . . . . . . 79

HUSBANDRYIguanas and Artificial Ultraviolet Light:How and How Much Made Simple — Well, Not Exactly Simple…Bob MacCargar . . . . . . . . . . . . . . . . . . . . 82

PROFILEHenry S. Fitch: A Legendary Passion forNatural HistoryRobert W. Henderson . . . . . . . . . . . . . . . 86

HISTORICAL PERSPECTIVEWhorl-tailed IguanasFranz Werner . . . . . . . . . . . . . . . . . . . . . . 89

BIOGRAPHICAL SKETCHRaymond L. Ditmars (1876–1942) . . . . 92

BOOK REVIEWThe Reptiles and Amphibians of Anguilla,British West IndiesAJ Gutman . . . . . . . . . . . . . . . . . . . . . . . . 93

Iguana Newsbriefs . . . . . . . . . . . . . . . . 94Editorial Information . . . . . . . . . . . . . . 96Letter from the President . . . . . . . . . . . 98

(See page 71)

INTERNATIONAL IGUANA SOCIETY

www.IguanaSociety.org

Phot

ogra

phs

by M

att

Mor

ton,

Dur

rell

Wild

life

Con

serv

atio

n Tr

ust.

Over a quarter of a century has passedsince we first investigated iguanaexploitation in Central America. A“friend” recently pointed out to us that

we were a couple of “old-timers” when it comesto that particular subject (see also the Profile on p. 86). Prior to collaborating on Ctenosaura andIguana research, we independently had done con-siderable field work with these lizards. Fitch stud-ied C. similis in Costa Rica in the late 1960s andHenderson worked with juvenile I. iguana andwith C. similis in Belize in 1970–1. We were bothwell aware of the human-induced pressureimposed on these lizards, and we were interestedin bringing their plight to the attention of our col-leagues, conservation organizations, legislativebodies in the countriesin which we wereworking, and to thegeneral public (see thesummary in Fitch et al.1982, full citation atthe end of this article).

In the UnitedStates, most folksfamiliar with iguanasassociate them with thepet trade or considerthem curios, and theirimportation for thesepurposes is a minorindustry. However,within the iguanas’ ranges in tropical America,these lizards are hunted for food and, in someareas, may be the principal game animals. In orderto determine the extent of iguana exploitation andlearn the methods used to hunt and subdue them,we visited each of the Central American countries,observed iguanas in their natural surroundings and

in mercados, and we interviewed hunters and gov-ernment officials.

Early records revealed that iguanas had beenhunted since ancient times by many of the nativecultures, but, thanks to their high reproductivepotentials, they had been able to maintain theirnumbers over hundreds of years despite exploita-tion. However, the widespread use of insecticidesto protect crops, habitat destruction resulting inlarge part from an increase in the number of peo-ple, and overhunting inevitably altered ecosystems.The result was that both Green Iguanas andctenosaurs have experienced rapid reductions innumbers in recent years and have disappeared frommany areas where they were formerly abundant.

63IGUANASeptember 2003

A Backward Glance at Iguana Exploitation

Henry S. Fitch1 and Robert W. Henderson2

1Henry S. Fitch Natural History Reservation, University of Kansas, Lawrence2Milwaukee Public Museum, Milwaukee, Wisconsin

All photographs by RWH.

Ctenosaura similis in the Belize City cemetery.

A ctenosaur emerging froma Belize City cemetery grave.

64 IGUANA Volume 10, Number 3

Both Green Iguanasand ctenosaurs havebeen important proteinsources for peoplelocally. Each kind ispreferred in certainparts of the range.Beyond its actual foodvalue, the flesh is val-ued for its supposedmystical power, andconsumers are pre-pared to pay muchmore than they wouldfor a similar amount ofbeef or chicken. TheCatholic Church has ruled that iguana flesh, likefish, is a permissible substitute for “meat” (flesh ofmammals or birds) on Fridays and during Lent.Over much of the range, ctenosaur flesh is highlyvalued as an aphrodisiac. In Belize, however, it isconsidered unfit for human food. The Belize Citycemetery supports a colony of Ctenosaura similisand many of the lizards have burrows in the gravemounds. This led to the belief that they feed onhuman corpses in their graves.

In the MercadosFrom the Isthmus of Tehuantepec along the

Pacific Coast to Lake Nicaragua, at least 25 townshave mercados where iguanas were formerly sold.El Salvador was a major importer. Having depletedits own population of iguanas, it received themfrom Honduras, Guatemala, and Nicaragua byboat and by bus. The neighboring countriespassed legislation banning the trade, but pedestri-ans carrying sacks or baskets full of reptiles in many

instances were allowed to cross the border into ElSalvador.

Cruel treatment of captured iguanas was rou-tine. In the mercados, buyers often requested thatvendors sell them unlaid eggs without the females.The vendors would slit the females’ abdomens andrip out the eggs. Many commonly believed thatsuch stripped females would survive and producemore eggs. Females stripped of their eggs anddying from the crude surgery were a commonsight where street vendors were plying their trade.Also, because ctenosaurs were liable to bite a han-dler, their mouths were routinely sewn shut. Inaddition, pulling off the claw of the fourth(longest) hind toe with pliers and drawing out thetendon was common practice. The exposed ten-don was then tied to that of the opposite side sothat the animal was immobilized. Iguanas wereoften stuffed in sacks, several layers deep, andmight be kept there for long periods without foodor water.

Tethered Green Iguanas at the Chinandega, Nicaragua market(February 1976).

Male and female ctenosaurs with their mouths sewn shut(Nicaragua, February 1976).

Visiting the mercados, we would at timesencounter many dozens of iguanas. Approachingthe vendors, we requested permission to examinethe lizards and to take measurements and canrecall being spurned only once; the rest were more

than willing to let us handle their merchandise. Wewould often work for several hours at a stretch,standing in open markets and recording data,while curious onlookers watched. I’m sure ourpeculiar behavior was attributed to the fact that wewere gringos, thereby making it perfectly accept-able. After handling several dozen adult GreenIguanas and, especially, ctenosaurs, our armswould be covered with scratches and cuts, some-times from the lizards’ claws, but more often fromtheir lashing tails.

Visting the HuntersThe iguana vendors provided us with the

names of villages from which the lizards origi-nated. Based on this information, we worked back-ward from the mercados and were able to trackdown several hunters in order to learn how theycaptured their prey. In hunting iguanas for homeconsumption, a .22 rifle was often used, but someCentral American countries banned the use offirearms or even slingshots. Professional huntersfor the commercial market in El Salvador capturedtheir iguanas alive and unharmed. This was usuallyaccomplished with a bamboo pole and noose. Thehunter slowly approached the basking lizard andextended the pole with noose toward it, with apiece of calf’s liver dangling as bait.

Hunters often used trained dogs, and thesewere especially effective in hunting female Green


A pile of iguanas: mostlyCtenosaura similis, but some Green Iguanas, too(Chinandega, Nicaragua;February 1976).

An iguana vendor at theChinandega, Nicaraguamercado (February 1976).

Iguanas during the nesting season. The hunterwould move stealthily along a stream, scanning theterrain ahead for signs of nesting activity. Severalfemales might be digging their nesting burrows ina sand bank, and some might have their forequar-ters in a hole so that they could not see dangerapproaching. At a signal from the hunter, the dogwould race ahead and might catch a female beforeshe could escape from the burrow or outrun herbefore she could reach the safety of a tree. Femalesbearing a heavy burden of eggs were relatively slowrunners. The dogs were trained not to injure thelizards.

Some hunters used shovels to dig iguanasfrom their burrows; excavation also could beaccomplished with a pointed stick and their hands.Females concentrating on nest excavation wereespecially vulnerable to capture. Again, a femalewith her head and forebody in the burrow was lesslikely to be aware of an approaching hunter than alizard involved in other activities.

We also became aware that iguanas weresometimes used merely for target practice. Theywould be shot with a .22 rifle from a vehicle asthey sat on roadside boulders or fence posts. If not

killed outright, ctenosaurs that were shot oftenescaped into nearby burrows, whether or not theirwounds were mortal. The “hunters” would makeno attempt to retrieve the dead or woundedlizards.

What’s Happenend Since?After our multi-year stint with iguanas, we

moved on to new projects or resumed old inter-ests. Fitch continued his studies of Anolis lizardsand his long-term work with Kansas snake ecology.Henderson moved on to the West Indies and itsintriguing snake fauna. We assume that the merca-dos of Central America no longer sell iguanas inthe numbers that we observed in the mid-1970s.Is it because of legislation protecting the tastylizards, or because population densities have dra-matically plummeted? We suspect the latter. TheU.S. now harbors denser populations of Cteno-saura similis and Iguana iguana than do someareas where they occur naturally.

On a brighter note, interest in iguana biologyhas never been greater, and exciting research isbeing conducted in many countries and with adiverse cadre of species. The long-term (20+ years)mark-and-recapture study of John Iverson onCyclura cychlura inornata in the Bahamas is espe-cially exciting. Also, efforts to protect species withdramatically declining numbers (e.g., Ctenosaurabakeri, Cyclura lewisi) are, on the one hand, dis-heartening because the numbers are so danger-ously low, but, on the other, gratifying that peoplecare enough about the iguanas to do something.Iguanas of all species are fortunate to have theInternational Iguana Society in their corners.

ReferencesFitch, H.S., R.W Henderson, and D.M. Hillis. 1982. Exploitation

of iguanas in Central America, pp. 397–417. In: G.M.Burghardt and A.S. Rand (eds.), Iguanas of the World: TheirBehavior, Ecology, and Conservation. Noyes Publ., Park Ridge,New Jersey.

Krysko, K.L., F.W. King, K.M. Enge, and A.T. Reppas. 2003.Distribution of the introduced Black Spiny-tailed Iguana(Ctenosaua similis) on the southwestern coast of Florida.Florida Sci. 66:141–146.

Valiulis, J.M., K.N. Hines, and J.B. Iverson. 2003. Adventures ofthe iguana patrol, pp. 39–48. In: R.W. Henderson and R.Powell (eds.), Islands and the Sea: Essays on HerpetologicalExploration in the West Indies. Soc. Study Amphib. Rept.Contrib. Herpetol., Vol. 20, Ithaca, New York.


A young iguana vendor and her merchandise (Chinandega,Nicaragua; February 1976).

Abundant remains of sea turtles, tortoises,iguanas, and even snakes in archaeolog-ical excavations throughout the WestIndies testify eloquently to the fact that

Amerindians frequently and successfully exploitedreptiles. At least some biogeographers have sug-gested that the first human inhabitants of theregion carried with them favored species, such asRed-footed Tortoises (Geochelone carbonaria),establishing populations on islands as theymigrated north from South America to serve as aready source of food during subsequent visits1.Aboriginal inhabitants have been implicated in theintroduction of iguanas on islands in the Gulf ofCalifornia (see Iguana Times, June 2002) and alsomay have established some West Indian popula-tions. Furthermore, the earliest Americans havebeen blamed for the extinction or extirpation of

Exploitation of Reptiles in theWest Indies: A Long History

Robert PowellDepartment of Biology, Avila University, Kansas City, Missouri


A mongoose (Herpestes javanicus) from Savane-à-Mulets,Soufrière, Guadeloupe. These efficient predators have beenimplicated in the extirpation of populations and even theextinction of diurnally active snakes and ground-dwellinglizards (photograph by Alain Fossé).

Today, the AmericanCrocodile (Crocodylusacutus) is endangered, butin the early 20th Century,exploiting the species forhides and meat waswidespread and “officically”sanctioned. Here, convictsskin a crocodile in theDominican Republic in 1922(photograph by G. K. Noble;courtesy of the Departmentof Library Services, AmericanMuseum of Natural History).

1 Some authorities, noting the abundance of tortoises onoceanic islands, their ability to float, and their resistance toexposure to salt water, disagree and argue that Red-footedTortoises arrived naturally in the islands by means ofoverwater dispersal from South America.

native tortoises and some populations of iguanas(along with a number of large, endemic mammals)that are known only from fossil or archaeologicalevidence. If true, exploitation of at least somespecies was not sustainable — and the popularimage of Native Americans living in harmony withnature clashes with reality.

The West Indies have been exposed continu-ously to Europeans longer than any other region ofthe Western Hemisphere. Columbus spoke ofabundant turtles and iguanas, and, beyond anydoubt, he, his men, and those that followed con-sidered them resplendent table fare. With theadvent of the slave trade, Africans, most of whomwere products of cultures that exploited wildlife intheir native lands, continued to do so in the NewWorld. Like the Amerindians before them, WestIndians of European and African descent areblamed for translocations, extirpations, and extinc-tions. Although many of the latter are attributableto habitat alteration and destruction or competitionwith and predation by feral mammals, exploitationlevels were undoubtedly high — and often haveexceeded sustainable levels into recent decades.

Food undoubtedly has been the primary moti-vating factor. Most large reptiles, especially thosethat evolved on islands in the absence of largepredators, were at least initially abundant — andare easy to catch, nutritious, and tasty (note theobvious reference in the name Iguana deli-

catissima, for example). Even easier to acquire arethe eggs of turtles, iguanas, and crocodilians. Inaddition, however, turtle shells and bones of manylarge species were used as tools. Shells and hideswere used for the construction of shelters or fordecorative purposes, and some animals or theirparts became important in ceremonial functions.Snakes still play a significant role, for example, invoodoo rituals. Crocodilian penises and some


In 1922, Iguana hunterson Isla Beata, DominicanRepublic, used dogs to catch HispaniolanRhinoceros Iguanas(Cyclura cornuta) (photo-graphs by G. K. Noble;courtesy of theDepartment of LibraryServices, AmericanMuseum of NaturalHistory).

internal organs were thought to possess aphrodisi-acal properties.

Exacerbating the impact of exploitation foruseful purposes were efforts at extirpation moti-vated by fear or dislike. On Martinique and St.Lucia, the only West Indian islands with nativepopulations of pitvipers, these are hunted actively,and bounties have been offered intermittently bythe authorities until recent years. On many islandswhere no species are dangerous to humans, snakesnevertheless are killed whenever encountered.Crocodilians are widely feared and killed, oftenwithout any effort to salvage the meat or other use-ful parts. Even on the few islands on which humanshave not traditionally eaten iguanas, they (andother species of large lizards) are frequently perse-cuted solely because they are perceived as beingugly and strange.

Although presented in a historical context,many of these practices continue largely unabatedtoday, despite the protected status of many species.Shells of sea turtles are piled high in many ports,and freshwater turtles and iguanas show up regu-larly in small towns or even city markets. Especiallypeople living in rural areas routinely supplementtheir diets with reptilian meat and eggs. Fear ofsnakes, large lizards, and crocodilians remainsresponsible for the deaths of many reptilesthroughout the region.

However, in addition to the traditional reasonsfor exploiting reptiles, one of the most pervasivecauses of local extirpations today is unregulatedcollection for the burgeoning pet trade.Authorities in essentially every island nation cancite example after example of unscrupulous collec-

tors catching, buying, and smuggling enormousnumbers of reptiles of all sizes and species — andthese commercial collectors are, of course, onlythose that have been caught. Strangely enough,scientific collectors in the 19th and early 20th cen-turies initially developed a common practiceemployed today by these criminals2. Offeringbounties to locals eager to supplement their mea-ger incomes and often very familiar with the habitsof the local species, scientists historically and, now,suppliers to the pet trade harvest huge numbers ofanimals. The fact that rarities demand the highestprices imposes particularly intense pressure onpopulations of threatened or endangered species.To their credit, today’s scientists have largely aban-doned this practice.

What, if anything, is being done? Legislationnow protects many of the most vulnerable speciesand some habitats, but enforcement is lax ornonexistent in many cases. Educational efforts onmany islands have had some positive impact, par-


2 Scientific collecting has not been implicated in anyextirpations or extinctions of West Indian reptiles and theknowledge acquired by scientists who take specimens foracademic purposes often has been invaluable in develop-ing conservation strategies for specific species or entirebiotic communities.

A Green Turtle (Chelonia mydas) awaits slaughter on thebeach of Isla Saona, Dominican Republic (photograph byRobert W. Henderson).

This snake hunter was photographed on St. Lucia in 1888.The snake is Bothrops caribbaeus and it may have beenkilled for a bounty, or merely because it was a snake andpotentially dangerous (photograph courtesy of PhilipWalwyn).

ticularly when coupled with successful efforts atdeveloping ecotourism. Animals become morevaluable as means of attracting free-spendingtourists than they would bein the market. Unfortu-nately, education takestime and, in many cases, isdirected at children. By thetime they are in a positionto develop, implement,and enforce enlightenedpolicies, many additionalpopulations and evenspecies of reptiles will havedisappeared. Even eco-tourism has its drawbacks,since many travelers sup-posedly enamoured withwildlife buy curios madefrom exploited reptiles anddemand accommodationsand amenities that areincompatible with sustain-ing viable natural habitats.

What can any one indi-vidual do? One shouldabide by all legislated efforts to conserve nature,avoid all actions that might in any way encouragethe exploitation of wildlife, support educationalefforts by governmental agencies and non-govern-mental organizations, if a pet reptile is to be pur-chased, select only captive-bred animals fromknown sources, and travel lightly on the land whenin the islands.

AcknowledgementsRobert Henderson’s comments on an early

draft improved this article immensely.

ReferencesNote that the references listed below are surveys that, in turn,provide citations to many additional pertinent resources, as do thevolumes in which they appear.

Hedges, S.B. 1999. Distribution patterns of amphibians in theWest Indies, pp. 211–254. In: W.E. Duellman (ed.), Patternsof Distributions of Amphibians: A Global Perspective. JohnsHopkins Univ. Press, Baltimore, Maryland.

Hedges, S.B. and R. Thomas. 1991. The importance of systematicresearch in the conservation of amphibian and reptilepopulations, pp. 56–61. In: J.A. Moreno (ed.), Status yDistribución de los Reptiles y Anfibios de la Región de PuertoRico. Depto. Recursos Naturales de Puerto Rico, Publ. Cient.Misc. 1:1–67.

Henderson, R.W. and R. Powell. In press. Thomas Barbour andthe Utowana expeditions (1929–1934) in the West Indies.Bonner Zool. Beiträge.

Powell, R. and R.W. Henderson. 1996. A brief history of WestIndian herpetology, pp. 29–50. In: R. Powell and R.W.Henderson (eds.), Contributions to West Indian Herpetology:A Tribute to Albert Schwartz. Soc. Study Amphib. Rept.Contrib. Herpetol., vol. 12. Ithaca, New York.

Powell, R. and R.W. Henderson. 2003. Some historical perspec-tives, pp. 9–20. In: R.W. Henderson and R. Powell (eds.),Islands and the Sea: Essays on Herpetological Exploration inthe West Indies. Soc. Study Amphib. Rept. Contrib. Herpetol.,vol. 20. Ithaca, New York.

Williams, E.E. 1999. Over 300 years of collecting in theCaribbean, pp. 1–30. In: B.I. Crother (ed.), CaribbeanAmphibians and Reptiles. Academic Press, San Diego,California.

Wing, E.S. 1989. Human exploitation of animal resources in theCaribbean, pp. 137–152. In: C.A. Woods (ed.), Biogeographyof the West Indies: Past, Present, and Future. Sandhill CranePress, Gainesville, Florida.

Wing, E.S. 2001. Native American use of animals in theCaribbean, pp. 481–518. In: C.A. Woods and F.E. Sergile(eds.), Biogeography of the West Indies: Patterns andPerspectives. 2nd ed. CRC Press, Boca Raton, Florida.

Woods, C.A. 2001. Introduction and historical overview ofpatterns in West Indian biogeography, pp. 1–14. In: C.A.Woods and F.E. Sergile (eds.), Biogeography of the WestIndies: Patterns and Perspectives. 2nd ed. CRC Press, BocaRaton, Florida.


A roadside vendor selling “Jicotecas”(Trachemys stejnegeri) collected in theLaguna de Saladillo, Parque NacionalMontecristi, Dominican Republic(photograph by Robert Powell).

Hatchling American Crocodiles(Crocodylus acutus) offered for sale totourists in a shop in Santo Domingo,Dominican Republic (photograph byDonald D. Smith).

“I’m optimistic that the St. Lucian Iguana will besaved, and that we will be able to protect themin such a way that they will be able to expandinto new areas. My dream is to see the iguanarepopulate a much wider area of the island…”

— Donald Anthony, Wildlife Officer, St. Lucia

St. Lucia is one of the Windward Isles in theLesser Antillean chain, which extends fromjust southeast of the Puerto Rico Bankalmost to South America. With an area of

610 km2, the island is topographically and biotically

diverse. Montane uplands are covered by rainforest,whereas coastal areas are characterized by dry wood-lands. Human influence is evident in the abundantagricultural areas and numerous communities.

Green Iguanas (Iguana iguana) are widelydistributed through Central and much of SouthAmerica, and have been introduced and becomeestablished in many tropical and subtropical areasthroughout the world. St. Lucia is the northern-most of the Antilles on which these animals occurnaturally.

Introduced Green Iguanas are all too commonand indeed, have become pests in parts of Florida(see Newsbriefs, p. 94). On some of the more

The St. Lucian Green Iguana — A Special Case?

John S. BendonBath, England


Large adult male St. Lucian Green Iguana. Note the pale ground color and the length of the dorsal crest scales.

northerly isles of the Lesser Antilles, they pose athreat to populations of their endangered relative,Iguana delicatissima, through competition forfood and habitat and by reproducing at a higherrate. In some areas, they also have threatened theintegrity of the unique I. delicatissima gene poolby hybridizing with the native animals.

Unlike the introduced populations, whichhave become established only in recent years, St.Lucian Iguanas may have been isolated from otherGreen Iguana populations for as long as hundredsof thousands of years — and consequently appear

to be a bit different. Most obviously, St. LucianIguanas have “horns,” dramatically enlarged scalesthat project conspicuously from their snouts.These vary considerably in size and seem to bemore prominent in older animals. Although rela-tively rigid and firm (much more so than the mid-dorsal crest spines, for example), they do “bend”from side to side. They also are thicker in diame-ter than dorsal crest scales, although, in cross-sec-tion, they are somewhat flattened laterally. Thecoloration and shape of some large scales on thelower jaw also differ from those of other Green


An adult male St. Lucian Green Iguana (a) and a young adult female St. Lucian Green Iguana (b) compared to male (c) andfemale (d) Green Iguanas from elsewhere in the species’ range. Note the horns and differing shapes of scales on the jaws.Drawings by the author.

A

B

C

D

Iguanas. Catherine Malone, Purdue University,has examined the DNA of many iguanids, andrecently has been concentrating on St. LucianIguanas. Her work is yet to be published, but pre-liminary results show distinct genetic differencesbetween St. Lucian and other Green Iguana pop-ulations.

Although based on a relatively small sample ofa few dozen adults and recognizing considerablevariation among individuals, St. Lucian Iguanasappear to undergo ontogenetic changes in colorand pattern. Hatchlings are bright or yellowishgreen with cloudy gray or black bands. Thesebands seem to fade in the first year or two of life,when the iguanas are a very intense green. As theanimals continue to age, the green (usually) losesits intensity and the bands appear to darken. In theoldest (or, at least, the largest) individuals, thegreen fades to a pale gray and the black bands arequite prominent.

Until recently, no research has been conductedon these iguanas. Now, with support from theDurrell Wildlife Conservation Trust, MatthewMorton of Bristol, England and local assistants areimplementing a project in cooperation with the

Forestry Department (part of the Ministry ofAgriculture) of the St. Lucian government. TheForestry Department has provided field assistanceand technical expertise, as well as vehicles andoffice facilities. Head of Department Brian Jamesand Deputy Michael Andrews have taken a keeninterest, and Donald Anthony and Alwin Dornellyfrom the Department’s Wildlife section are work-ing closely with Morton. Karen Graham of theSedgwick County Zoo (Kansas), Bill Toone of theSan Diego Zoo, and Catherine Malone also havecontributed greatly by assisting with the UnionZoo’s (St. Lucia) exhibit, education and commu-nity outreach planning, and blood sampling fromwild and captive iguanas for DNA analysis. Laborand travel expenses for Graham and Toone werecovered by their respective institutions, andMalone received funding through theInternational Iguana Foundation and the MiamiMetro Zoo to cover material costs for improve-ments to the zoo exhibit and educational posters.

The goals of the project are to: (1) determinethe population size and distribution of St. LucianIguanas, (2) establish the population’s taxonomicstatus, and (3) protect and conserve current pop-ulations.

Blood is drawn for DNA analysis and PIT tags(transponders) are inserted in each individual. Thelatter can be read using a type of barcode reader,and a unique number permanently identifies theanimal, even if the tag turns up in the belly of apredator. Some iguanas have been equipped withtransmitters and radiotracked.


A female iguana in the Limiere area.

Over 1000 hatchlings were observed on the main nestingbeach this year, but survival rate is very low due to predators.

The largest St. Lucian Iguana processed todate weighed over 5 kg and had a snout-ventlength of 45 cm. Current estimates of adult popu-lation size range in the hundreds, certainly lessthan 1000. Over 1000 hatchlings have been

observed on the main nesting beach, but very fewof that number survive to maturity.

St. Lucian Iguanas are restricted to the north-eastern part of the island, including the beaches.Several unsubstantiated reports also place at least afew animals in the southeast. Sightings or recentreports of iguanas are absent for the entire interiorof the island (moist forest) and the entire westerncoast. The small number of older reports that areavailable are difficult to interpret. At least some ofthese areas should be suitable for iguanas, so whyaren’t they there? Any number of factors might beresponsible.

“Dogs kill adult iguanas, either in hunts withpeople (a few people still eat them for food) or,more commonly, it seems, because they areallowed to run around unattended, killingwhatever they can. Hunting (by people) wasreportedly more common in the past, and manylocal people attribute perceived declines iniguana numbers at one site in particular to


An adult male St. Lucian Green Iguana that was recapturedon several occasions.

Illustration by John Binns.

over-hunting. The dreaded mongoose takeseggs and hatchlings; this year (unlike last) a catwas found, killing hatchlings, at the main nestsite. Natural predators of hatchlings seem to bepredominantly various species of herons and theAmerican Kestrel. Manicou (opossums) and ratsmay theoretically take hatchlings, though thereis no direct evidence of this. There is a singleobservation of a Boa constrictor in the act ofswallowing an adult, though generally boas aresmall here and I think this is likely to beuncommon.

“Traditionally there has been heavy sand miningat both the beaches where iguanas nest. Thesesites are also used by Leatherback Turtles and it isknown that their nests have been destroyed inthis way. So, it seems likely that iguana nestswould likewise be at risk, although sand miningis now less intense than in the past at these sites.At one of these beaches ‘semi-feral’ pigs are

reported to open turtle nests, though there is noevidence of the opening of iguana nests as yet.

“Clearance of large areas of land for bananagrowing in the last few decades may have hadan effect (during a habitat-use study usingradiotracking last year, all nine tracked iguanas‘pointedly’ avoided banana plantations).”

— Matt Morton

“The main things that are killing iguanas aredogs – especially dogs – cats and people.”

“When I see them, I’m feeling happy. They seeme, they give me a lot of signs – they’re shakingtheir heads [head bobbing]. They’re fine withpeople … Let it be, in the wild, like anyone else.”

— ‘Seko’, a St. Lucian field assistant on the project


Sign funded by the International Iguana Foundation and theDurrell Wildlife Conservation Trust; one of many posters madeby parents and teachers for pre-school age local children, andan iguana lacing trainer, also made by an adult for children.

The International Iguana Foundation and theDurrell Wildlife Conservation Trust have fundedsigns asking people to keep dogs on leashes in themain areas occupied by iguanas.

“Roots Farm is a cooperative farm, run by Rastasin one of the areas on St. Lucia where iguanasare still found. They practice organic farming andgrow a wide range of mixed crops, in contrast,for example, to monocultures of bananas, whichare commonplace in many areas. What drew ourattention to this place is that, unlike most placeson the island, this was one where you had afairly good chance of seeing an iguana. Andthat’s because of the Rastas’ gentle andsympathetic attitude toward iguanas and allwildlife. Hunting and unleashed dogs are nottolerated on their land.”

— Matt Morton

“I think, in this culture people, not knowing too much about wildlife, do not have anunderstanding of their importance and, as aresult, do not appreciate those animals aspeople who are wildlife-sensitive do. We’ve had,in the past, to do extensive work in terms ofeducation with the St. Lucia Parrot, and we can

see the successes of that. We know that evenschoolchildren know some of the basic factsabout the parrot and appreciate it. We wouldlike to create that sort of attitude in peopletoward the iguana, but we know that, rightnow, some people just look at the iguana as alizard, that it’s not good looking, and it mightnot be friendly.”

— Alwin Dornelly, Wildlife Officer, St. Lucia

Clearly, this mirrorsan attitude that unfor-tunately is commonthroughout tropicalregions of the world.People fail to appreciatetheir wildlife resourcesand are likely to dislike,misunderstand, ormerely exploit them.The only effectivemeans of changing atti-tudes is education.Posters designed as partof a local school’s activ-ity program and otheractivities such as games,coloring books, andchildren’s lacing train-ers made by parentsand teachers are meth-ods that are being used


An iguana burrow on St. Lucia.

Young adult St Lucian Iguana.


As part of the effort to protect and conserve thecurrent population of St. Lucian Green Iguanas,Matthew Morton of the Durrell Wildlife ConservationTrust has been regularly checking hatchlings on thenesting beach at Louvet. Among this year’s hatch-lings, Morton observed a number of anomalies.

Multiple “nest sites” are located at Louvet, at mostof which multiple females have nested. Four of themain sites and five of the less heavily patronized oneshave been fenced. Thus, certain effects can bedescribed as “per nest-site” and others as “perclutch,” and have a higher probability of being dueto nesting conditions or to circumstances originatingwith the parents, respectively. However, causativeagents remain in all cases speculative.

Some individuals appeared grubby andunhealthy with patches of gray scales in various loca-tions on their bodies. Some clutches seem to havemany individuals with this condition, whereas mosthave no affected animals. This condition also wasobserved in a few of last year’s hatchlings. This doesnot appear to be normal shedding and preliminaryobservations suggest that the affected hatchlings also

tend to have hatching weights toward the low endof the scale.

One suggested cause for the grey patches was abacterial infection acquired in the nest. This wouldaccount for some clutches being affected more thanothers. The mother could pass a bacterial infection tothe developing embryos. Bacteria will incubate in thedeveloping eggs and can produce systemic infectionsin the hatchlings. Secondary opportunistic organismscan localize topically and result in the skin discol-oration. Fungal infections also may be responsible,particularly given the damp nesting environment.

A different potential cause is poor nutritional sta-tus of the mother. The absence of necessary traceminerals such as selenium or vitamins such as B1 maysingly or in combination result in problems such asmalabsorbed yolk sacks or too little yolk to support ahatchling in the last hours before hatching and thefirst days thereafter.

Morton also observed some overt birth defects,mainly tails that were kinked very slightly in the ter-minal few millimeters and occasionally a more pro-nounced kink higher up. Again this seemed to affectseveral individuals in some clutches and none in oth-ers. This is most likely attributable to extreme incuba-tion temperatures, although the possibility ofinbreeding within a small population exists as well. Asingle individual displayed what appeared to behydrocephaly, a condition that, in humans at least,

Anomalous HatchlingsAll photographs by Matthew Morton.

These hatchling St. Lucian Iguanas emerged from asingle nest; the missing portions of their tails probablyfell victim to a rat or land crab invading their nest.

Hatchling St. Lucian Iguanas: the individual on the leftsuffers from patches of gray scales that most likely werecaused or exacerbated by bacterial or fungal agents, thehatchling on the upper right suffers from possiblehydrocephaly or an abscess, whereas the youngster onthe bottom right is quite healthy.

to get the message to adults and children at thesame time.

On some islands people are brought up eatingiguana as a delicacy. This is not sustainable.

“Actually, we’ve always recognized that the St.Lucian Iguana was in dire straits in terms of itsnumbers and its chance of survival becausewe’ve had a history of hunting iguanas in St.Lucia. Even [though] we had the wildlife lawspassed in 1980 with a fine of 5,000 [EasternCaribbean] dollars for anyone convicted ofcatching or killing an iguana [St. Lucian law alsoallows for fines and imprisonment for possessionof an iguana], it is very difficult to catchsomebody ‘in the act,’ because the areas arevery remote and [representatives of the ForestryDepartment] are stretched in terms of numbers:we don’t have the Forestry Officers to policeeverywhere. When you go to investigate anarea, people are tight-lipped — they won’tspeak out. It’s hard to get any firm evidence. Wehave been trying to source funding for a longtime to get extra help. In the Wildlife Section, Iam the only person working there until recently.Now Alwin Dornelly is my assistant. With aidfrom Durrell Wildlife [and funding grants from the Iguana Specialist Group and theInternational Iguana Foundation], we’re nowable to do the things we always wanted to do… doing surveys, finding out where thepopulation is, finding the threats where they arenesting, and other things.

“I can say right now that we have a pretty goodhandle on where the iguanas are located and apretty good idea as to where they are nesting,

and I’m sure that we will get a pretty good ideaof the size of the population.”

— Donald Anthony, Wildlife Officer, St. Lucia

Much work remains. This is only the second yearof research and plans are in place to conduct exten-sive observations of hatchlings on the main nestingbeach this year. With time, a much better pictureregarding the status of these lizards will emerge.

Species are disappearing at an alarming rateand we now know that we must act immediatelywhen warning signs are seen. The California GiantCondor (Gymnogyps californianus) and Spix’sMacaw (Cyanopsitta spixii) are two high-profilespecies that have disappeared in the wild. TheGrand Cayman Blue Iguana (Cyclura lewisi) alsois on the brink of extinction.

On St. Lucia, hope prevails. Enough iguanassurvive at the moment and the means by whichtheir numbers are being depleted are now wellknown. The right people are in place to deal withthe situation. This beautiful and distinctive GreenIguana is one of the jewels of the Caribbean —and it deserves a future.

AcknowledgmentsI am very grateful to Matthew Morton for

providing interviews, valuable information, andphotographs. Brian James, Michael Andrew,Donald Anthony, Alwin Dornelly, CatherineMalone, Karen Graham, Bill Toone, and JohnHartley have read and commented on variousdrafts of this article.

All illustrations © J.S. Bendon & lizardwizard 2003.All photographs © Matthew Morton/Durrell Wildlife 2003.


can be congenital. However, one observer suggestedthat this might simply have been an abscess.

Yet another unusual observation was made ini-tially in nine hatchlings, all of which emerged from asingle, closely watched nest with various portions oftheir tails missing. Without a stabilizing tail, these ani-mals were a lot slower and clumsier, fishtailing fromside to side as they attempted to run. Consequently,they were much easier to catch (and photograph).The most likely culprit in this instance is a rat or a landcrab that burrowed into the nest, a conjecture sup-

ported by the fact that more hatchlings with missingtail portions were subsequently observed emergingfrom other nests.

Acknowledgments

Jenny Daltry (FFI), Matthias Goetz (Durrell WildlifeConservation Trust), Karen Graham (SedgwickCounty Zoo), Rick Hudson (Fort Worth Zoo), JeffLemm (San Diego Zoo), and Juliann Sweet respondedto inquiries regarding the possible causes for the var-ious anomalies.

Although few iguanas in the genusCtenosaura are actually black (thisoccurs only in some individual C. acan-thura and C. pectinata), one of the com-

mon names by which they are collectively knownis “Black Iguanas.” This name is actually mislead-ing when referring to some of the smaller, morecolorful species (e.g., C. alfredschmidti, C. defen-sor, and C. flavidorsalis). Another common nameused in English for species in this genus is “Spiny-tailed Iguanas.”

In Central America, Black Iguanas are called“Garrobos” or “Iguanas Negras.” Local inhabi-tants of the Islas de la Bahia (Utila, Roatan, andGuanaja), located off the northern coast ofHonduras, call the species that occur there,“Wishiwillies,” and, on the Isla de la Providencia,they are known as “Ishillies.”

The phylogenetic relationships and taxonomyof iguanas have provoked considerable controversyin recent years. In 1988, eight monophyletic2

groups were shown to exist within what had beenconsidered the Family Iguanidae. Shortly there-after, these were elevated to eight distinct families,largely because no evidence could be found for themonophyly of the entire family. This new classifi-cation was heavily criticized and eventuallyreversed on the basis of molecular genetic studies.However, after a very recent revision of the origi-nal study, which, among other things, nowshowed eleven monophyletic groups of iguanianlizards, the controversy continues. Some experts

believe that all pleurodont iguanian lizards shouldbe included in a single family (Iguanidae sensu lato)and that the monophyletic subgroupings shouldbe treated as subfamilies (e.g., the subfamilyIguaninae). In contrast, other experts think thatreal relationships are better portrayed by recogniz-ing eleven separate families, one of which includesthe “true” iguanas (Iguanidae sensu stricto).

Accordingly, lizards in the genus Ctenosauraare placed either in the subfamily Iguaninae (in theFamily Iguanidae sensu lato) or in the FamilyIguanidae (sensu stricto). Regardless of taxonomicrank, this group also includes the Marine Iguanas(Amblyrhynchus), Rock Iguanas (Cyclura), FijiIguanas (Brachylophus), Galapagos Land Iguanas(Conolophus), Desert Iguanas (Dipsosaurus), GreenIguanas (Iguana), and Chuckwallas (Sauromalus).The genus Ctenosaura is differentiated from theother genera within the family (or subfamily) bythe presence of an extremely long subocular scaleand whorls of enlarged, spiky scales on the tail.Ctenosaurs are most closely related to lizards inthe genera Iguana and Cyclura.

In recent years, tremendous advances havebeen made in the systematics of Ctenosaura.Seventeen species are currently recognized:C. acanthura, C. alfredschmidti, C. bakeri, C. clarki, C. conspicuosa, C. defensor, C. flavidor-salis, C. hemilopha, C. macrolopha, C. melanos-terna, C. nolascensis, C. oaxacana, C. oedirhina, C. palearis, C. pectinata, C. quinquecarinata, andC. similis. As recently as 1928, 13 species were rec-ognized, but subsequent workers determined thatthe species C. brachylopha, C. brevirostris, and C. parkeri were actually C. pectinata; and, morerecently, C. erythromelas was subsumed within C. defensor, reducing the number to only nine.More recently described species (the most recent in2001) have elevated the number to the current 17.

Black Iguanas: Name and Systematics1

Gunther Köhler1, AJ Gutman2, and Robert Powell3

1Forschungsinstitut Senckenberg, Frankfurt a/M., Germany; 2West Hartford, Connecticut, USA;

3Department of Biology, Avila University, Kansas City, Missouri, USA.


1 Adapted from Köhler, G. 2002. Schwarzleguane.Lebensweise, Pflege, Zucht. Herpeton Verlag Elke Köhler,Offenbach, Germany. Translated from German by AJGutman with additional commentary by Robert Powell.2 A “monophyletic” group includes an ancestor and all ofits descendants.

In the past, some species (C. bakeri, C. palearis, C. quinquecarinata, C. clarki, and C. defensor) were split off from Ctenosaura andplaced in a separate genus, Enyaliosaurus, but thisdivision is no longer widely recognized. Morerecent studies, based on genetic and morphologi-cal data, have presented an hypothesis of relation-ships within the genus Ctenosaura that recognizesthree monophyletic species groups designated assubgenera (Ctenosaura, Enyaliosaurus, andLoganiosaura).

The subgenus Ctenosaura includes the largerspecies, C. acanthura, C. conspicuosa, C. hemilopha,

C. macrolopha, C. nolascensis, C, pectinata, and C. similis, all of which lack the enlarged, heavilykeeled to spiky scales on the dorsal surface of theupper thigh (along with some other characters).The males in these species attain total lengths ofover 1 meter. The morphological criteria con-tained in the initial descriptions of three of theselarger species, C. acanthura, C. pectinata, and C. similis, were apparently quite variable andresulted in no definitive morphological charactersby which these three taxa could be distinguishedfrom one another. As recently as ten years ago,some authorities suggested that further study of


the status of the C. similis-acanthura-pectinatagroup would reveal that only a single speciesoccurred from southern Sinaloa (Mexico) all theway to Panama. This assumption proved to beincorrect. Morphological and genetic studies haveshown that these three species are all valid and canbe distinguished by scalation and coloration. Fieldstudies also have provided a more detailed pictureof the geographic distribution of the three speciesin the Isthmus of Tehuantepec, where their rangesapproach each other.

The subgenus Loganiosaura includes themedium-sized species, C. bakeri, C. melanosterna,C. oedirhina, and C. palearis, which attain maxi-mum total lengths of 80 cm. Of these species, C. bakeri, C. melanosterna, and C. palearis possessa well-developed pendulous dewlap, whereas C. oedirhina has a transverse gular fold. Thespecies in this subgenus also differ from otherctenosaurs in several skull characters, which rein-forces the view that this group is monophyletic.

The subgenus Enyaliosaurus includes the smallspecies, C. alfredschmidti, C. clarki, C. defensor, C. flavidorsalis, C. oaxacana, and C. quinquecari-nata. These species are characterized, among otherthings, by a reduction of the number of postmen-tal scales (from four to two) and possession of arelatively short, very spiky tail, which, at its widestpoint, is broader than it is high. Within this sub-genus, C. alfredschmidti, C. clarki, and C. defensorare notable for the reduction of the parietal eye,which is barely distinguishable with the naked eye(this trait is very evident in other species as the dis-tinct spot in the middle of the parietal scale on thetop of the head). These three species also havewhorls of well-developed, enlarged, spiky scales allthe way to the tips of their tails (approximately thedistal third of the tail is without whorls in the otherspecies).

ReferencesBailey, J.W. 1928. A revision of the lizards of the genus

Ctenosaura. Proc. U.S. Natl. Mus. 73:1–55.

Böhme, W. 1990. Buchbesprechung. Zschr. Zool. Syst. Evol. Forsch.28:315–316.

Buckley L.J. and R.W. Axtell. 1990. Ctenosaura palearis Stejneger.Cat. Amer. Amphib. Rept. (491):1–3.

Buckley L.J. and R.W. Axtell. 1997. Evidence for the specific statusof the Honduran lizards formerly referred to Ctenosaura palearis.(Reptilia: Squamata: Iguanidae). Copeia 1997:138–150.

de Queiroz, K. 1987. Phylogenetic systematics of iguanine lizards:a comparative osteological study. Univ. California. Publ. Zool.118:xii + 203 p.

de Queiroz, K. 1987. A new Spiny-tailed Iguana from Honduras,with comments on relationships within Ctenosaura(Squamata: Iguania). Copeia 1987:892–902.

de Queiroz, K. 1990. Ctenosaura bakeri Stejneger. Cat. Amer.Amphib. Rept. (465):1–2.

de Queiroz, K. 1990. Ctenosaura oedirhina de Queiroz. Cat. Amer.Amphib. Rept. (466):1–2.

Etheridge, R.E. 1982. Checklist of the iguanine and Malagasyiguanid lizards, pp. 7–37. In: G.M. Burghardt and A.S. Rand(eds.), Iguanas of the World. Their Behavior, Ecology, andConservation. Noyes Publ., Park Ridge, New Jersey.

Etheridge, R.E. and K. de Queiroz. 1988. A phylogeny ofIguanidae, pp. 283–368. In: R. Estes and G. Pregill (eds.),Phylogenetic Relationships of the Lizard Families: EssaysCommemorating Charles L. Camp. Stanford Univ. Press,Stanford, California.

Frost, D.R. and R. Etheridge. 1989. A phylogenetic analysis andtaxonomy of iguanian lizards (Reptilia: Squamata). Misc. Publ.Mus. Nat. Hist. Univ. Kansas 81:1–65.

Frost, D.R., R. Etheridge, D. Janies, and T.A. Titus. 2001. Totalevidence, sequence alignment, evolution of polychrotid lizards,and a reclassification of the Iguania (Squamata: Iguania).Amer. Mus. Novit. (3343):1–38.

Gicca, D.F. 1982. Enyaliosaurus clarki (Bailey). Cat. Amer. Amphib.Rept. (301):1–2.

Gicca, D.F. 1983. Enyaliosaurus quinquecarinata (Gray). Cat. Amer.Amphib. Rept. (329):1–2.

Grismer, L.L. 1999. An evolutionary classification of reptiles onislands in the Gulf of California, Mexico. Herpetologica55:446–469.

Köhler, G. 1993. Schwarze Leguane—Freilandbeobachtungen,Pflege und Zucht.. Herpeton Verlag, Offenbach.

Köhler, G. 1995. Eine neue Art der Gattung Ctenosaura (Sauria:Iguanidae) aus dem südlichen Campeche, Mexiko.Salamandra 31:1–14.

Köhler, G. 1996. Freilanduntersuchungen zur Morphologie,Verbreitung und Lebensweise des Yucatan-Schwarzleguans(Ctenosaura defensor). Salamandra 32:153–162.

Köhler, G. and C.R. Hasbun. 2001. A new species of Spiny-tailedIguana from Mexico formerly referred to Ctenosauraquinquecarinata (Gray 1842)(Reptilia: Squamata: Iguanidae).Senckenb. Biol. 81:257–267.

Köhler, G. and K. Klemmer. 1994. Eine neue Schwarzleguanartder Gattung Ctenosaura aus La Paz, Honduras. Salamandra30:197–208.

Köhler, G., W. Schroth, and B. Streit. 2000. Systematics of theCtenosaura group of lizards (Reptilia: Sauria: Iguanidae).Amphib.-Rept.. 21:177–191.

Köhler, G. and B. Streit. 1996. Notes on the systematic status ofthe taxa, acanthura, pectinata, and similis of the genusCtenosaura. Senckenb. Biol. 75:33–43.

Lazell, J.D. 1992. The family Iguanidae: disagreement with Frostand Etheridge (1989). Herpetol. Rev. 23:109–112.

Macy, J.R., A. Larsen, N.B. Ananjeva, and T.J. Papenfuss. 1997.Evolutionary shifts in three major structural features of themitochondrial genome among Iguanian lizards. J. Mol. Evol.44:660–674.

Schulte, J.A., II, J.R. Macy, A. Larsen, and T. Papenfuss. 1998.Molecular tests of phylogenetic taxonomies: a generalprocedure and example using four subfamilies of the lizardfamily Iguanidae. Mol. Phylogen. Evol. 10:367–376.


Readers will recognize that, in order tostay healthy, iguanas require UV-B(ultraviolet radiation in the “B” range)as much as they need the complex salads

we prepare for them daily. Without UV-B, the all-important mineral, calcium, cannot be effectivelyabsorbed — and calcium is as critical as nutritiousfood, water, and heat. In responsible husbandry,we must replicate the UV spectrum required forthe photochemical process involved in metaboliz-ing vitamin D3 (which mediates calcium absorp-tion) — while never forgetting that this will be oflittle importance if the diet, heat, etc. are less thanoptimal. Also remember that this is a discussion onwhat is best for reptiles, NOT humans.

Since extensive research published in reputablescientific journals has yet to be performed, the rec-ommendations that follow are based on logic,

atomic physics, engineering principles, existingknowledge about the natural habitat of iguanasand their biology, and my own personal experi-ence.

UV-B is part of the electromagnetic spectrum.This spectrum includes everything from radiowaves at one end to gamma rays on the other.Visible light is somewhere in the middle.Wavelengths are read in nanometers from 0.001nm (x-rays) to 100 billion nm (radio waves). Areptile bulb can be subjected to testing by an ultra-violet radiometer and spectrographs in order tomeasure precisely the wavelengths that are beingproduced.

The UV range is from 180–400 nm, the Brange specifically from 280–320 nm. However, weare most concerned with readings from 290–300nm. Why? This is the D-UV range, which triggersthe miracle of photo-biosynthesis and is responsi-ble for creating pre-vitamin D3 (cholecalciferol).We think that about 80% of this photochemicalreaction is triggered by ultraviolet waves in thisrange. Ingested vitamin D (7-dehydrocholesterolor 7-DHC) in the reptile’s skin absorbs the UV-Bprotons, which allows the photochemical reactionthat converts the 7-DHC to cholecalciferol. Thelatter is then converted in the liver to 25-hydrox-ycholecalciferol (25-HDCC). The final steptoward becoming biologically active vitamin D3(1,25- dihydroxycholecalciferol or 1,25 DHCC)takes place in the kidneys through the process ofthermal isomerization. Biologically active vitaminD3 is stored in the liver and kidneys and its pri-mary function is to regulate calcium metabolism.This gives you some idea of why reptiles sufferingfrom metabolic bone disease (the lack of sufficientD3 to metabolize calcium) also suffer from formsof liver and kidney disease.

Having mentioned reptilian skin, let’s quicklyconsider the amount of exposure of an animal to

H U S B A N D R Y82 IGUANA Volume 10, Number 3

Iguanas and Artificial Ultraviolet Light: How and How Much Made Simple —

Well, Not Exactly Simple…1

Bob MacCargarThe Greenhouse in New York

All photographs by the author.

Proper placement of bulbs will provide an iguana access toheat and UV light and permit proper thermoregulation byallowing the animal to move toward or away from the heatsource.

1 All references to brand names reflect the author’spersonal experiences and should not be construed as anendorsement by the IIS.

natural sunlight that is necessary for this chemicalreaction to take place. The assumption that a fewminutes a day is sufficient is based entirely on stud-ies with humans. Keep in mind that the protonsfrom the UV radiation have to penetrate the skindeep enough to reach the capillaries underneaththe skin in order to produce this reaction. Anyonewith experience around iguanas can appreciatehow tough and thick their hide is. Several hours aday of natural sun exposure is much more appro-priate for a large reptile.

However, the ultraviolet we need to supply forour animals must be “useable” (in other words, inthe D-UV range). How can we measure and beconfident that we have supplied our creatures withample amounts of D-UV? By far the most reliablemethod is to test specifically for blood levels of 25-HDCC (wild iguanas have levels of 175–275nmol/L of this form of D3 in their blood). Thistest can be performed by a qualified veterinarianand is the final say on whether we have providedoptimum husbandry for our iguanas. This is muchmore reliable than using a standard blood paneltest (which measures only calcium and phospho-rus levels). Even iguanas that have what appear tobe good blood panels have been found to be onthe low end of the active 25-HDCC level. Thishelps to explain why iguanas in the wild can drop20 feet out of a tree and hit the ground running,whereas stories of captive Green Iguanas falling 5feet and ending up with compound fractures arecommon.

Just what kind of ultraviolet levels are thesecreatures exposed to in their natural environment?The standard for measuring intensity of the UVspectrum is read in microwatts per square cen-timeter (µW/cm2). The USDA recently did astudy with an Ultraviolet Pyranometer and foundreadings of UV-B in Florida on June 1st thatreached 450 µW/cm2 (remember, however, thatonly a certain percentage of this falls into our“usable” D-UV). Hobbyists have the ability totake their own UV-B readings with a simple hand-held ultraviolet radiometer from Solarmeter(model 6.2). The measurements taken by theUSDA are extremely close to the readings that Ihave been taking for the last two years in a studyof reptile lamps and I have used this as a constantfor meter calibration.

Although iguanas may not be exposed to asmuch as 450 µW/cm2 on a continuous basis, they

will spend several hours a day exposed to relativelyhigh numbers. My studies, as well as those of oth-ers, have found that, even in the shade, global UV-B readings reach 30–50 µW/cm2. These numbersgive us an idea of the minimum and maximumultraviolet B exposure levels in nature.

Interestingly, another study has proven thatD3 biosynthesis is a naturally self-limiting process.Without getting too technical, this “safety valve”ensures that toxic levels of vitamin D3 are not cre-ated, and that the excess is broken back down intoinert ingredients (but please see the references atthe end of this article). Basically this means that aslong as we do not expose our iguanas to any moreUV than that to which they are exposed in theirnatural environment, we will stay within safeperimeters.

Two styles of reptile UV-B bulbs are available.One is the fluorescent tube and the other is themercury vapor (MV) reptile lamp. Both stylelamps use the heavy metal mercury as a catalyst forproducing ultraviolet radiation. An electricalcharge passing through liquid mercury excites themolecules until they vaporize (when the mercurycools, it resumes liquid form). In the fluorescenttube, the mercury must combine with high-gradephosphorus to achieve the ultraviolet results.

Over the past two years, I have studied failureand decay rates of mercury vapor lamps and con-ducted a general study of the major brand fluores-cent tube reptile bulbs. I am constantly asked if Ihave tested a certain fluorescent brand bulb byindividuals who have seen them listed inexpen-sively — in spite of the fact that, for many years, Ihave been telling people that quality UV is notcheap (unless we’re talking about the great andwondrous sun). In a recent conversation with


The low-cost hand-heldultraviolet radiometer fromSolarmeter (model 6.2)provides hobbyists with theability to take their own UV-B readings.

Voltarc Technologies, one of the largest manufac-turers of reptile fluorescent bulbs in the nation, theengineers confirmed the fact that money buys UV.

Production costs rise with more exacting spec-ifications (how much UV-B and where precisely itis to be delivered). In order to build a tube thatwill provide not only high UV-B readings, but onethat will generate output in the useable 290–300nm range, manufacturing tolerances have to bevery precise. Two different fluorescent tubes canemit equal amounts of total UV-B, yet one will doa much better job keeping your pet healthy thanthe other (even though both bulbs might havebeen manufactured by the same company, but fortwo different distributors’ specifications).

The best fluorescent tubes tested emit 12–15µW/cm2 at 12" after initial burning. A variety ofgood fluorescents (as well as some absolutely ter-rible ones) are on the market, but ZooMed 5.0 isbuilt to the most exacting tolerances according toall of the manufacturers with whom I have spoken.

The other choice in artificial UV is the mer-cury vapor reptile lamp, which comes in a varietyof styles and wattages. Anyone who has done reha-bilitation work has seen the effects of these bulbscompared to even the best fluorescent tubes. Whyis this the case? Do they emit huge amounts ofUV? Are they reliable? Is one brand better thananother?

Self-ballasted MV lamps suffer from a 50% fail-ure rate over the first six months and a 70% decayrate in total UV-B emitted. The best-selling styleMV bulb, the 160 watt FLOOD, emits much lessUV-B than stated on their endorsements afterdecay. These bulbs settle in at about 12 µW/cm2

at 12". Why, then, do we see such incredible resultswith MV lamps when compared to fluorescents?After all, two high-quality fluorescents will produce25–30 µW/cm2 at 12". The answer is simple: MV

lamps emit less total UV-B than fluorescents — butmore “usable” UV. Studies have shown that MVlamps produce the same percentages of D-UV (andUV-A, which is another subject) as a percent oftotal energy emitted as the sun.

Self-ballasted SPOT-style lamps produce muchhigher UV readings than any other reptile bulb onthe market. They still are subject to the failure ratesof all self-ballasted MV lamps. These lamps settlein at about 100–150 µW/cm2 (at 12"), but havea much narrower disbursements of UV-B. Theseare excellent rehabilitation bulbs for treating ani-mals suffering from severe metabolic bone disease.Regarding concerns that iguanas under this styleof lamp will need “goggles” to prevent blindnessfrom “excessive” UV exposure, remember theexposure levels in their natural environment. Wehave five iguanas that have spent two years exclu-sively under mercury vapor SPOT lamps with nonegative impact on vision. However, the distancefrom any MV lamp to the basking area must beregulated in order to provide optimum tempera-tures, regardless of any distances stated by the dis-tributor!


ZooMed 5.0 fluorescent tubes are built to the mostexacting tolerances.

Erving, a rescued Green Iguana, made a remarkablerecovery following six months of therapy under UV lamps(top). Note the severe spinal scoliosis (center) and evidenceof severe metabolic bone disease (bottom) when he wasinitially rescued.

At this time I recommend only 100- and 160-watt T-Rex UV Heat, 160-watt ZooMedPowersun, and the Westron Lighting 60-watt in-line ballast MV lamps. Others performed poorly inmy tests. A new 60-watt in-line ballasted MV lampfrom Westron Lighting produces excellent UV-Bafter decay, and without the problem of failure (noself-ballast to fail). Its primary drawback is limitedheat production, although this could be an assetfor keepers of smaller reptiles kept in glass habitats.It also generates poor-quality visible light.According to the manufacturer, these limitationsof the current bulb will not apply to models thatwill soon be available.

So, what is best for iguanas? I recommend30–50 µW/cm2 at 12" for 8–12 hours per day(comparable to minimal natural exposures in thewild). The best way to realize this number is to usean ultraviolet radiometer (such as the Solar Meter6.2 hand-held version; see references) to measurethe amount of UV-B available to your iguana. TheZooMed 5.0 fluorescent lamps are at the top ofthe ladder in terms of meeting these specificrequirements (i.e., the best “usable” UV-B). Useat least two of these lamps to achieve the desired

exposure levels. The 100- or 160-watt T-RexFLOOD lamps or the 160 watt ZooMed FLOODlamp also may be used in conjunction with aZooMed 5.0 fluorescent tube. If you do not havea meter, you should rely on the security providedby the quality of the 5.0 bulbs. Alternately, T-RexSPOT bulbs (100- and 160-watt) will emit50–150 µW/cm2 (at 12" after break-in) — aslong as they burn. No other supplemental UV-Bsource is necessary with these lamps. Finally, theWestron Lighting MV in-line ballast lamp can pro-duce 30–50+ µW/cm2 after break-in. This lampwill need additional heat and full-spectrum com-plements, but should not fail as quickly or fre-quently as other MV lamps.

References and Resources (listed by topic)

25 HDCC blood level study with radiograph (and much moreinformation on ultraviolet bulbs), http://www.myiguana.com

25 HDCC vitamin D Test, University of Michigan, Animal HealthDiagnostic Laboratory, PO Box 30076, Lansing, MI 48909(517-353-0621)

Calcium physiology, Bogoslavsky, B. 2002. Calcium metabolism iniguanas. Iguana Times (Journal of the International IguanaSociety) 9(1&2):32–34.

Electromagnetic spectrum,http://csep10.phys.utk.edu/astr162/lect/light/spectrum.html

Photochemistry and biology, http://www.photobiology.com

Ultraviolet radiometers, www.solarmeter.com

UV-B and D3, http://home.att.net/~chameleons/zoomedUV-B.html

Vitamin D discussion, “The Merck Manual,” 17th ed., p. 35


The Westron Lighting MV in-line ballast lamp requirescomplementary heat and full-spectrum lighting but outlastsother MV lighting.

ZooMed’s Powersun 160-Watt lamp is anotherlamp that performed wellwhen tested.

The T-Rex UV Heat 100- and160-Watt lamps are amongthose recommended.

Ihad not spoken with Henry Fitch in quitesome time, and not since his wife, Virginia,had passed away in December 2002 — butBob Powell had requested that we do some-

thing for Iguana, and I wanted to say “hello” andto see if he was interested in collaborating onsomething for that publication.

I called in the morning. Henry answered thephone right away and actually sounded pleased tohear from me. We reminisced briefly, and the sub-sequent conversation went something like this:

RWH: You’re 92 now, right?

HSF (chuckling): No, 93.

RWH: So, how’re you feeling?

HSF: I’m pretty crippled-up and don’t getaround much.

RWH: Sorry to hear that. I guess you’re notgetting into the field then?

HSF: Well, I’m just starting a radio telemetryproject with Timber Rattlesnakes. We putthe transmitters in the snakes this morning.

RWH: (Silent incredulity).

And that pretty much tells you what HenryFitch is all about.

———

When I first arrived at the University of Kansasas an incredibly naive undergraduate in 1967, Ididn’t know Henry Fitch existed, much less thathe was doing exactly the kind of research Idreamed of doing myself. The attraction for me atKansas was Bill Duellman and his amazingly activeprogram of herpetology in the American tropics.But now I had access to a remarkable library(Duellman’s), and I somehow found some of

P R O F I L E86 IGUANA Volume 10, Number 3

Henry S. Fitch: A Legendary Passion for Natural History

Robert W. HendersonMilwaukee Public Museum, Milwaukee, Wisconsin

Henry Fitch with a Boa constrictor in Chiapas, Mexico(February, 1972). Photograph by R.W. Henderson.

Fitch’s publications. It was a real epiphany to real-ize that someone was doing (and had been doingfor many years) exactly what I wanted to do. I devoured his classic Autecology of the Copperhead(not knowing what “autecology” meant when Istarted to read it) and anything else snake-related.

Dr. Fitch had been on sabbatical with his fam-ily in Costa Rica, and he returned to Kansas in thefall of 1968. I enrolled in his vertebrate natural his-tory course and, although I don’t recall the details,I’m sure I forced myself upon him very early on. I became a frequent visitor to the natural historyreservation that now bears his name, often arrivingunannounced on a weekend afternoon when hewas trying to relax and spend time with his family.Often he allowed me to make runs, either withhim or on my own, to check the tins under whichcongregated handfulls of Diadophis punctatus(Ringneck Snakes). I think he quickly realized thatwhat I lacked in IQ points I easily compensated forin enthusiasm. He encouraged me to initiateresearch as an undergrad, and eventually becamemy advisor when I entered graduate school atKansas. While still at Kansas, and long after I hadmoved on to the Milwaukee Public Museum, wespent time in the field. We worked on Anolis inMexico, Great Corn Island off the coast ofNicaragua, and the Dominican Republic, and oniguana (Ctenosaura and Iguana) ecology, behav-

ior, and conservation in Belize and Nicaragua. Wedescribed new species of anoles, presented detailson the social behavior of anoles and iguanids, anddocumented the exploitation of iguanas in CentralAmerica (see article on p. 63).

Cumulatively, we’ve spent months together inthe field. One of my most vivid memories wasspending six weeks traveling in southern Mexico inhis truck and camper. Every dinner for those sixweeks consisted of boiled potatoes and fried Spam.I still like Spam, but I cannot stomach a boiledpotato. While in Mexico, we ate cold cereal withpowdered milk for breakfast each morning. Onemorning we were out of milk and I suggested wehave sandwiches. Dr. Fitch looked incredulous andreplied: “For breakfast?” He opted to have hiscereal with plain water and admitted that it wasn’tvery good. Henry is a creature of habit.

Long before I knew him, Henry Fitch wasalready a living legend. His long-term field studiesof snake populations at the University of Kansasfield station are data-rich and amazingly thorough.He brought mark and recapture studies of snakesinto the forefront of herpetology, and he was atrue pioneer in the use of radio-telemetry to studymovement ecology and habitat use by a variety ofsnake species. In 1999, he published his landmarkbook, A Kansas Snake Community: Compositionand Changes Over 50 Years (Krieger Publ. Co.).


Henry Fitch and Bob Henderson measuring a Haitian Boa (Epicrates striatus) at Paraiso, Barahona, Dominican Republic (March1985). Photographs by N.E. Kraucunas.

The likelihood of another researcher being in aprofessional situation to conduct a similar study,much less to have the drive, enthusiasm, and goodhealth to do it, are virtually nil.

It wasn’t until 1965 at the age of 55 that Dr.Fitch finally made it to the tropics. Once there,however, tropical fieldwork became his passion.He eventually spent a solid year in Costa Rica withhis family in order to study the population ecologyof a wide variety of lizards. After that, any year(especially during the long Kansas winters) that didnot include at least one trip to somewhere (any-where!) in Central or South America or the WestIndies was considered disappointing. He was espe-cially interested in Anolis ecology and behavior,and has published on a wide range of topics relatedto their biology, including taxonomy, behavior,and ecology.

In 1976, we were awarded a small grant tostudy iguana exploitation in Central America. Weworked together in Nicaragua for about six weeksbefore I returned to the U.S. At about 65 years ofage, when other folks are thinking about retire-ment and taking it easy, Henry set-off on his ownto visit markets all over Central America in orderto document the extent of exploitation ofCtenosaura similis and Iguana iguana. By inter-viewing vendors and hunters, he was able to mapa network of iguana black marketeers, with truck-loads of the lizards going across borders from onecountry to another. The next year, we spent sev-eral weeks in the Belize City cemetery watchingthe interactions of a healthy population of C. similis (known locally as Wish-willies) usingtombstones for basking sites, look-outs for territo-rial males, and for retreats.

During all of the time I spent with the man,whether on the natural history reservation inKansas or somewhere in the tropics, I never sawhis enthusiasm wane. He was always ready for anew adventure, to collect the next lizard, or recordmore data. Although I’m 35 years younger thanHenry, I would welcome an occasional break inthe day to put my feet up and have a cold drink.Dr. Fitch could do that too, but only for about 11minutes. Then it was: “Bob. Let’s go to such andsuch a place and see what we can find. It’s thelocality for such and such a critter.” — and, ofcourse I would go because I knew that if I didn’tI was going to miss seeing something worthwhile

and an opportunity to learn something fromsomeone who has forgotten more about lizardsand snakes than I’ll ever know.

I’m proud to have been Henry Fitch’s studentand frequent collaborator, and I can’t tell you howmuch it pleases me that a 93-year-old man who(supposedly) can barely get around is excited aboutinitiating a new project with Timber Rattlesnakes.Decades after it no longer mattered to his profes-sional “status,” Dr. Fitch has continued to conductnatural history studies in Kansas and the Americantropics and to publish in scientific journals. He con-tinues to do it because his desire to add to ourknowledge of squamate ecology has never waned.His life is testimony to passion and doing what youcan to satisfy that passion.


Henry Fitch weighing a Ctenosaura similis in Nicaragua(February 1976). Photograph by R.W. Henderson.

The Whorl-tailed Iguanas (Cyclura Harl.[translator’s note: “Harl.” Refers toRichard Harlan, an American paleontolo-gist and comparative anatomist who

described many American and “exotic” species ofamphibians and reptiles]) are distinguished fromCommon Iguanas [= Iguana spp.] by their denti-tion, the smaller dewlap, and the more weaklydeveloped gular fold. Pattern and scalation is sim-ilar to that of other iguanas, except that the dorsalsurface of the tail is distinguished by a ring ofenlarged scales following three or four rows of typ-ical scales. Although these enlarged scales are notparticularly long, their collective tips form a whorl-like band of erect spikes around the tail. The dor-sal crest may be interrupted in the shoulder andhip regions. The teeth, which appear to increase innumber with age, are not serrated, but are three-sided, as in Common Iguanas, and those on the

pterygoid and sphenoid are small but numerous.Femoral pores are distinct and form a long row oneach leg. All species are terrestrial and are foundprimarily in sandy areas where they live in burrowswhich they excavate for themselves.

The best-known species is Cyclura lophomaGosse [this “species” is now known to consist ofseveral distinct taxa, including C. collei, C. nubila,and C. cychlura; Philip Gosse was the author of thedelightful A Naturalist’s Sojourn in Jamaica, pub-lished in 1851] of Cuba, Jamaica, and theBahamas. This form may reach a total length of

H I S T O R I C A L P E R S P E C T I V E89IGUANASeptember 2003

Whorl-tailed Iguanas*

Franz Werner

* excerpted and translated from Franz Werner (1913. Die Lurche und Kriechtiere von Alfred Brehm. ZweiterBand: Kriechtiere (Schuppenkriechtiere). In O. z. Strassen(ed.), Brehms Tierleben. Allgemeine Kunde des Tierreichs.4th ed. Bibliographisches Institut, Leipzig and Wien.Translated by R. Powell.

Adult male Jamaican Iguana (Cyclura collei); at the time Werner wrote this article, the Jamaican species plus populations inCuba (C. nubila ) and the Bahamas (C. cychlura) were all known as C. lophoma. Photograph by Jeff Lemm.

1.2–1.3 m, of which the tail comprises about 70cm, and is distinctive because of its relatively lowdorsal crest, which is composed of linked, some-what sickle-shaped spines, and which may be con-tinuous at the nape and base of the tail or distinctlyseparated at one or both places. The snout is cov-ered by three pairs of large, multi-sided, tubercu-late plates that are separated from one another bysmall scales. Plates of variable size, those that aremedially located the most prominent, characterizethe forehead, and on each side of the lower jaw isa row of large, quadrangular, heavily keeled scales.Overall color of the body and limbs ranges fromslate gray to brownishgreen. A few diagonallines passing over theshoulder and severalbroad crossbands thatextend from the dorsalcrest to the belly aredark olive-brown. Thetail is circled by regu-larly spaced light anddark olive-green bands.

According toGosse, Whorl-tailedIguanas live in only afew places on Jamaica.They were relativelycommon on the lime-stone hills that extendfrom Kingston harborto the so-called Goat Island and are well knownfor the large numbers of feral goats, pigs, andchickens that live there. Otherwise, one may findthese iguanas with some regularity in the flats thatseparate the coastal hills from higher mountains ofthe interior, but only in areas that feature largenumbers of old, hollow tree trunks. These lizardsappear to have no particular fondness for water,although, like their relatives, they know how toswim rather well. A fortunate circumstance madeit possible to learn more about the Whorl-tail thanmany of its relatives. Two individuals lived in anold Acacia on the estate of a Mr. Minot for 16months, providing him with the opportunity toobserve them for an extended period of time. Afriend of the owner had fortuitously discovered thetwo animals, but a blow with a riding crop fright-ened them so much that for weeks thereafter they

emerged from their lair in only the most surrepti-tious fashion and immediately sought refuge in thehollow tree when any human approached tooclosely. Subsequently, Minot forbade any furtherdisturbance of the animals and, over time, they for-got their fright and eventually became so tame thatthey consented to regular and proximate visits bytheir landlord. As the day warmed, one of the ani-mals would emerge from its tree hollow and hangfrom the bark or crawl out on a thin, dry branchto bask in the sun. It would often spend the entireday doing little else, with nary a care for the rest ofits environment. Minot never saw it hunting

insects and only oncedid he surprise a lizardwhile it was eating.This occurred whenthe sun broke throughthe dark clouds after aheavy rain and the veg-etation had largelydried. One of theWhorl-tails left thetree, covered about 10m of ground with slowsteps, moving only oneleg at a time, until itapproached a stand ofso-called Guinea-grass.Tearing off mouthfulsat a time, it swallowedlarge masses without

any trouble. Startled by the observer whom it sud-denly noticed, it rushed back to the tree, not somuch running or walking as moving with a rapidseries of frog-like hops. Once there, it ducked intoits hole and quickly disappeared from sight.

Minot made particular note of the fact that thefleeing Whorl-tail showed no inclination to seek outwater, as other iguanas are wont to do under simi-lar circumstances. As a matter of fact, these Jamaicananimals showed little interest in water of any kindand seemed to tolerate even the driest conditionswithout drinking. The two inhabitants of the treewere obviously a pair, since they differed from oneanother in size and coloration. They also lived har-moniously together, but they were never seen outof the burrow at the same time. Eventually, a good-for-nothing youngster ended the observations, lay-ing near the tree and heartlessly shooting the two


Head of a Rhinoceros Iguana, Metopocerus cornutus Daud.Photograph by L. Medland, F.Z.S. – Finchley, N.

harmless creatures one after the other. Minotopened the carcasses of the shamefully murderedanimals and found both stomachs filled with thepreviously mentioned grass.

Barbour [Thomas Barbour, an American her-petologist who traveled widely throughout theWest Indies during the first half of the 20th cen-tury] recently noted that the Whorl-tail has beenbrought to the verge of extinction by the presenceof the introduced Mongoose (Herpestes griseus[= Herpestes javanicus]) and is now found only onGoat Island, which the interloper has yet to reach.

Although generally retiring, preferring toretreat in the face of humans whenever possible,the Whorl-tail is quitebrave and quite capableof defending itself in anemergency. The tail is aweapon that shouldnot be underestimatedand which can be usedwith great effect. Easilyaroused like all iguanas,the Whorl-tail willrespond with consider-able wrath when cor-nered, inflating itself,erecting the dorsalcrest, gaping to showits sharp teeth, staringdarkly at its antagonist,and readying itself foran attack. If one con-tinues to threaten thebeast, it will turn around quickly and deliver ahefty blow with its tail, and then immediately turnthe other way in anticipation of delivering anotherstrike from the other side. Hill [an unknown per-son] had been alerted by natives to the dangerposed by this animal and had been clearly warnedagainst a careless approach. The spikes on thepowerful tail are sufficiently sharp that the animalcan readily inflict a most painful wound. Dogs thatapproach one carelessly often are severely injured.

Whorl-tails, possibly as a consequence of theirdiet, have such an unpleasant odor that even antsrefuse to touch them — or at least to strip the fleshfrom a carcass thrown on their colony. These ani-mals appear not to be hunted as eagerly as theirrelatives in Mexico.

The Rhinoceros Iguana, Metopocerus cornutusDaud. [Metopocerus is now considered synony-mous with Cyclura; “Daud.” refers to François-Marie Daudin, a French naturalist who wroteHistoire Naturelle des Quadrupèdes Ovipares in1800, in which he formally described theRhinoceros Iguana], is a mighty beast found inSan Domingo [a reference to Hispaniola and,more precisely, to the Dominican Republic]. Itresembles the Green Iguana (Iguana tuberculata[= Iguana iguana]) in having teeth with serratedcrowns and the Spiny-tailed Iguanas [= Cteno-saura spp.] of Middle America in color, size, build,and lifestyle — but is readily distinguished by the

three large, cone-shaped, horn-like scales on thetop of the snout of adult males. Old males also arecharacterized by their bulging jowls and exceed-ingly large heads and make a considerable impres-sion that is enhanced further by their proud pos-ture and lively eyes. The powerful bite and thestrong, spiky tail make the Rhinoceros Iguana aworthy opponent.

Rhinoceros Iguanas are overwhelmingly terres-trial, consume both vegetable and animal matter,and in captivity can be readily maintained on a dietof fruit, juicy leaves, and strips of raw meat.Ditmars [see the appended biographical sketch]indicated that they even overpower rats and youngchickens, shaking larger prey until it is torn intopieces. Given enough space and heat, these lizards


Rhinoceros Iguana.


B I O G R A P H I C A L S K E T C H : Raymond L. Ditmars

(1876–1942)Raymond Ditmars was America’s first great popularizer of reptiles. He was born in Newark, New Jersey

and caught his first snakes at age 12 at Gravesend Bay near Brooklyn, New York. In 1893, Ditmars was hiredby the entomology department at the American Museum of Natural History, but resigned in 1897 to takehigher paying positions unrelated to his interest in animals. In 1898, as a reporter for The New York Times,one of his beats was the newly-formed New York Zoological Society. Shortly after the zoo opened, Ditmars,barely 23 years old, was hired as Assistant Curator of Reptiles and his private collection become the nucleusof the zoo’s reptile displays.

Ditmars published eight herpetological books, plus a dozen more about other animals or tales of his work or travels. The Reptile Book, published in 1907 (an expanded edition was published in 1930) established his reputation. This volume and several others (Reptiles of the World, 1910, revised 1933; Snakesof the World, 1931; Reptiles of North America, 1936; and Field Book of American Snakes, 1939) had anenormous public influence, kindling a widespread interest in reptilesand resulting in several generations of professional herpetologists,including many still active today, who can trace their interest to oneor more of Ditmars’ books.

Ditmars was often dismissed as a mere showman, mainlybecause he wrote highly popular books for the general public, but he also made substantive scientific and public contributions.He pioneered techniques for treating diseases of reptiles, he was a major organizer of the first effort to establishan antivenin institute in America, he produced influ-ential nature films, and he led the development of theBronx Zoo into the first rank of the world’s zoos.

Source: Adler, K. 1989. Herpetologists of the past, pp. 5–141. In K. Adler (ed.), Contributions to the History of Herpetology.Society for the Study of Amphibians and Reptiles, Contributions to Herpetology, Number 5. Ithaca, New York.

Raymond L. Ditmars (photographcourtesty of Kraig Adler).

do well in captivity and have become staples inzoos that maintain collections of reptiles.Rhinoceros Iguanas are lively, shy, and careful, anddo not become tame in captivity for some time. Amale, kept by Werner [the author] in a cage forseveral years, gaped when approached, exposing itsdark purple throat, but did not bite, instead

retreated farther into the cage and, only whenapproached even more closely, did it turn andstrike with its tail. When removed from its cage, itexhibited an extraordinary strength, resisted des-perately, and flailed its tail violently.

Hodge, Karim V. D., Ellen J. Censky, andRobert Powell. 2003. The Reptiles andAmphibians of Anguilla, British West Indies.Anguilla National Trust, The Valley. Softcover,$15. 72 pp., over 90 illustrations, nearly all incolor.

This delightful volume is the second in a series ofguides published by the Anguilla National Trust ontheir native flora and fauna. The nation of Anguilla iscomposed of one main and several small islands, whichcollectively form the northern end of the Lesser Antilles(the island chain that extends from the Puerto RicoBank to South America). Unlike most other LesserAntillean islands, which are mountainous, much ofAnguilla is low-lying and dry. Rather than accumulatingon the surface, rainfall tends to seep through the karstlimestone from which the islands are formed. Thus thenative forest must be both drought and hurricane resis-tant. What remains, after extensive foraging by theubiquitous goats, is mostly degraded scrub forest, yetthe islands still host over 500 varieties of plants and, sur-prisingly, over 20 species of reptiles.

The guidebook features species accounts ofAnguilla’s three frogs or toads (all introduced), four seaturtles, one terrestrial turtle (possibly introduced), threesnakes (two introduced), and 13 lizards. The lizardaccounts address two iguanas (one native and one intro-duced), three ameivas (all native and two endemic), twoanoles (one native and one introduced), a skink(native), and five geckos (one endemic species has notbeen formally described and one species is probablyintroduced). These subjects range in size from massiveLeatherback Sea Turtles to tiny Dwarf Geckos. My per-sonal favorite is the Lesser Antillean Iguana, Iguanadelicatissima, of which an excellent photograph byGlenn Gerber graces the cover. Of particular interest isthe conservation status of this animal. Although threat-ened throughout its range, the Anguillian population iscritically endangered. The usual factors (habitat destruc-tion and competition with feral animals) are responsi-ble, but these may be exacerbated by potential compe-tition and hybridization with Green Iguanas,introduced as pets or on a raft of floating debris thatarrived on Anguilla shortly after Hurricane Luis in1995.

This book is an excellent resource for both herpenthusiasts and newcomers. A brief description of thenature of reptiles and amphibians and an excellent glos-sary of terms provide help for the uninitiated. Eachspecies account includes colorful photographs and illus-trations along with notes on the distribution, origin,biology, and conservation status of each animal. I par-ticularly liked the sections on the origins of the various

species. Aside from mentioning the usual methods ofintroduction (e.g., rafting, land bridges, or humanmediation), one notation suggested that some geckosmay have been “purposely introduced by sorcerers tosome islands in order to terrify their subjects.”

Most significantly, a section on conservationdescribes the reasons why many species have becomethreatened and suggests measures for improving the sit-uation. Among other things, the authors suggest sus-tainable land use and the conservation and enhance-ment of animal habitat. Perhaps the most importantmeasure suggested, however, is increasing public aware-ness, and, in this, the book itself provides an excellentresource. In fact, the Trust intends to utilize it inAnguillian schools to enhance an awareness and appre-ciation of the nation’s natural resources.

Because printing costs were covered by a grantfrom the United Kingdom’s Foreign andCommonwealth Office Environmental Fund forOverseas Territories, all proceeds from the sale of thebook will support the efforts of the Anguilla NationalTrust to preserve that nation’s natural and cultural her-itage. Copies may be ordered directly from the Trust(Anguilla National Trust, P.O. Box 1234, The Valley,Anguilla, B.W.I.; [email protected]) or fromBibliomania! (http://www.herplit.com/) for US $15(EC $40) plus shipping and handling.

AJ Gutman

B O O K R E V I E W93IGUANASeptember 2003

First Breeding ofCyclura nubilanubila andCyclura n. caymanensis x C. lewisiin the UnitedKingdomRoger Lamb of Solihull,West Midlands, UK has 18years experience in keepingand breeding three gener-ations of Green Iguanas(Iguana iguana) and onegeneration of RhinocerosIguanas (Cyclura cornuta).Since 1997, Lamb also hasbeen working with twopairs of Cuban Iguanas (C. nubila nubila) and apair of hybrid C. lewisi ×C. n. caymanensis.

The Cuban Iguanasproduced their first fertileeggs in 2002, with threehatchlings successfullyemerging. The femalehybrid laid seven eggs, two of which were fertile.Following a 78-dayincubation, the hatchlings

had pipped the shell andtheir heads were out. One egg, much to Lamb’ssurprise, contained twins.

Evidence ofGreen IguanaReproduction in AnguillaKarim V. D. Hodge,Environmental Officer,Government of Anguilla,reported the discovery of ahatchling Green Iguana(Iguana iguana) on 30July at East End Village.This suggests that repro-duction of Green Iguanas isoccurring in the wild.

The hatchling was foundless than a mile from thesite where Green Iguanaslanded on a raft of floatingdebris after Hurricane Luisin September 1995.Consequently, the possibil-ity exists that survivors ofthat event may still live inthe area and are reproduc-ing. However, anotherAnguillian population ofGreen Iguanas near the

southwestern end of theisland is known to be the result of escaped (orintentionally released) pets.The fact that this hatchlingcould be a product of suchanimals moving or beingcarried into the area cannotbe ruled out.

Regardless of the source,active reproduction ofGreen Iguanas in Anguillaposes a potentially seriousthreat to the native,critically endangeredpopulation of LesserAntillean Iguanas (Iguanadelicatissima) by means ofcompetition for limitedresources or pollution ofthe native gene poolthrough hybridization.

Hodge currently is attempt-ing to assess the extent ofthis new threat by searchingfor other hatchlings andadults in the area where thisindividual was found.

Banner Year forBlue IguanaHatchlings(Cyclura lewisi)Fred Burton, Director ofthe Blue Iguana RecoveryProgramme, reported that82 eggs were collected thisyear. Burton attributedmuch of the success toWarden D. Ebanks, whoclosely monitored breedingand nesting sites at theQueen Elizabeth II BotanicGarden on Grand CaymanIsland. In addition, Ebanksmaintains a vegetable gardendesigned to meet the dietaryneeds of iguanas held at thebreeding facility in the park.The position was fundedwith donations from MaplesFinance in Cayman and theInternational IguanaFoundation.

Eggs were exhumed,placed in an incubator, and are being carefullymonitored by Burton, whonoted that the due date forthe first batch of nine eggsis imminent. The success

of the captive breedingprogram emphasizes theneed to find a safe andsuitable site for thehatchlings to grow up in the wild.

— Caymanian Compass, 6 August 2003

South Florida ison the Menu forHungry IguanasWith few natural predatorsin South Florida, reptilianimmigrants from Mexicoand Central and SouthAmerica are free to romp— and chomp — whereverand whatever they please.South Florida has had asmall population of wildiguanas for many years, butonly recently have theirnumbers grown out ofcontrol. So far, plant lovershave not found an effectiveway to stop them.

Valerie Cassidy, presidentof the Gardens at CrandonPark Foundation, estimatedthat several dozen Spiny-tailed Iguanas (Ctenosauraspp.) gobbled up severalthousand dollars worth ofplants in a 48-hour pig-outin the flower beds at theCrandon Gardens duckpond on Key Biscayne.“They’re huge. They’rehungry. And they’re eatingparts of South Florida.”

I G U A N A N E W S B R I E F S94 IGUANA Volume 10, Number 3

Hatchling Green Iguana (Iguana iguana) found at East End Village,Anguilla. Photograph by Karim V. D. Hodge.

Twin hybrid Cyclura lewisi x C. nubila caymanensis emerging fromthe egg. Photograph by Roger Lamb. Hatchling Blue Iguana (Cyclura

lewisi) emerging from an eggat the Blue Iguana RecoveryProgramme at QueenElizabeth II Botanic Garden,Grand Cayman. Photographby Fred Burton.

At Fairchild TropicalGarden, iguanas feast onthe filet mignons andcaviars of the plant world— rare and expensive plantssometimes brought fromthe other side of the world— much to the frustrationof the horticulturists.“They like the prettiest andrarest flowers, so they’reobviously creatures of somediscriminating taste,” saidMike Maunder, Fairchild’sdirector of horticulture.Park groundskeepers havetried many tactics to driveoff the lizards, includingspraying the plants with achemical deer repellent, but the reptiles, who atCrandon happily gobble uptoxic Philodendron leaves,were not deterred.

The lizards have only beennibbling on BrowardCounty, with smallpopulations at FlamingoGardens, Snyder Park, The Bonnet House, andEasterlin Park in OaklandPark. “Right now we’reOK,” said Ross Dovey,director of Easterlin Park.“But it’s a potentialproblem I’m afraid willonly get worse.”

— Daphne Duret, Miami Herald,

31 July 2003

Anegada IguanaPopulationAssessment 2003The IUCN IguanaSpecialist Group (ISG)conducted a populationassessment of Cyclurapinguis (Stout Iguana) onAnegada Island, BritishVirgin Islands from 14–29June (see also IGUANA10(2), June 2003). TheInternational IguanaFoundation (IIF),Institute of Museum andLibrary Services (IMLS),and private contributionsby John Binns, JoeBurgess, and GeorgeWaters funded the survey.

Anegada is approximately16 km long and 2–3.5 km

wide (about 39 km2).Although not a large island,it presented a significantchallenge to the smallteam’s ability to properlysurvey the island in alimited amount of time.The strategy was to focuson the western half of theisland. Any remaining timewould be devoted tosurveying the area betweenthe central ponds and thesouthern, developed coastand some areas east of theSettlement. Results of East

End surveys conducted byGlenn Gerber, Binns, andothers in 2001 wouldround out the data neededto estimate the totalpopulation size.

Formal results of the surveywill be presented at theannual ISG meeting in theTurks and Caicos Islandsthis November. However,the status of C. pinguis onAnegada remains criticaland population numbersappear to be in a continu-

ing decline. The coreiguana area, Bone andWindlass bights, containedbulldozed access cuts,which have destroyed anumber of known iguanaburrows. Citron Bush, thesite of Michael Carey’sresearch in 1968, in whichhe described healthypopulations of C. pinguis,consisted solely of heavilydisturbed habitat, withmost of the damage causedby feral cattle and goats.Neither iguanas nor signsthereof were found.Overall, iguana sightingswere few, especiallydisappointing to thoseteam members who hadnever seen Stout Iguanasand who had traveled farand worked hard for nomore than a fleeting glanceof a disappearing shadow.

Current plans call for alimited release of head-started C. pinguis, presentlyheld at the Anegada head-starting facility, on FallenJerusalem, a small island inthe British Virgin Islands. If successful, this wouldprovide a further hedgeagainst the extinction ofthis species. As a part ofthat relocation plan,Gerber, Kelly Bradley, andLee Pagni conducted ahabitat survey on FallenJerusalem after completingthe survey on Anegada.They found no feralmammals and the islandappears to be an ideallocation for release.

I G U A N A N E W S B R I E F S95IGUANASeptember 2003

One of the few photographsof Cyclura pinguis takenduring the survey. Photographby Joe Burgess.

The 2003 survey team: (back row, left to right) George Waters (IIS),Dr. Glenn Gerber (SDZ/CRES/ISG), and John Binns (IRCF/ISG/IIS);(front row) Roberto Maria (ZooDom/ISG), Kelly Bradley (DallasZoo/ISG), Tarren Wagener (Ft. Worth Zoo), and Joe Burgess (IIS).Lee Pagni (SDZ/ISG/IIS; not shown) periodically assisted in thesurvey, but his primary agenda was to interview local residentsregarding Stout Iguanas in order to develop public awareness andschool educational programs. Lee Vanterpool (National Parks Trust,BVI/Head Head-starting Keeper) and Sallie Davis, a member of anarcheological team conducting research on the East End, alsoassisted the team. Photograph by Tarren Wagener.

Each morning, the team members surveyed designated transectareas, battling dense brush and cacti in 100° heat. The habitatshown in this photograph is typical of what team membersencountered during the survey. Left: Joe Burgess, center: SallieDavis, right: Roberto Maria. Photograph by John Binns.

I.I.S. Board of DirectorsJoseph Wasilewski, PresidentHomestead, FL

Joseph Burgess, Vice-presidentJacksonville, FL

A.C. Echternacht, Ph.D., TreasurerUniversity of Tennesee, Knoxville, TN

AJ Gutman, Executive Secretary andAssociate Editor of IGUANAWest Harford, CT

John & Sandy Binns, Directors, AssociateEditor of IGUANA and Internet OperationsInternational Reptile Conservation Foundation,San Jose, CA

Gunther Köhler, Ph.D. and Elke Köhler,DirectorsForschungsinstitut und NaturmuseumSenckenberg & Herpeton, Frankfurt, Germany

Robert Powell, Ph.D., Director and Editorof IGUANAAvila University, Kansas City, MO

I.I.S. Editorial/AdvisoryBoardAllison C. Alberts, Ph.D.IUCN/SSC Iguana Specialist Group andZoological Society of San Diego, San Diego, CA

John Bendon, Illustrator for IGUANABath, England

Gordon M. Burghardt, Ph.D.University of Tennessee, Knoxville, TN

Frederic J. BurtonBlue Iguana Recovery Program, Grand Cayman,Cayman Islands

Jorge FerrariOrganización de Rescate y Protección deReptiles y Anfibios, Tegucigalpa, Honduras

Matthias Goetz, Dipl.-Biol.Durrell Wildlife Conservation Trust, Jersey,British Isles

Alexander Gutsche, Dipl.-Biol.Humboldt Universität, Berlin, Germany

Richard HudsonInternational Iguana Foundation and Fort Worth Zoo, Fort Worth, TX

John B. Iverson, Ph.D.Earlham College, Richmond, IN

Chuck KnappUniversity of Florida, Gainesville, FL

Michael Ripca, Managing Editor of IGUANALeaping Lizards Graphic Design, Atco, NJ

Thomas Wiewandt, Ph.D.Wild Horizons, Tucson, AZ

FounderRobert W. EhrigBig Pine Key, FL

Statement of PurposeThe International Iguana Society, Inc. is a not-for-profit corporation dedicated to preserving the biological diversity ofiguanas. We believe that the best way to protect iguanas and other native plants and animals is to preserve naturalhabitats and to encourage development of sustainable economies compatible with the maintenance of biodiversity.To this end, we will: (1) engage in active conservation, initiating, assisting, and funding conservation efforts incooperation with U.S. and international governmental and private agencies; (2) promote educational efforts relatedto the preservation of biodiversity; (3) build connections between individuals and the academic, zoo, and conserva-tion communities, providing conduits for education and for involving the general public in efforts to preserveendangered species; and (4) encourage the dissemination and exchange of information on the ecology, populationbiology, behavior, captive husbandry, taxonomy, and evolution of iguanas.

Membership InformationIguana, the Journal of The International Iguana Society, is distributed quarterly to members and member organizations.Annual dues:Individual U.S. and Canadian Membership . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . $25.00Individual Foreign Membership . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . $35.00U.S. and Canadian Organizational Membership* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . $35.00Foreign Organizational Membership* . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . $45.00

(*receives double copies of Iguana) Additional copies are available at a cost of $6.00 including postage.

JOIN ON-LINE AT: www.IguanaSociety.org

Membership questions? Call AJ at 860-236-8203, or write to: The International Iguana Society, Inc., 133 SteeleRoad, West Hartford, CT 06119

SolicitationsMembers of the I.I.S. are encouraged to contribute articles, letters to the Editor, news items, and announcements forpublication in Iguana. General articles can deal with any aspect of iguana biology, including conservation, behavior,ecology, physiology, systematics, or husbandry. Submission of photographs to accompany articles is encouraged.

Manuscripts may be submitted via e-mail (send to [email protected]). For any contribution, please include yourname, address, phone number, and e-mail address. Authors of one page or more of print are entitled to five copies ofthe issue in which their article appears.

Advertising Policy of IguanaWe advertise only non-living products (except feeder insects). All products have been examined and been found to behigh quality and fairly priced. Contact Sandy Binns, Advertising Director, at [email protected] or 3010 MagnumDrive, San Jose, CA 95135.

Iguana Copyright ©2003 by The International Iguana Society, Inc., 133 Steele Road, West Hartford, CT 06119 USA. ISSN # 1098-6324. The contents of Iguana, excluding Research Articles, Research Updates, andreprinted articles, may be reproduced for inclusion in the newsletters of other herpetological societies, provided the material is reproduced without change and with appropriate credits, and a copy of the publication issent to The International Iguana Society. Occasional exceptions may be made to this policy.

Editors’ RemarksIGUANA has undergone substantive changes within the past year. The name has changed, the cover went to blackand white, new features have been added, and now the color covers are back and the page count increased — yetthe message remains the same: iguanas of all kinds are fascinating animals, whether we experience them in theirnatural habitat or keep them in our homes — and they deserve our attention. Many wild populations are threatenedor endangered and nearly all are declining. Our emphasis on conservation remains unchanged and we will continueto focus on populations in need of our assistance, bringing news of setbacks and advances, and profiling projectsand persons that endeavor to help these magnificent animals. However, because many of us came to appreciatethese reptiles by caring for them in captivity, we also will strive to provide information that will enhance our ability tocare for them in the most humane and efficient manner.

In this issue, we present some shocking images of iguanas and other reptiles exploited primarily for food in their nativetropical habitats. Although some might argue that this differs little from the exploitation of cattle, swine, or poultry intemperate regions, one major distinction exists: iguanas are wild animals and continued exploitation threatens theirvery existence. As iguana enthusiasts, regardless of whether we relate best to a pet we have named and with whichwe share our home or to the fleeting glance of a wild lizard fleeing through the tropical thorn scrub, we must set highstandards in our support of conservation efforts and in responsible ownership of even a single animal.

This means following the guidelines set out at the end of the article on the exploitation of reptiles in the West Indies(those “rules” apply everywhere), avoiding impulsive purchases of animals (see the letter from the President), being aresponsible pet owner (see, for example, the consequences of escaped iguanas in Newsbriefs), supporting animalrescuers, and, above all, seeking to educate the public and other pet owners by taking every opportunity to inform

them of conservation needs and appropriate husbandry concerns.

The IIS did just that at this year’s National Reptile Breeder’s Expo inDaytona Beach, Florida. Representatives of the Society enjoyedspeaking to many current IIS members and welcomed 32 newmembers! In the course of the weekend, hobbyists, breeders,rescuers, veterinary professionals, and conservationists shared ahigh level of concern for the well-being of an extraordinary groupof animals.

Bob Powell, AJ Gutman, and John Binns

The IIS in Daytona (left to right): Joe Wasilewski, John Binns, Liza Greenberg, Sandy Binns, AJ Gutman, and Joe Burgess. Visit our website (www.IguanaSociety.com) for more photos from Daytona. Photograph by John Binns.


L E T T E R F R O M T H E P R E S I D E N T

Iguanas make great pets — for those who make acommitment before they purchase one. In fact, acommitment should precede the acquisition of anyexotic or domestic pet. Pets are completely dependenton their owners for all of their needs, including food,shelter, and veterinary care. Domestic dogs have a lifeexpectancy of 12–15 years, yet animal shelters take inthousands of dogs daily. Iguanas, on the other hand, if properly maintained, will live into their mid-twenties.

During the mid-1980s, hundreds of thousands of Green Iguanas were imported for the pet industry.Most came from Central and South American iguana“farms,” where gravid female iguanas were releasedinto huge enclosures and their eggs were collected.Initially, iguanas sold for $50 or more. Once themarket was saturated, however, the price of a babyiguana dropped to the point where some pet shopswere offering free iguanas with the purchase of a ten-gallon aquarium. Importers were selling large num-bers of iguanas for as little as 75 cents apiece. Thisonly served to cheapen the quality of care theseanimals received.

Iguanas belong to a group of animals classified as“impulse” pets. People may visit a pet shop, see aniguana at an inexpensive price, and the next thing theyknow — they’re pet owners. A quick-talking salesper-son can outfit a customer with everything needed toraise an iguana. Sadly, the novelty quickly wears off formost people, and the animals suffer from neglect. Fewwill actually receive proper care and live long, healthylives. Contrary to popular belief, zoos and animalattractions cannot take those that survive, and theyend up in the hands of the few wildlife rehabilitatorswho are willing to deal with them. Consequently,many are dumped back on the pet shops or released.Depending on the local climate, the latter may or maynot thrive.

In South Florida, the climate is well-suited toexotic wildlife, and the Green Iguana population hasexploded (see Newsbriefs, p. 94). Healthy populationsof Green Iguanas range from Southern Palm Beachinto the Florida Keys. They have taken up residence inseveral tourist attractions, local parks, and botanicalgardens. Even southern Florida is a bit too far northfor Green Iguanas to occur naturally, because severaldays throughout the year are too cold. The past fiveto seven years in Florida have been unusually warm,causing iguana numbers to grow at an alarming rate.

Despite the warm trend, the temperature dippedto under 40° for several nights this past January, andmany Floridians thought that many iguanas wouldperish. On one cold Saturday morning, I visited alocal tourist attraction with a healthy population of

iguanas. The curator said he had found several dozeniguanas sprawled on the ground and that they hadbeen collected and put into boxes. By the time Iarrived, it was near noon, and there I stood, camera inhand, ready to document dozens of iguana carcasses.To my surprise, the staff walked out with several hugeboxes and began putting them in my truck. Muchscratching came from the boxes, and, upon inspec-tion, I discovered that the iguanas were alive! Coldand slow, but alive. So, when you look at the accom-panying photo, note that the only way to pose with10–15 iguanas is for them to be half-frozen.

Remember, the next time you go to a pet shop,beware of impulse purchases.

Joe Wasilewski

Phot

ogra

ph b

y Li

za G

reen

berg

.

Adult m

ale Spiny-tailed Iguana (Ctenosaura sim

ilis) from U

tila, Bay Islands, Honduras (stories on pp. 63 &

79). Photograph by John Binns.

iguana volume 10, number 3 september 2003 us$6 · diverse cadre of species. the long-term (20+...

Documents