IT 9.1-2 B&W pages · 2017-01-15 · Iguana Times THE JOURNAL OF THE INTERNATIONAL IGUANA SOCIETY g T VOLUME 9, NUMBERS 1 AND 2 MARCH/JUNE 2002 $12.00 A large male spiny-tailed iguana,

Iguana TimesTHE JOURNAL OF THE INTERNATIONAL IGUANA SOCIETYWWW.IGUANASOCIETY.ORG

Iguana TimesVOLUME 9, NUMBERS 1 AND 2

MARCH/JUNE 2002$12.00

A large male spiny-tailediguana, Ctenosauraconspicuosa, basks atop acardon cactus, Pachycereuspringlei, on Isla SanEsteban, a small island inthe Gulf of California. Photograph: L. Lee Grismer

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Ageneral paradigm among many biolo-gists and lay people alike is that “millionsof years” are required for species toevolve. In many cases this is true, but

numerous endemic species on young landbridgeislands from throughout the world are only thou-sands of years old, indicating that the speed of evo-lution is not constant across or within taxonomicboundaries. Population genetics has shown us thatevolutionary rates vary according to populationsize and the selection pressure on that popula-tion. Such phenomena are most efficientlystudied and best exemplified in island popula-tions such as those in the Gulf of California.This 1000-km-long inland sea lies between theBaja California Peninsula and continental mainlandMéxico and harbors a diverse array of islands thatvary in geological origin, age, and size.

The herpetofauna of the Gulf of California hasreceived considerable attention for nearly 50years (e.g., Case 1975, 1983; Cliff 1954a,b;Dixon 1966; Grismer 1994a,b,c,d1999a,b,c; Hews 1990a,b; Hollingsworth1998; Murphy 1975, 1983a,b; Murphyand Ottley 1984; Petren and Case 1997;Radtkey et al. 1997; Robinson 1972,1974; Savage 1960; Soulé 1964, 1966;Soulé and Sloan 1966; Upton andMurphy 1997; Wilcox 1978). Recently,Cryder (1999) presented a phylogenyof the four species of spiny-tailediguanas of the Ctenosaura hemilophagroup (sensu Grismer 1999a) based on

mitochondrial DNA sequences of portions of thecytochrome b and cytochrome oxidase III genes(Fig. 1). These species are endemic to the circum-Gulf of California region of northwestern Méxicoand occur on a number of islands in the Gulf ofCalifornia (Grismer 1999b; Fig. 2). Cryder’s(1999) results supported previous hypotheses con-cerning the human-facilitated origin of Ctenosauraon Isla San Esteban (Grismer 1994b) and indi-

cated that overwater dispersal byctenosaurs to other islands is

still a relativelyrecent (and

perhaps

Spiny-tailed Iguanas, Insular Evolution, and Seri Indians:

How Long Does it Take to Make a NewSpecies and Does it Matter Who Makes it?

L. Lee GrismerDepartment of Biology

La Sierra UniversityRiverside, California 92515-8247

3Iguana TimesMarch/June 2002

Ctenosaura conspicuosa. Photograph: L. Lee Grismer

ongoing) phenomenon. Cryder’s (1999) studyused individual lizards from each of the four rec-ognized species as the unit of analysis. Because hisdata were obtained from the maternally inheritedmitochondrial genome, the resulting branchingdiagram produces a tree depicting the relationshipsbetween individual lizards. These relationships arethen inferred to represent the relationships of thespecies to which those individuals belong.

The relationships between Ctenosaura nola-scensis of Isla San Pedro Nolasco, C. conspicuosaof islas Cholludo and San Esteban, and C. macrolopha of mainland México (Fig. 1) pro-vide the fodder for some intriguing and inextrica-bly related issues concerning the consequences ofinsular isolation, human introduction, and speciesconcepts. All individuals of C. conspicuosa analyzedgrouped together (i.e., they were more closelyrelated to each other than to any other individuals

of other species). Grismer (1994b) hypothesizedthat the population of ctenosaurs on islas SanEsteban and Cholludo were introduced by SeriIndians. This was based on a number of circum-stantial lines of evidence. First, the coastal distri-bution of C. macrolopha (a presumable continentalsource) in mainland Sonora approaches Guaymasat its northern extent (Smith 1972), approximately115 km south of the nearest coastal mainlandlocality adjacent to Isla San Esteban. The absenceof Ctenosaura on the large, geographically inter-mediate island of Tiburón (Fig. 2) suggests that, ifCtenosaura had ever occupied more northerlyareas of coastal Sonora, it would be present on IslaTiburón, as is every other species of reptile thatoccurs on the adjacent Sonoran mainland (Grismer1994c). If the presence of C. conspicuosa on IslaSan Esteban is a natural occurrence, then ahypothesis is necessary to explain its absence from

4 Iguana Times Volume 9, Numbers 1 and 2

Figure 1. Relationships of the Ctenosaura hemilopha group based on cytochrome b and cytochrome oxidase III.

Isla Tiburón and adjacent mainland México.Second, Isla San Esteban is the only island in theGulf of California where the chuckwalla (genusSauromalus) and Ctenosaura are sympatric. Thepotential sympatry of these large, herbivorous gen-era exists on at least 13 other major islands in theGulf of California given the continental and penin-sular distributions of these genera (Fig. 2). Two ofthese islands (San Pedro Nolasco and Cerralvo)have only Ctenosaura, whereas the other 11islands (Espirítu Santo, Partida Sur, San Francisco,San José, San Diego, Santa Cruz, Santa Catalina,Monserrate, Carmen, Danzante, and Coronados)have only Sauromalus (Grismer 1999b). Third,the genetic data of Cryder (1999) were consistentwith the Seri Indian introduction hypothesis inthat C. conspicuosa was most closely related toanother insular endemic, C. nolascensis, rather thana peninsular or continental population, whichwould have more likely been a source of origin hadits continental distribution been farther north. Weknow that the Seri Indians collected and trans-ported both Sauromalus and Ctenosaura as foodsources (e.g., Case 1982, Felger and Moser 1985)and would release them on small islands where

they could easily be recaptured (see Grismer1994b for discussion). Fourth, and most com-pelling, is Gary Nabham’s work on the ethnoher-petology of the Seri Indians (book manuscript inpress). Nabham learned the Seri language andinterviewed Seri elders about their use and viewson the local herpetofauna. They informedNabham that the Seri had always collected iguanasfrom Isla San Pedro Nolasco as a food source andbrought them back to their families living on IslaSan Esteban. This provides the best explanation todate for the presence of Ctenosaura on Isla SanEsteban and the close relationship of C. conspicu-osa and C. nolascensis (Fig. 1) on islands that areseparated from one another by 146 km (Fig. 2)and share no common geological history.

Another paradigm often associated with insu-lar biogeography is one in which island coloniza-tion is considered to be something that has hap-pened rather than something that is happening.Wong et al. (1995) demonstrated recent coloniza-tion of zebra-tailed lizards (Callisaurus dra-conoides) on Isla Danzante in the Gulf ofCalifornia. Cryder (1999) demonstrated that allindividuals of Ctenosaura macrolopha from main-


Ctenosaura hemilopha. Photograph: L. Lee Grismer

land México grouped together, but found thattwo individuals of C. nolascensis from Isla SanPedro Nolasco were more closely related to theindividuals of C. macrolopha than to the otherindividuals of C. nolascensis (Fig. 1). The most par-simonious interpretation of these data is that thesetwo individuals represent recent colonizers fromthe adjacent C. macrolopha gene pool on mainlandSonora to Isla San Pedro Nolasco. Isla San PedroNolasco lies 12 km off the coast of Sonora oppo-site significant drainages that purge large quanti-ties of jetsam into the Gulf of California followingheavy storms. A large arboreal lizard rafting onfloating vegetation for short distances is not incon-ceivable.

The most debatable issue concerningctenosaurs and man is how long does it take for aspecies to evolve? And, if such an event is the resultof a human introduction, does it make it any less anatural event — especially if the human populationwas living as an aboriginal entity of nature. In part,this depends on one’s concept of a species. In ele-vating the subspecies of Ctenosaura hemilopha tofull species, Grismer (1999a) was using a lineage-based concept, emphasizing a population’s inde-


Ctenosaura macrolopha. Photograph: L. Lee Grismer

Figure 2. Distribution of the species of the Ctenosaurahemilopha group.

pendent evolutionary history (as determined by itspossession of unique characteristics) and its poten-tial to continue to evolve separately from all otherpopulations (as a result of its allopatry) as evidencefor species status (see Grismer 1999a for discus-sion). Under this species concept, C. conspicuosa isrecognizable as a valid species. Does it matter thatthe population may be the result of human intro-duction? My opinion is no. The genetic conse-quences of allopatric speciation are going to pro-ceed as a mechanistic process of recombinationoperated on by selection unmindful of the cause ofthat selection process. Of course, one can take thisto levels of absurdity and opine that fruit flies in abottle could be considered valid species under thismodel. The uselessness of this extreme is obvious,but it does touch on more realistic issues such asthose presented above. I do not know where theline should be drawn, but I do believe it lies some-where between the two examples. Birds (a naturalentity of nature) frequently contribute to seed dis-persal on islands and I see no difference betweenthis entity and aboriginal human populations.

The next issue is one of time. Some “experts”are of the opinion that thousands of years may betoo short a time period for a species to evolve.Indeed, I have been criticized for describing asnew some of the most distinctive and bizarrespecies from landbridge islands in the Gulf ofCalifornia. The critique was not one of characteranalysis but one of time, in that landbridge islandsare too young (8–10 thousand years) to havespecies (but they could have subspecies!). Thisimplies that evolution happens at a constant ratefor all taxa. However, evolution will proceed as fastas necessary within the genetic capabilities of thepopulation in order to avoid the extinction of thatpopulation. The potential competition betweentwo large herbivorous lizards on a small desertisland could potentially be severe and serve todrive evolutionary change. Case (1982), however,noted little in the way of competition betweenSauromalus varius and Ctenosaura conspicuosa.The change in C. conspicuosa may simply begenetic drift following a severe bottleneck.

The beauty of Cryder’s (1999) work is that itprovides us with an insight into the evolutionaryhistory of this group of spiny-tailed iguanas that,using a morphological data set, was unobtainablyimbedded in its phylogeny. Combining phylo-

geography with natural history, plate tectonics, andethnoherpetology, we can begin to piece togethera coherent scenario from a vast diversity of sourceswhose corroboration strengthens our interpreta-tions of this group’s history.

Literature CitedCase, T. J. 1975. Species numbers, density compensation, and

colonizing ability of lizards on islands in the Gulf of California.Ecology 56:3–18.

Case, T. J. 1982. Ecology and evolution of the insular giantchuckawallas, Sauromalus hispidus and S. varius, pp.184–212. In G. M. Burghardt and A. S. Rand (eds.), Iguanasof the World. Their Behavior, Ecology, and Conservation.Noyes Publications, Park Ridge, New Jersey.

Case, T. J. 1983. The Reptiles: Ecology, pp. 159–209. In T. J. Caseand M. L. Cody (eds.), Island Biogeography in the Sea ofCortéz. University of California Press, Berkeley, California.


Ctenosaura nolascensis. Photograph: L. Lee Grismer

Cliff, F. S. 1954a. Snakes of the islands in the Gulf of California,Mexico. Transactions of the San Diego Society of NaturalHistory 12:67–98.

Cliff, F. S. 1954b. Variation and Evolution of the Reptiles Inhabitingthe Islands in the Gulf of California, Mexico. Ph.D.Dissertation, Stanford University, Stanford, California.

Cryder, M. R. 1999. Molecular Systematics and Evolution of theCtenosaura hemilopha Complex (Squamata: Iguanidae). M.S.Thesis, Loma Linda University, Loma Linda, California.

Dickerson, M. C. 1919. Diagnosis of twenty-three new speciesand a new genus of lizards from Lower California. Bulletin ofthe American Museum of Natural History 61:461–477.

Dixon, J. R. 1966. Speciation and systematics of the gekkonidlizard genus Phyllodactylus of the islands in the Gulf ofCalifornia. Proceedings of the California Academy ofSciences, 4th series 33:415–452.

Felger, R. S. and M. B. Moser. 1985. People of the Desert andSea. Ethnobotany of the Seri Indians. The University ofArizona Press, Tucson, Arizona.

Grismer, L. L. 1994a. The origin and evolution of the peninsularherpetofauna of Baja California, México. HerpetologicalNatural History 2:51–106.

Grismer, L. L. 1994b. Geographic origins for the reptiles on islandsin the Gulf of California, México. Herpetological NaturalHistory 2:17–40.

Grismer, L. L. 1994c. The Evolutionary and EcologicalBiogeography of the Herpetofauna of Baja California and theSea of Cortés, México. Ph.D. Dissertation, Loma LindaUniversity, Loma Linda, California.

Grismer, L. L. 1994d. Three new species of intertidal side-blotchedlizards (Genus Uta) from the Gulf of California, México.Herpetologica 50:451–74.

Grismer, L. L. 1999a. An evolutionary classification of reptiles onislands in the Gulf of California, México. Herpetologica55:446–69.

Grismer, L. L. 1999b. Checklist of amphibians and reptiles onislands in the Gulf of California, México. Bulletin of theSouthern California Academy of Sciences 98:45–56

Grismer, L. L. 1999c. Phylogeny, taxonomy, and biogeography ofCnemidophorus hyperythrus and C. ceralbensis (Squamata:Teiidae) in Baja California, México. Herpetologica 55:28–42.

Hews, D. K. 1990a. Resource Defense, Sexual Selection, andSexual Dimorphism in the Lizard Uta palmeri. Ph.D.Dissertation, The University of Texas, Austin, Texas.

Hews, D. K. 1990b. Examining hypotheses generated by fieldmeasures of sexual selection on male lizards, Uta palmeri.Evolution 44:1956–66.

Hollingsworth, B. D. 1998. The systematics of chuckwallas(Sauromalus) with a phylogenetic analysis of other iguanidlizards. Herpetological Monographs 12:38–191.

Murphy, R. W. 1975. Two new blind snakes (Serpentes:Leptotyphlopidae) from Baja California, Mexico with acontribution to the biogeography of peninsular and insularherpetofauna. Proceedings of the California Academy ofSciences, 4th series 40:87–92.

Murphy, R. W. 1983a. Paleobiogeography and genetic differentia-tion of the Baja California herpetofauna. Occasional Papers ofthe California Academy of Sciences 137:1–48.

Murphy, R. W. 1983b. The reptiles: origin and evolution, pp.130–158. In T. J. Case and M. L. Cody (eds.), IslandBiogeography in the Sea of Cortéz, University of CaliforniaPress, Berkeley, California.

Murphy, R. W. and J. R. Ottley. 1984. Distribution of amphibiansand reptiles on islands in the Gulf of California. Annals of theCarnegie Museum 53:207–230.

Petren, K. and T. J. Case. 1997. A phylogenetic analysis of bodysize evolution and biogeography in chuckwallas (Sauromalus)and other iguanines. Evolution 51:206–219.

Radtkey, R. R., S. M. Fallon, and T. J. Case. 1997. Characterdisplacement in some Cnemidophorus lizards revisited: Aphylogenetic analysis. Proceedings of the National Academyof Sciences 94:9740–9745.

Robinson, M. D. 1972. Chromosomes, Protein Polymorphism, andSystematics of Insular Chuckwalla Lizards (Genus Sauromalus)in the Gulf of California, Mexico. Ph.D. Dissertation,University of Arizona, Tucson, Arizona.

Robinson, M. D. 1974. Chromosomes of the insular species ofchuckwalla lizards (genus Sauromalus) in the Gulf ofCalifornia, Mexico. Herpetologica 30:162–167.

Savage, J. M. 1960. Evolution of a peninsular herpetofauna.Systematic Zoology 9:184–212.

Smith, H. M. 1972. The Sonoran subspecies of the lizardCtenosaura hemilopha. Great Basin Naturalist 32:104–111.

Soulé, M. E. 1964. Evolution and Population Phenetics of theSide-blotched Lizards (Uta stansburiana and its relatives) onthe Islands in the Gulf of California, Mexico. Ph.D.Dissertation, Stanford University, Stanford, California.

Soulé, M. E. 1966. Trends in an insular radiation of a lizard.American Midland Naturalist 100:47–64.

Soulé, M. E. and A. J. Sloan. 1966. Biogeography and distribu-tion of the reptiles and amphibians on islands in the Gulf ofCalifornia, Mexico. Transactions of the San Diego Society ofNatural History 14:137–156.

Upton, D. E. and R. W. Murphy. 1997. Phylogeny of the side-blotched lizards (Phrynosomatidae: Uta) based on mtDNAsequences: support for a midpeninsular seaway in BajaCalifornia. Molecular Phylogenetics and Evolution 8:104–113.

Wong, H., E. Mellink, and B. D. Hollingsworth. 1995. Proposedrecent overwater dispersal by Callisaurus draconoides to IslaDanzante, Gulf of California, México. Herpetological NaturalHistory 3:179–182.

Wilcox, B. A. 1978. Supersaturated island faunas: a species-agerelationship for lizard faunas on post-Pleistocene land-bridgeislands. Science 199:996–998.



In a recent study of the phylogeography ofCyclura, Malone et al. (2000) presented aphylogram (Figure 1) that resulted from ananalysis of mitochondrial DNA sequence

data. In concurrence with other recent molecularstudies (Sites et al., 1996; Norell and de Quieroz,1991; Petren and Case, 1997), these data stronglysupport the monophyly of the Iguanidae (sensustrictu) and the antiquity of Brachylophus.Dipsosaurus resolved as the sister group to a cladecomposed of Cyclura, Ctenosaura, Conolophus,Amblyrhynchus, Sauromalus, and Iguana. Cycluraalso is supported as a monophyletic lineage,equally related to Ctenosaura, Conolophus,Amblyrhynchus, Sauromalus, and Iguana. Withinthe latter assemblage, the data strongly supportthat Iguana andSauromalus are sistertaxa, contrary to theconclusions of Wiensand Hollingsworth(2000, and referencestherein), who sug-gested that Iguanaand Cyclura were sorelated. Within Iguana,lineages from differentgeographical areaswere highly differenti-ated (not shown here),and may result in taxonomic distinctionswith further study.

As expected,endemism wasextremely high withinCyclura, with each lin-eage restricted to oneisland or island group.Hispaniola is the only

island that supports two distinct lineages, possiblyreflecting separate origins on the two paleoislandsthat joined to form the current island. The distinctlineages cluster geographically (Figure 2). An“eastern” clade is composed of C. pinguis, the old-est extant lineage, which historically inhabitedmost of the Puerto Rico Bank. A “central” clade iscomposed of C. cornuta from Hispaniola and IslaMona and its sister group, containing C. ricordiifrom Hispaniola and C. carinata from the Turksand Caicos Islands. A “western” clade is composedof C. collei from Jamaica, C. rileyi from the west-ern Bahamas, C. cychlura from the easternBahamas, and C. nubila, as traditionally defined,from Cuba and the Cayman Islands. This contra-dicts the topology of Schwartz and Carey (1977),

who considered the C.ricordii + C. carinatagroup basal and clus-tered C. cornuta withC. rileyi, C. collei, C. nubila, and C. cychlura. This data setalso brought to lightthe need for furtherinquiry into the relationship betweenCyclura nubila lewisifrom Grand Caymanand populationspresently consideredto be conspecific, C. n.nubila from Cuba andC. n. caymanensis fromLittle Cayman andCayman Brac. Alsonoteworthy are thevery close associationsbetween populationscurrently recognized as

Comments on a Phylogeny ofIguanid Lizards

Catherine L. Malone1 and Robert Powell2

1Department of Forestry and Natural Resources, Purdue University, West Lafayette, Indiana 479072Department of Biology, Avila College, Kansas City, Missouri 64145

Figure 1. The maximum-likelihood estimate of relation-ships among iguanid lizards represented as a phylogram.The asterisk (*) indicates that support for this node is weakand that, pending further data, the relationship betweenCtenosaura, Conolophus, and Amblyrhynchus might beconsidered a trichotomy.


Figure 2. A phylogeographic distribution of the genus Cyclura, with the maximum likelihood estimate of historical relation-ships within Cyclura superimposed onto a map of the Greater Antilles, Bahamas, and Turks and Caicos Islands. The dottedarrow represents the probable historic range of C. pinguis on Puerto Rico proper. Photographs by John Binns (Cyclura nubila,C. collei, C. carinata, C. ricordii), Carl Fuhri (C. cychlura, C. rileyi), Robert Powell (C. nubila lewisi, C. cornuta, C. c. stejnegeri),and Glenn Gerber (C. pinguis).

Cyclura nubila lewisi Cyclura nubila caymanensis Cyclura cychlura Cyclura rileyi

Cyclura collei Cyclura carinata Cyclura ricordii Cyclura cornuta

Cyclura cornuta stejnegeri

Cyclura pinguis

Phylogeography of Cyclura


subspecies of C. cychlura, C. rileyi, and C. cornuta;with these data arguing strongly against species-level recognition of populations of the latter fromHispaniola and Isla Mona (e.g., Powell and Glor,2000).

Forthcoming work involves a closer look atthe relationships within the Iguana iguana com-plex, within Ctenosaura (C.R. Hasbun, pers.comm.), and between C. cornuta from Hispaniolaand Isla Mona. Until these data are adequatelyanalyzed, we suggest that the phylogram presentedherein constitutes the best currently available rep-resentation of relationships among iguanid lizards.

Literature CitedMalone, C.L., T. Wheeler, J.F. Taylor, and S.K. Davis. 2000.

Phylogeography of the Caribbean Rock Iguana (Cyclura):Implications for conservation and insights on the biogeo-graphic history of the West Indies. Molecular Phylogeneticsand Evolution 17:269–279.

Norell, M.A., and K. de Quieroz. 1991. The earliest iguaninelizard and its bearing on iguanine phylogeny. AmericanMuseum Novitates (2997):1–16.

Powell, R. and R.E. Glor. 2000. Cyclura stejnegeri. Catalogue ofAmerican Amphibians and Reptiles (711):1–4.

Schwartz, A. and M. Carey. 1977. Systematics and evolution inthe West Indian iguanid lizard genus Cyclura. Studies on theFauna of Curaçao and Other Caribbean Islands 53:15–97.

Sites, J.W., S.K. Davis, T. Guerra, J.B. Iverson, and H.L. Snell.1996. Character congruence and phylogenetic signals inmolecular and morphological data sets: a case study in theliving iguanas (Squamata, Iguanidae). Molecular Biology andEvolution 13:1087–1105.

Wiens, J.J. and B.D. Hollingsworth. 2000. War of the iguanas:conflicting molecular and morphological phylogenies andlong branch attraction in iguanid lizards. Systematic Zoology49:143–159.

Cyclura nubila nubila (above), C. nubila caymanensis, and populations in the C. cychlura complex appear to be more closelyrelated to one another than to C. nubila lewisi, which probably is a distinct species. Photograph by John Binns.


0730 h, temp 72°F, wind NE at 15 knots,seas 2–3 feet

A group of Earlham College students yawn,rub their eyes, and awaken to the aroma of break-fast aboard the Bahamas Star. The boat has com-pleted the five-hour journey to a set of remoteislands located in the northern portion of theExuma Island chain of the Bahamas. Here stu-dents will learn the essentials of herpetologicalfieldwork while working with the endemic iguanas.

Characteristic of lizards in general, the iguanasdo not start their day until the sun’s rays warm thesand on this group of three islands, known as theAllen’s Cays. Consequently, the students’ day willfollow the same pattern, initially by waiting for thefirst iguana to make its way onto the beach to

bask. A leisurely breakfast is followed by the appli-cation of sunscreen and the packing of personalgear: sunglasses, more sunscreen, a towel, durableshoes, snorkeling equipment, Joy dishsoap (for anocean-friendly scrub), Neosporin for scratches (onpeople and iguanas), photocopied maps of theisland, and a Sharpie. Also loaded into the trans-port dinghy are rulers, scales, PIT-tagging equip-ment, large scissors, nooses, nets, pillowcases, andbait. These are the tools that the crew, headed byDr. John. B. Iverson, will use to capture and exam-ine Cyclura cychlura inornata.

Dr. Iverson began this study in 1980, a year inwhich some of the students who are currently par-ticipating in the adventure were not yet alive, andthe rest were too young to have relevant recollec-

Figure 1. Cyclura cychlura inornata on the beach at Allen’s Cays (photographs by Lynne Pieper, May 2001).

Sun, Students, and Scratches:Research on Allen’s Cays Iguanas

Samantha C. Larimer1,2 and Stesha A. Pasachnik1

1Department of Biology, Earlham College, Richmond, Indiana 473742Current address: Department of Psychology, Cornell University, Ithaca, New York 14850


tions. In this ongoing natural history study of theendangered iguanas (C.I.T.E.S, Appendix 2),Earlham College students annually get an oppor-tunity to be a part of this important researchendeavor, learning about the iguanas whose storythey’re helping to document. Issues regarding thisspecies’ habitat, learning about other nativespecies, the “challenge” of living on a boat in theCaribbean, and tourism are all parts of this expe-dition to the Exumas.

Cyclura cychlura inornata is a large iguana,reaching more than 120 cm in total length andoften living for more than 30 years. Breeding pop-ulations, until recently, have occurred only onAllen’s Cays. Here, they are found predominantlyon Leaf Cay (4 ha) and U Cay (3 ha), although 11individuals have been recorded from Allen’s Cay(7 ha) (Iverson, 2000). The latter are all at least 10cm larger than the largest individuals on the otherislands. This is not only an amazing sight tobehold but also a phenomenon yet to be

explained, although one may speculate that a dif-ference in diet and/or reduced intraspecific com-petition might play a role. Between 1988 and1990, eight individuals were transferred toAlligator Cay (1.8 ha), located in the Exuma CaysLand and Sea Park, Bahamas. This was done as asafeguard against the Allen’s Cays populationbeing destroyed by a stochastic event such as ahurricane, an epidemic, or the introduction of aferal mammalian predator. Since their introduc-tion, these individuals have been very prolific andwere, at last count in 1998, close to carryingcapacity (Knapp, 2001).

For 21 years, Dr. Iverson has been takinggroups of students to the Exumas from the smallliberal arts college in Richmond, Indiana, where heteaches biology. Collectively, these expeditionshave been able to gather data essential to under-standing the life history and survival of this species.Eventually he would like to be able to documentlife expectancies, age classification, mating sched-ules, reproductive parameters, causes of death, andthe means of their prevention. Basically, he wantsto know just about everything one can learn witha minimum amount of intrusion. As part of thislong-term data-collection project, temperaturerecorders (tidbits) are placed on the island in vari-ous habitats to monitor year-round environmentalconditions. Dr. Iverson recently (June 2001)launched a more intensive study of the iguanas’reproductive behavior and capacity by spendingthe month-long mating season on the islands.

0830 h, and temps have risenIguanas become visible as dark masses pepper-

ing the sandy shore. The crew is ready for the day,and 11 former or current students and Dr. Iversonhead to land to begin their work. Some memberswait for a ride in the dinghy; others, eager tobegin, load their bags into the boat and plungeinto the water to swim the 150 meters to shore.Fifteen minutes later, catching begins in earnest.Students work quickly, walking the beach singly orin pairs, and looking especially at sites whereiguanas aggregate while waiting for tourist hand-outs. Many of these iguanas have become accus-tomed to the tourists who travel in boatloads tosee them, leaving their vessels for 10 minutes inorder to throw some food scraps to these fascinat-ing creatures before taking off for another island.

Figure 2. John Iverson measures an iguana; size is just onebit of data he and the students gather for each individual(photograph by Lynne Pieper, May 2001).


This feeding obviously is not a natural event, andits effects are something that Dr. Iverson hopes tosomeday understand. The massing behavior trig-gered by these handouts appears to have elimi-nated much of the territorial behavior that prob-ably characterized the iguanas’ ancestors. Also, thefood, though easy to obtain, is not kind to theseanimals’ digestive systems, often passing throughtheir guts less completely digested than their nor-mal diet of leaves and fruit (J.B. Iverson, unpubl.data). Whether this altered diet has an effect onthe life span of this species is something thatrecently collected data comparing iguanas fromdifferent sites will hopefully be able to determine.Because limited areas of each island receive visitsfrom tourists, detailed locality data will allow com-parisons of growth rates between individualsreceiving altered diets and those that are notaffected by tourism.

In the short term, tourists are both helpful anda hindrance to the researchers. The familiarity ofthe iguanas with humans makes them initially easyto capture. Many can be approached easily, and thefriendliest individuals can be captured with a sweepof the net. Others are a bit more cautious, andneed to be lured with bait. Small rocks or shellsfound on the beach are sometimes enough, thefalling objects possibly mimicking ripe fruit fallingfrom a tree. For the most wary individuals, leftoverfood from the boat works well. Orange peels withtheir strong citrus smell and bright orange col-oration are favorites; however, grapes, apples,bread, and lettuce all work well. More renewablethan the food are “swing baits,” which wereinvented by Dr. Iverson and his crew a decade ago.Large red Legos® tied to string can be tossedrepeatedly to fall with just the right force to inter-est an iguana, and the red color is also thought to

Figure 3. Too heavy for the scales, handling this huge iguana, caught after hours of stalking by boat Captain Bruce onAllen’s Cay, required two people, in this case John Iverson and Geoff Smith. For unknown reasons, iguanas on Allen’s Caytend to grow much larger than those on Leaf or U Cays (photograph by Lynne Pieper, May 2001).


excite them. An added advantage of the swingbaits is that they can be pulled in slowly, luringreluctant individuals from the bushes to a net lyingon the ground, which is flipped over the distractedlizard. The struggling iguana is then extractedfrom the net. This is often a two-person task, espe-cially for larger individuals, and painful scratcheshave been known to result from the effort.Eventually, the lizard is placed into a pillowcase,which is tied and placed in the shade until pro-cessing. Variations on the catching theme arenumerous, with each student developing his or herown technique over the course of the week. Largenooses of heavy cord attached via fishing swivels toextendible golf ball retrieving rods are sometimesmore successful on particularly cautious animals.Their curiosity at the swinging cord often causesthem to sit patiently while the noose is adjusted.Occasionally the animals will virtually noose them-selves as they attempt to bite the cord.

Regardless of capture technique, the baggedCyclura are taken to a workstation where they areprocessed. Each lizard is weighed, sexed, measured(both tail and snout-vent lengths are recorded),marked, and released. Marking is done in a varietyof ways. All captured iguanas are toe-clipped and

many, depending on size, are PIT-tagged. PassiveIntegral Transponders are tiny glass beads withwire coils inside. They are injected subcutaneously,allowing the animal to be identified with thesweeping of a reader which shows an identificationcode, much like the system used by supermarketsto ring-up prices on groceries. Together, these twomethods of marking provide very reliable identifi-cation (Iverson, 2000), even allowing dead anddecaying animals to be recognized. Prior torelease, the iguanas are marked temporarily withnon-toxic paint. This identifies them as previouscaptures and eliminates wasted time chasing ani-mals for which data have already been recorded.

The response of tourists to this fieldwork variesfrom wanting to be a part of the effort to extremeanger. A few folks become quite offended by themarking of individuals, possibly because it destroysotherwise perfectly natural looking pictures.However, because they too want to help preservethese magnificent animals, others comment on theresearchers’ intrusive methods, and cite signs onthe islands that threaten dire consequences tothose who interfere with these vulnerable crea-tures. Dr. Iverson, in fact, was instrumental inputting up the very signs that the tourists quote,

Figure 4. Students Colin Meeks and Stephen Jones “touch, catch, chase, and harass” a young iguana, ignoring the sign inthe background that prohibits such activities — but it’s all in the name of science (photograph by John Iverson, March 2000).


and the research he is undertaking has beenapproved by all of the appropriate permit-grantingagencies. Still, we were pleased that touristsacknowledge the protection afforded these animalsand are willing to take action.

Although many people encountered in thefield are doing what they can to help protect theseanimals, many others are still causing harm, some-times unknowingly. Humans are the only signifi-cant threat to this species, whether it is in the formof feeding by tourists, poaching, or illegal exploita-tion for international trade. Each action has itsown serious effects. This species is protected undernational Bahamian Law, but enforcement is lack-ing because no warden works in the area and theseislands are extremely secluded (Iverson, 1999). As

for the future, conservation plans include addi-tional studies of the introduced individuals onAlligator Cay and possible captive-breeding pro-grams at the Ardastra Zoo in Nassau and theNature Centre Different on Abaco (Iverson,1999). These are hopeful prospects, but actionstaken by individuals are even more important forthe preservation of this species. Taking time toreport illegal treatment of this species and educat-ing oneself and others are methods by whicheveryone can help save the endangered Allen’sCays Rock Iguanas.

1800 h, temperatures are droppingThe crew piles into the dinghy to return to the

boat for dinner. Many have spent the entire day onthe island and this will be the first time since break-fast that they will be back onboard. After dinner,

Figure 5. After more than an hour of trying to coax thisiguana out of the brush using every type of bait available,student Chaundra Schofield was on the verge of giving upwhen the previously reluctant beast came running down toeat a grape out of the hand of a newly arrived Italiantourist. A quick sweep of the net and two people to helpdetangle it, and the year’s largest iguana from Leaf Cay wasin hand (photograph by Samantha Larimer, May 2001).

Figure 6. Students James Rose and Brendan Gallaghermark an iguana with car touch-up paint to avoid recaptur-ing the same individual later that trip. The paint is eventu-ally shed with the skin and causes no permanent damage(photograph by Lynne Pieper, May 2001).


students gather on deck to listen to Captain Ron,John, and alumni tell stories of past trips, laughingand gasping appropriately. Although most studentsparticipate in only one week of this long-termstudy, these stories allow them to grasp all thatgoes on in a research project of this intensity andduration. This undoubtedly is a new concept forsome, as each party is composed of a wide age-range of individuals with varied interests and back-grounds in science. For some, this will be theironly exposure to science-in-action (as opposed tomerely learning about it from books or television).

Besides the interesting stories, students loung-ing on deck also entertain themselves with goodbooks, decks of cards, or by recording the day’sexperiences in their personal journals, jotting downrecollections and impressions of the day’s excitingevents. The iguana-catching process itself isthrilling. It gives students an opportunity to relive

their childhood fantasies of exploration and adven-ture, their survival dependent upon outsmartingthe wild animals they stalk. However, students domore than work, taking occasional short breaks toexplore tide pools, go snorkeling, or try their luckat retrieving coconuts from trees. Finally, withheads filled with images of the day’s experiencesand eagerly anticipating tomorrow, the studentsmake their way to the bunks to rest in preparationfor the next day’s exhilarating activities, dreamingabout that one iguana that just couldn’t be caught,new capture techniques, a section of the island yetto explored, whether or not enough iguanas canbe captured to shatter last year’s record, …

Note: Stesha Pasachnik and Samantha Larimer wereparticipants on trips in March 2000 and May 2001,respectively. They thank Bob Powell and John Iversonfor comments on this paper, and the latter for mak-ing such trips available to students.

ReferencesIverson, J.B. 1999. Allen’s Cays Iguana, Cyclura cychlura inornata,

p. 34–36. In A. Alberts (comp. & ed.), West Indian Iguanas:Status Survey and Conservation Action Plan. IUCN, Gland,Switzerland and Cambridge, UK.

Iverson, J.B. 2000. Results of Allen’s Cays Iguana study (18–25 March 2000) http://www.earlham.edu/%7Ebiol/iverson/Iguana.html.

Knapp, C.R. 2001. Status of a translocated Cyclura iguana colonyin the Bahamas. J. Herpetol. 35:239–248.

Figure 7. An iguana, excited by a rock that fell onto thesand, comes running to investigate, only to fall prey to anet lying on the ground near the well-placed stone(photograph by Samantha Larimer, May 2001).


I. Binomial Nomenclature.

The classification of animals began in a ratherhaphazard manner. In the early days (prior toLinnaeus), most creatures were given a descriptivename. Unfortunately, such names varied accordingto the language used and even by the circum-stances that prevailed when the organism was “dis-covered” — often independently by several differ-ent people. Consequently, the names that appliedto the same type of animal might equal the num-ber of persons who had encountered it. Some feelfor the confusion that prevailed is evident whenpeople from different parts of the world try tocommunicate while relying exclusively on com-mon names. For example, Europeans speak of an“elk,” which North Americans call a “moose.”The animal called an “elk” in America is referredto as a “wapiti” by Europeans — and the “Irishelk” is a different beast altogether. All “robins” arebirds, but English “robins” are quite different thanthose found in the Western Hemisphere (whichare called “wandering thrushes” in Europe). AGerman herpetologist might speak of a“Wüstenleguan,” to the consternation of anEnglish speaker, who would refer to the same ani-mal as a “desert iguana” (Dipsosaurus dorsalis).

Such an unwieldy system soon became over-burdened when Europeans, during the “Age ofDiscovery,” began to carry back from their voy-ages samples of wildlife from all over the world. Aremarkably simple and flexible alternative wasdeveloped and popularized by Carl Linné, aSwedish botanist (Linnaeus was the Latinized ver-sion of his name under which he published hismany volumes on the plants and animals of theworld). In his system, organisms are ordered in ahierarchical manner, based largely on the numberof common traits possessed by various forms.Thus, animals sharing but a few fundamental sim-ilarities were placed in the same higher categorybut were segregated at the next lower rank. Onlythose individuals that were essentially alike shared

the full gamut of categories and were placed in thesame “species.” Linnaeus’ system was easilyadapted to the vast quantities of new knowledgemerely by adding additional categorical ranks.

Fundamental to the Linnaean system is the“binomen” (= two names), which is used to des-ignate any species (this is the reason we refer to a“binomial system of nomenclature”). The first partof the name is that of the genus to which the ani-mal belongs. Genera (the plural form), accordingto Linnaeus, were groups of species which sharedmore characteristics with each other than withspecies placed in any other genus. The genericname is a proper noun and should always be capi-talized (as should all names of higher categories).As nouns, generic names have gender, whichaccounts for many of the different endings (e.g.,many “feminine” nouns end in –a). The secondhalf of the name is the trivial or specific name.Used to distinguish species within a genus, these

Understanding Animal ClassificationRobert Powell

Department of Biology, Avila College, Kansas City, MO 64145(parts I, III, and IV were adapted from an essay by E.O. Wiley. 2001. Kansas Biology Teacher 10:14–17)

Subadult Iguana delicatissima from Anguilla. The criticallyendangered Anguillian population represents the northern-most extent of the species’ range. The meaning of thisspecies’ trivial name is self-evident. Interestingly, althoughconsidered a delicacy throughout most of its range,Anguillian animals have not been exploited for food by localinhabitants. The precarious state of the population is duealmost entirely to habitat alteration and competition withferal livestock. Photograph: Stesha Pasachink


names are adjectives or possessives that modify thegeneric name. Trivial names must agree in genderwith the generic name and should never be capi-talized. The entire binomen is always set off insome distinct way, usually italicized (in print) orunderlined.

An additional feature of the Linnaean systemis that all names are Latin (or Latinized; manyactually are derived from the ancient Greek or evenfrom various modern languages). Because Latin isno longer the vernacular of any culture, it is anextremely stable language and the grammaticalrules are fixed. Consequently, a Pole or anArgentinean could name a species, and the otherwould immediately recognize it as a scientific nameand maybe even understand its meaning — eventhough they might otherwise be totally incapableof communicating.

Applications of this system give us names likeDipsosaurus dorsalis (the “desert iguana”).“Sauros,” the root of the generic name means“lizard” in Greek. “Dipsos,” also from the Greek,means “thirsty,” an obvious reference to the deserthabitats in which these lizards are found. The triv-ial name “dorsalis” is an adjectival form of “dor-sum,” a Latin word that means “back,” an allusionto the crest of enlarged scales that runs down themiddle of the back in these animals. “Iguana” pre-sumably was modified from an Amerindian wordmeaning a kind of large lizard; “delicatissima” isfrom the Latin, meaning “very delicious.”Descriptions of new species should include an ety-mology, which provides the origin and meaning ofthe new name. Unfortunately, at the time whenmany common species were first described, the‘rules’ were not always observed; as a result, we’reoften forced to guess the meaning of a name if itsderivation is not obvious.

Although daunting when first encountered,mainly because the Latin (or Latinized) namesappear to defy pronunciation, the system makes alot of sense. Compare the binomial to a person’sname. The surname (at least in English-speakingcultures) is equivalent to the generic name (thefamily to which a person belongs is analogous tothe genus to which a species is assigned), and aperson’s first name equates to the specific name(identifying that individual much like the trivialname identifies a unique species).

II. Rules of Priority.

In Part I, I mentioned the confusion thatcould arise when different people at different timesgave different names to the same animal.Unfortunately, this problem is not unique to com-mon names. On more than one occasion, severalproperly constructed scientific names have beengiven to the same species. This may have occurredbecause someone was unaware of an earlierdescription, possibly published in an obscure jour-nal, maybe even in another language. Other rea-sons have included descriptions as different speciesof males and females of the same sexually dimor-phic form, or descriptions as different species ofdifferent stages in a life cycle (tadpole versus meta-morphosed amphibian or a juvenile with a dis-tinctly different appearance than adults of eithersex), or descriptions as different species of individ-uals from different regions or merely individualsdemonstrating the considerable variation known tooccur in some animals. Regardless of the reason,the problem of multiple names led to the necessityfor synonymies, lists of all of the names that havebeen applied to a particular taxon. Usually, the firstname is given precedence over all subsequentnames, but exceptions sometimes occur. The“rules” that govern animal classification areenforced by an International Commission onZoological Nomenclature, and the Commissionmust approve any exceptions. Better than a pro-longed explanation is a relevant example takenfrom a recent issue of the Bulletin of ZoologicalNomenclature (2001. 58:37–40), the formal pub-lication of the Commission (reproduced below inits entirety, except for the list of references and theaddresses of the authors):

Case 3143

Euphryne obesus Baird, 1858 (Reptilia,Squamata): proposed precedence of the specificname over that of Sauromalus ater Duméril,1856

Richard R. Montanucci, Hobart M. Smith, KraigAdler, David L. Auth, Ralph W. Axtell, Ted J.Case, David Chiszar, Joseph T. Collins, RogerConant, Robert Murphy, Kenneth Petren, RobertC. Stebbins


Abstract. The purpose of this application is toconserve the long used and well known specificname of Sauromalus obesus (Baird, 1858) for thechuckwalla (family IGUANIDAE) from the southwestof North America by giving it precedence over thelittle used name S. ater Duméril, 1856.

Keywords. Nomenclature; taxonomy; Reptilia;Squamata; IGUANIDAE; Sauromalus ater;Sauromalus obesus; chuckwallas; southwesternNorth America.

1. In 1856 Duméril (p. 536, pl. 23, figs. 3 and3a) described a new genus and single newspecies of iguanid lizard as Sauromalus ater onthe basis of a single specimen presented byLieutenant M. Jaurès to the Muséum Nationald’Histoire Naturelle, Paris. The holotype(MHNP 813), which lacks locality data, wascollected somewhere in western Mexico dur-ing a world circumnavigating voyage of theFrench frigate La Danaïde.

2. The absence of a type locality for Sauromalusater has remained an acknowledged problemfor systematists working with Sauromalus (seeSchmidt, 1922; Shaw, 1945; Hollingsworth,1998). Shaw (1945, p. 273), unable to studythe holotype due to political conditions inEurope, drew upon descriptive information inDuméril & Bocourt (1870) and Mocquard(1899), and concluded that the holotype must

have originated from one of the islands off thesouthern coast of the Baja California penin-sula. Hence, in referring to the type locality,Shaw (1945, p. 284) stated: ‘Not definitelyknown but undoubtedly one of the severalislands in the southern part of the Gulf ofCalifornia where this species is known tooccur’, Subsequently and without justification,Smith & Taylor (1950) further restricted thetype locality to Isla Espiritu Santo.

3. Two years after Duméril, Baird (1858, p. 253)described the new genus and single newspecies Euphryne obesus and noted that it was‘abundant in the canons of the Colorado, ofCalifornia, collected by Maj. Thomas,Mex[ico] Boundary Survey, and Lt. Ives’Expedition’. The type specimen was given asUSNM 4172 in the U.S. National Museum,Washington. Subsequently, Baird (1859, p. 6,pl. 27) indicated the locality of USNM 4172as ‘Fort Yuma’. Van Denburgh (1922) andShaw (1945) correctly noted the location ofFort Yuma in California. Montanucci (2001)discussed the confusion caused by Baird’spiecemeal publication of data and clarified theparticulars relating to the collector and typelocality. Cope (1864) commented that thename Euphryne Baird, 1858 was a synonym ofSauromalus Duméril, 1856, but both genericnames continued to be used in the literatureuntil Cope (1875) and Coues (1875) placed

Subadult Cyclura nubila from the U.S. Naval Base at Guantanamo Bay, Cuba. For many years, this species was thought to con-tain three subspecies; however, new evidence indicates that the Grand Cayman population is distinct at the species level.Photograph: Robert Powell


Euphryne as a synonym of Sauromalus (seeHollingsworth, 1998, p. 40). Sauromalus hasbeen used since that time.

4. Prior to 1922, the name Sauromalus ater, andnot S. obesus, was used in most papers, includ-ing checklists and distributional accounts.Most notable among these publications areCope (1875, 1900), Stejneger’s (1891)description of a new species of Sauromalus, thechecklists of Yarrow (1882) and Stejneger &Barbour (1917), and Van Denburgh’s (1922)The Reptiles of Western North America. Therecognition of S. ater and S. obesus as separatespecies came with publication of Schmidt’s(1922, pp. 640–641) study of the amphibiansand reptiles of lower California, and was fol-lowed by the later checklists of Stejneger &Barbour (1923, 1933, 1939, 1943). The tax-onomic treatment of the genus Sauromalus byShaw (1945) reinforced the concept that S.ater and S. obesus are separate species, a viewheld by virtually all subsequent workers exceptHollingsworth (1998).

5. In his recent monographic revision ofSauromalus, Hollingsworth (1998) placedSauromalus obesus in the synonymy of S. ater,and restricted the type locality of S. ater tosouthern Sonora. However, Montanucci(2000) argued that Hollingsworth’s analysis todetermine the provenance of the type speci-men was unconvincing due to limitations inhis statistical data, leading to ambiguousresults and an unsubstantiated conclusion.Accordingly, Montanucci (2000) concludedthat, in the absence of any new, compellinginformation, the type locality of S. aterremained open to speculation and conjecture.

6. The literature using the name Sauromalus obe-sus is substantially more abundant and signifi-cant than that using the name S. ater. Beaman,Hollingsworth, Lawler & Lowe (1997) listed626 titles of technical and popular articles per-taining to the genus Sauromalus. Out of thistotal, the name S. ater is used in about 46papers; most of these (34) were publishedbefore 1950, and nearly all pertain to taxon-omy and/or distribution. The literature for S.obesus is profoundly more extensive by com-parison, being conservatively estimated to beabout 90% of the total literature for the genus

as a whole, or some 550 papers. The name S.obesus is used, almost to the exclusion of S.ater, in the literature dealing with physiologi-cal ecology and thermoregulation of chuck-wallas (about 133 papers), most of the basicecological works (about 71 papers), as well asmorphological studies (about 92 articles).Over 100 papers dealing with distribution usethe name S. obesus. While the name S. ater hasbeen little used and is essentially restricted topublications in technical journals, the name S.obesus appears in numerous papers, magazinesand books, ranging from technical to popular.Clearly, the name S. obesus has had a long his-tory of usage to the present, and is deeplyentrenched in both the scientific and popularliterature. Hence, any proposed change of thislong-recognized name would certainly createextensive confusion and instability.

7. We propose that, if the names Sauromalus aterDuméril, 1856 and S. obesus (Baird, 1858) areconsidered to be synonyms, obesus should beconserved for the combined taxon by giving itprecedence over ater. If the two names areconsidered to refer to different taxa (species orsubspecies), then both names are available foruse. If the application is approved by theCommission both names will be placed on theOfficial List. As mentioned in paras. 1 and 3above, the holotypes of both nominal taxa arein existence.

8. The International Commission on ZoologicalNomenclature is accordingly asked:(1) to use its plenary power to give the name

obesus Baird, 1858, as published in thebinomen Euphryne obesus, precedence overthe name ater Duméril, 1856, as publishedin the binomen Sauromalus ater, when-ever the two are considered to be syn-onyms;

(2) to place on the Official List of GenericNames in Zoology the name SauromalusDuméril, 1856 (gender: masculine), typespecies by monotypy Sauromalus aterDuméril, 1856;

(3) to place on the Official List of SpecificNames in Zoology the following names:(a) obesus Baird, 1858, as published in the

binomen Euphryne obesus, with the


endorsement that it is to be given precedenceover the name ater Duméril, 1856, as pub-lished in the binomen Sauromalus ater, when-ever the two are considered to be synonyms;

(b) ater Duméril, 1856, as published inthe binomen Sauromalus ater, withthe endorsement that it is not to begiven priority over obesus Baird, 1858,as published in the binomen Euphryneobesus, whenever the two are consid-ered to be synonyms.

Comments on this case are invited for publication (subjectto editing) in the Bulletin; they should be sent to theExecutive Secretary, I.C.Z.N., c/o The Natural HistoryMuseum, Cromwell Road, London SW7 5BD, U.K. (e-mail:[email protected]).

III. Changing Paradigms.

Throughout most of history, animals weregrouped into categories using as criteria readilyobservable morphological similarities. Nearly allearly naturalists, who were educated in systemsbased on literal interpretations of the Bible, viewedclassification as a means to better understandingGod’s plan. The underlying basis for the resultinggroups, however, has changed. In the latter half ofthe 19th century, almost entirely in response toCharles Darwin’s work, the interpretation of clas-sification by most scientists changed dramaticallyfrom one based on similarity and a revelation ofGod’s plan to one based on “descent with modifi-cation.” That one could switch from interpretingthe existing Linnaean system of classification as ascheme of similarity to one based on an evolution-ary history is testament to the system’s innate flex-ibility.

Yet, even today, the transition from one schoolof thought to the other is incomplete. This is pred-icated by the reality that animal classification con-tains two distinct and sometimes mutually exclu-sive entities: (1) a purely technical aspectconcerned with establishing a set of rules thatdefine how classifications will be implemented(and which is unconcerned with the philosophy ofthe person describing a new taxon), and (2) a con-ceptual aspect that seeks, above all, to determinejust what a classification means (i.e., identifyingand understanding the underlying factors respon-sible for the relationships that a classification

implies). Some insight into the first aspect was pro-vided by the petition to the InternationalCommission on Zoological Nomenclature in PartII of this series. The second aspect is more diffi-cult, in practice if not in principle.

Willi Hennig, a German systematist, proposedin 1950 a major reform of existing animal classifi-cation schemes, a new approach that has becomeknown as “phylogenetic systematics.” Until itsimplementation and widespread (though not uni-versal) acceptance by practicing taxonomists, therecognition that our understanding of animalgenealogies is woefully incomplete resulted in ageneral willingness to use criteria that rely heavily(or solely) on similarities to define categories. Inother words, “relationships” reflected in taxonomymight be relationships of similarity or relationshipsof genealogical descent or even some mixture ofthe two. Hennig suggested that “relationships,” ina rigorous evolutionary sense, must include onlyconsiderations of descent, defined as the relation-ship between an ancestral species and its descen-dents. No longer could the placement of a taxoninto a particular category be justified by argumentsbased on similarities. Thus, the association of croc-odilians with lizards, because they are superficially

Subadult Iguana iguana from southern Veracruz, México.Traditionally considered a single, widely distributed species,new evidence suggests that I. iguana actually constitutes aspecies complex. As details are resolved, these new data willeventually be reflected by taxonomic changes that moreclosely approximate the reality in nature. Photograph:Robert Powell


similar, was no longer acceptable (genealogically,crocodilians share a much closer common ancestrywith birds and dinosaurs than with lizards andother squamates). Although sound in principle,the new paradigm was difficult to implement inmany instances — mainly because we still lack theevidence necessary to understand many animalgenealogies and, to a lesser degree, because weoften are unwilling to reject familiar categoriesbased on similarities or flawed interpretations ofevolutionary relationships.

Another refinement of classification emergedfrom Hennig’s work. Systematists previously hadrecognized two kinds of groups. “Monophyletic”groups were composed of the descendants of a sin-gle common ancestor and “polyphyletic” groupscontained organisms that did not share a commonancestor. For example, Mammalia is monophyletic,but Homeothermia (a group composed of mam-mals and birds based on their ability to physiolog-ically maintain elevated body temperatures) ispolyphyletic because the two constituent groupshave different “reptilian” ancestors. Hennig notedthat many recognized “monophyletic” groupswere not really monophyletic at all, and recog-nized a third type of group. Because these assem-blages did not contain all of the descendants of acommon ancestor, he called them “paraphyletic.”Phylogenetic systematists asserted that paraphyleticgroups were as unnatural as polyphyletic groups.

The most easily understood example is the ClassReptilia, as traditionally defined. Reptilia consti-tutes a paraphyletic group because some descen-dents (mammals and birds) are left out. Anotherexample is the Family Pongidae, which consists ofthe great apes but excludes humans (who areplaced in their own family, Hominidae). Becauseparaphyletic groups are logically inconsistent withthe phylogenies that classification should reflect,the idea that we should abandon traditionalschemes that include paraphyletic groups is grow-ing in acceptance (although almost every textbookin print presents classifications that include para-phyletic groups).

How would a revised, purely phylogeneticclassification of an iguana look? Actually, theanswer is less than obvious, because the criteriathat define the higher categories in particular areoften vague and frequently subject to revision.However, one scheme (modified from a phylogenypresented in Pough et al. 1998. Herpetology.Prentice-Hall, Inc., Upper Saddle River, NewJersey) might look a lot like this:

Chordata (animals with notocords, pharyngeal gill slits, anddorsal, hollow nerve cords)

.Vertebrata (chordates with at least rudimentary braincases andvertebral columns)

..Sarcopterygii (lobe-finned fishes and their descendents)

…Tetrapoda (amphibians and their descendents)

….Amniota (those tetrapods that reproduce via cleidoic eggs;excludes amphibians)

…..Reptilia (excludes the common ancestor of mammals and allof its descendents)

……Diapsida (excludes “reptiles” with anapsid, synapsid, andeuryapsid skull structures)

…….Lepidosauria (the common ancestor of squamates andrhynchocephalians and all of its descendents)

……..Squamata (lizards and snakes)

………Iguania (non-scleroglossine lizards; includes acrodonts aswell as iguanids in the broad sense)

……….Iguanidae (the oldest common ancestor of all species inthe genera Amblyrhynchus, Brachylophus, Conolophus,Ctenosaura, Cyclura, Dipsosaurus, Iguana, andSauromalus and all of its descendents)

………..Iguana (the oldest common ancestor of I. iguana and I.delicatissima as currently defined and all of its descen-dents)

………...Iguana delicatissima (the binomen, indicating a groupconsisting of the oldest common ancestor of allpopulations now assigned to I. delicatissima and all of itsdescendents)

Adult male Cyclura ricordii from Parque Nacional IslaCabritos, Dominican Republic. The generic name (“ring-tail”)is based on the whorls around the tail, which are very evi-dent in this specimen. The trivial name is a possessivepatronym (indicated by the -ii ending), which honorsAlexandre Ricord, who collected the type specimen.Photograph: Robert Powell


An examination of this example quickly pointsto a problem inherent in a phylogenetic classifica-tion, namely that the number of groups can easilyexceed the number of Linnaean categories. Somesystematists have proposed alternatives that includeusing numerical prefixes instead of categoricalnames or even a “rank-less” system in which thehierarchy is indicated merely by indenting (asabove). Unfortunately, these alternatives are notwithout problems of their own. Numerical systemscan be applied only to one phylogeny withoutbecoming confusing (does a particular numberhave the same meaning in each context?), andrank-less systems quickly become so cumbersomethat determining who is related to whom is impos-sible (note that the example includes only onealternative at each level; if all chordate groups wereincluded, the classification easily would fill manylarge volumes).

IV. What Classification Can —

and Cannot — Do

Modern classification should demonstrate therelationship between phylogeny and taxonomy byidentifying and naming hierarchical groups thatdefine genealogies. That recognition acknowledgesthe underlying hypotheses that a classificationseeks to examine, namely that groups reflectgenealogical relationships. If a classification com-pares favorably with a phylogenetic tree, thehypothesis is supported. Unfortunately, the lack ofavailable evidence for relationships of many groupsoften results in classifications based on similaritiesor even the intuition of the systematist. Theseoften are merely efforts to organize diversity untila group can be adequately studied. Obviously,newly acquired evidence renders many of thesetentative schemes unacceptable, and this is at thevery root of many taxonomic changes. The dis-tinction between classifications based on greatquantities of pertinent evidence and those thatreflect mere “guesses” is impossible — unless thereader is very familiar with the group(s) being clas-sified. However, in both instances, a reader shouldremember that any classification is based only onhypotheses addressing the group relationships thatexist between the organisms being classified.

The rank of a group does not necessarilyreflect the distinctiveness of that taxon. A system-

atist may raise the rank of a group to reflect herviews of its distinctive nature, but rank in and ofitself does not provide that information. For exam-ple, when Frost and Etheridge (1989. A phyloge-netic analysis and taxonomy of iguanian lizards(Reptilia: Squamata). Univ. Kansas Mus. Nat. Hist.Misc. Publ. (81):iv + 65 pp.) elevated what hadbeen considered “iguanines” or “Iguaninae” tofull familial rank (“Iguanidae”), the intent was tobetter reflect historical events, not to suggest thatthe group was any more distinctive than had beenpreviously recognized.

Emphasizing the fact that rank does notdenote degrees of distinction is the reality thatranks cannot be and are not applied equitably todifferent groups. In other words, the ranks in anygiven classification represent genealogical relation-ships by subordinating descendent groups to thoserepresenting older (ancestral) taxa; thus, the num-ber of ranks merely reflects the relationships in aparticular group rather than being definitive arbi-trators of a certain level of distinction. In otherwords, a family of insects is not comparable to afamily of squamates (as a matter of fact, if equiva-lency were the goal, an insect family may equatewith an order or even a class of vertebrates).Rather than being a defect of modern classificationschemes, these inequalities testify to the system’sflexibility. The idea that unrelated (actually dis-tantly related) groups of comparable rank are bio-logically equivalent dates to the antiquated per-ception that all types of organisms representedrungs on a “ladder of life” that represented a scaleculminating in “perfection” (in these ancientschemes, humans invariably resided on the toprung; what a humble species we are…).

Finally, classification cannot remain stable andunchanged. All taxonomic decisions are, in fact,hypotheses subject to additional testing and poten-tial rejection when new contradictory data areuncovered. Consequently, like science in general,classification must be dynamic — and this is astrength rather than a flaw of the system. If classi-fication were stagnant, it could not be scientific.Although we all bemoan the need to constantlyrelearn new classifications, we should instead betoasting the changes that reflect new knowledge,scientific progress, and are probably closer to thetruth represented in nature.


V. Do ‘Species’ Exist In Nature?

When I was an undergraduate, I was taughtthat the species was the only “real” taxonomicrank, based on the assumption that it could besubjected to testing by applying criteria applicablein nature. Of course, the concept of species towhich reference was being made was the “biolog-ical species,” defined as a group of similar, repro-ductively interacting (or potentially interacting)organisms reproductively isolated from all othersuch groups. The emphasis on reproduction cer-tainly was appropriate. I often tell my students thatthe primary role of individual organisms in natureis to reproduce, and that all other functions aresecondary. Also, the contention that reproductiveisolation was testable appeared reasonable — butwas it? I began to have doubts when discussionsveered to the many exceptions. Obviously, theprincipal criterion was not applicable to asexuallyreproducing organisms or to parthenogeneticorganisms (those all-female species in which off-spring are produced by development of an unfer-tilized ovum), but could it even be applied ade-quately to sexually reproducing populations withnon-overlapping distributions? If organisms aregeographically isolated (allopatric), is any test ofreproductive isolation appropriate? When I beganto examine examples with which I was familiar(allopatric “subspecies” of North Americanamphibians and reptiles or insular populations ofWest Indian forms), I found that assumptionsbased on degrees of morphological similarity orsimple guesswork invariably were substituted forempirical testing. This conclusion then led me intoa review of the voluminous literature pertaining tospecies concepts.

The best summary of my search for answerscame when I ran across several papers whichemphasized that speciation was a process, and that“species,” variously defined, could exist at anystage in that progression. The biological speciesconcept applied accurately only to groups of sexu-ally reproducing organisms that had essentiallycompleted the process. At the other extreme werepopulations of common ancestry that had onlyrecently become isolated (geographically or eco-logically) and had barely begun to differentiate asa consequence of differing selective pressures,mutations, or random changes in genetic compo-

sition often associated with small population sizes.Some of these isolates will never diverge to anextent sufficient for recognition as a distinctspecies, others may do so eventually — even to theextent of becoming reproductively isolated. Manyof the competing species concepts actually addresssuch “works in progress.” For example, the “evo-lutionary species concept” recognizes populationsas species that have diverged enough to be diag-nosable and which, due to geographic (or ecolog-ical) isolation, have unique evolutionary trajecto-ries. Against the argument that such divergingpopulations would readily interbreed if they shouldcome back into contact with one another, propo-nents of the “evolutionary species” rightly notethat science cannot deal with events that have yetto occur.

So, where does that leave the biologist who istrying to assess the biodiversity of a region (typi-cally expressed as the number of species present),establish a conservation plan for a particular taxon(generally, species-level taxa are provided greaterconsideration than subspecies or isolated popula-tions when management plans are developed),develop a captive-breeding program with thepotential for the eventual release of progeny intothe wild (releasing hybrids into naturally occurringpopulations will merely dilute or pollute the exist-ing gene pool), or even engage in research usingas a model a particular species, the identity ofwhich is crucial (if only to provide an accurate labelfor the model)? Actually, the answer is not that dif-ficult once agreement can be reached on a com-mon definition of “species.” The best I have foundis that a species is a natural entity that derives itsexistence from historical evolutionary (ancestral)relationships (a phylogenetic emphasis), inter-breeding (the traditional biological species con-cept), or some combination of both. If we canaccept that definition (recognizing as we do thatspeciation is a dynamic process), then the “generallineage concept” should be acceptable to all. Thisconcept recognizes species as segments of popula-tion-level lineages — and all contemporary speciesdefinitions, in one way or another, consider speciesto be segments of population lineages.Consequently, all species concepts are componentsor variations of this main underlying concept,although they may variously emphasize differentdiagnostic criteria.


What does thismean in the “realworld?” It means thatspecies definitions willvary to some degreedepending on the cri-teria applied by theresearcher and heragenda (if interested inconservation, shemight be more inclinedto recognize a popula-tion as a full speciesthan, for example,another researcher whomerely wants a label forhis model, and wouldprefer that which ismost familiar to his col-leagues). To avoid suchvarying standards,some systematists havesuggested substitutingthe term “operationaltaxonomic unit” forspecies. This apparentlyunconventional approach might seem strange atfirst, but has the very distinct advantage of settingaside disagreements over which species-definingcriteria to emphasize by focusing on the actualentity — the population(s) that exist in nature.

How does this apply to iguanas? CatherineMalone (2000. Mol. Phylogen. Evol. 17:269–279)presented data suggesting that green iguanas fromdifferent regions in the Western Hemisphere aregenetically distinct. These distinctions may or maynot lead to interpretations suggesting that Iguanaiguana, as currently defined, is actually a complexof separate species. However, the conservationistand the breeder can no longer, in good con-science, treat all green iguanas as if they were inter-changeable. Conservation of unique insular popu-lations, regardless of taxonomic rank, should be ahigh priority when management plans are devel-oped and implemented. Allowing animals fromdifferent areas to breed could result (and hasundoubtedly in many instances resulted) in hybridsthat are no more “natural” than the “designersnakes” bred by some hobbyists.

In that same paper, Malone demonstrated avery close relationship between populations ofCyclura on Hispaniola (C. cornuta) and Isla Mona(C. stejnegeri). I have recently advocated thespecies-level recognition of the Mona Island pop-ulation based primarily on allopatry and differencesin morphological and biological (reproductive)traits. If additional data indicate that these popula-tions do not differ sufficiently to warrant full-species status (suggesting that the differencesmerely reflect local conditions and that the twoentities have only recently become separated), thegeneral lineage concept would still apply. Any con-servation plans that emphasize the preservation ofboth sets of populations would not be any lessvalid, nor would efforts to separate any captivepopulations by their origin be any less important.

Do ‘species’ exist in nature? Sure, but theiridentity may or may not correspond to our effortsat defining them. Plus, the animals don’t read ourtextbooks and they certainly don’t care what wecall them.

Adult Cyclura stejnegeri from Isla Mona. This population was originally described as a speciesdistinct from populations of C. cornuta on Hispaniola. Recent evidence indicates, however,that Mona Island animals are not genetically distinct, suggesting a very recent (possibly evenhuman-mediated?) colonization of the island by Hispaniolan stock. Photograph: RobertPowell


“

You mustn’t speak to the animals or theywon’t take you seriously. These peopleare scientists working with criticallyendangered species.” I was having this

conversation with my dear friend and house guestat the time, John Bendon. John has been workingwith both the International Iguana Society and theWest Indian Iguana Specialist Group for manyyears, and is well acquainted with the scientistsworking at the San Diego Zoo. My husband wasattending a medical conference in San Diego andJohn had set up appointments for me to meetsome important people and some important ani-mals at the Zoo and the affiliated Centre forReproduction of Endangered Species (CRES).Like many of our readers, I knew only aboutspecies such as Iguana delicatissima, Cyclurapinguis, and Brachylophus fasciatus from the pages

of Iguana Times and other reptile publications,and this was my best and probably only chance ofever seeing live specimens.

I was determined to appear as “serious” aspossible, so I went rifling through back issues andunearthed articles on all of the above species.None of the material was unfamiliar to me becauseI’ve been teaching students, from preschoolers touniversity-level, about iguanas and conservationfor years. Still, I was certain that I would somehowmanage to embarrass myself.

Between the Zoo and the Wild Animal Park, Ispent my first four days in San Diego admiring allkinds of remarkable animals, wonderful exhibits,and excellent presentations, and reading about theexciting work being done with the pandas,California condors, and other endangered species.As awe-inspiring as the whole experience was, I

was saddened to seepeople persistentlymisunderstanding ani-mals that were differ-ent in any way. I recallthe family (mother,father, and child)watching three mag-nificent condors thatwere perched togetheron a rock. They werepicking at each otherand the child thoughtthey were fighting.The father startedegging them on andthe mother pulled herfamily away from the“disgusting” display. Ihad to stop myselffrom shouting at them.Couldn’t they see thatthis was a mother,father, and a juvenilethat were grooming

A Day at the ZooAJ Gutman

West Hartford, CT

Jeff Lemm of the San Diego Zoo holding a large Cuban iguana (Cyclura nubila nubila).Photograph: Steve Steward


each other and enjoying a lovely afternoon ontheir rock? How much more do people fail toidentify with the plight of the reptiles?

By the day of my appointments most of thegloom had vanished, but I was still a little appre-hensive about appearing sufficiently “serious.” Atthe CRES, I was met by Jeff Lemm, the ResearchAnimal Coordinator. Jeff struck me immediately assomeone who took great pride in his work andcared very deeply for the animals. I suspect he alsohas had plenty of experience entertaining goofyvisitors, because he made me feel quite welcome ashe introduced the animals and let me know whichones I could hug and which were being kept withas little human contact as possible in order toencourage breeding.

I was first introduced to three substantialwhite-throated monitors that are used in behav-ioral experiments. Then I had the pleasure ofobserving Ruby, a beautiful Komodo dragon,being fed her lunch of specially prepared “dragonmeatballs.” Ruby was quite intent on her meal anddidn’t seem the least bit troubled by anyone’sboots.

Subsequently, I got to meet the iguanas.Gitmo the Cuban iguana (Cyclura nubila nubila)has had his handsome face featured on Reptilesmagazine. At 15 years of age and an impressive 17pounds, he is a relaxed and charming spokesiguanaoriginally from Guantanamo Bay. The Cubaniguanas of Guantanamo Bay were one of the firstpopulations studied by CRES scientists to test con-servation strategies such as headstarting and cap-tive breeding. Information collected from this con-

Above: A large Jamaican iguana (Cyclura collei).Right: Male and female Iguana delicatissima.Photographs: Jeff Lemm


tinuing study has proven invaluable in assistingother endangered species of Cyclura.

The story of the Jamaican Iguana, Cycluracollei, has been well documented in the pages ofIguana Times since their rediscovery in 1990 aftersupposed extinction to the successful headstartprogram at the Hope Zoo in Kingston. Mostrecently, I recall an article about the neoprene vestsproduced by Nike to hold the radiotransmitters fortracking the headstarted iguanas. Even these high-tech vests have proven inadequate over time. Anew strategy involving surgically implanting thetransmitters has been tested and shown not toimpair egg laying or mobility in captive-heldCuban iguanas. Three pairs of C. collei have beenat CRES since 1992, including the youngest in anyUS zoo. One of the females nested successfully inthis, her first lay year, although her nine eggs wereinfertile. Still, this was an important first step anddefinitely cause for optimism. Even though I knowthat the orange col-oration typical ofJamaican iguanas pho-tographed in the wildis caused by the highiron content in the soilof their native habitat, Ithought it odd to see“clean” C. collei sport-ing the usual attractiveCyclura striping.

The Anegada or“Stout” iguana, Cyclurapinguis, has been a par-ticular favorite of minesince reading an articleby James Lazell inIguana Times in 1997.This animal, Lazellwrites, is described inlocal folklore as attain-ing “weights of morethan 60 pounds andlengths over 6 feet.”The species varies in anumber of ways fromother members of thegenus Cyclura. Mostsignificantly, it lacksrings of enlarged cau-

dal scales, a defining featuring of “Cyclura,” whichliterally means, “ring tail.” Both the teeth and theskull bones also differ from those of other species,and the young C. pinguis, although displaying theusual striped patterning of young rock iguanas arevarious shades of green and become very dark,almost black as they mature. Some researchershave speculated that C. pinguis may be the stempopulation from which all the other species ofCyclura are descended.

At one point, fewer than 200 C. pinguis werethought to remain in the wild. Only isolated pop-ulations survived the hunting of previous cen-turies, and the remaining animals have had to con-tend with development and competition from ferallivestock. Translocated populations on Guana andNeckar islands have experienced some success, butCRES has concentrated its efforts on a headstartprogram on Anegada, with already 50 animalsbeing raised for eventual release.

John Kinkaid of the San Diego Zoo holds a Fijian banded iguana (Brachylophus fasciatus).Photograph: AJ Gutman


CRES has housed three pairs of Stout Iguanassince the fall of 2000. These animals had beenlegally purchased and in the possession of a privatebreeder for ten years before coming to the zoo. Iwas impressed by their dark, glowering looks, butI suppose I was expecting them to be bigger thanthey were. Jeff was overwhelmingly skeptical aboutthe 6 foot-60 pound iguana story, claiming thelargest he had worked with was only a little over 1m long and about 6–7 kg. Personally, I like theidea too much to abandon it just because the reallybig one has so far managed to elude everyone.

Iguana delicatissima is another remarkablespecies. It is the only other member of the genusIguana beside the green iguana. It is somewhatsmaller than its closest relative. Juveniles are brightgreen, but the adults become predominantly greywith the males acquiring pink highlights in themating season. Wild populations of I. delicatissimaface the usual obstacles caused by introducedpredators, habitat destruction, and fragmentationof populations, but they also are faced with com-petition from and hybridization with greeniguanas. Iguana delicatissima produces clutchesthat are intermediate in size between those ofgreen iguanas and species of Cyclura. Eggs also areintermediate in size. Only seven animals are in cap-tivity in only three institutions in the world. Thepair at the Jersey Wildlife Trust have bred success-fully (see report in Iguana Times 8(4)) but twoeggs from a successful mating this year in SanDiego failed to hatch.

I could have spent days at CRES, marveling atall the animals and asking a million questions, butI had only five minutes to run over to the zoo formy next appointment at the Reptile House. I wasonly a little sweaty and no more disheveled thanusual by the time I arrived, struggling to regain my“serious” demeanor.

Again, I was in for a big treat. John Kinkaid,the Animal Care Manager, was clearly as enthusi-astic about his animals as I was. “What have youseen so far? You can take out any of the Bandediguanas if you want to photograph them. Is thereanything else you’d like to see?” John spent thenext 2 1⁄2 hours showing me all of his charges. I sawlong-necked tree monitors and endless species ofboth rare and common snakes. I held young blue-tongued skinks that looked like fat sausages andwas loath to disturb the young Corucia who

climbed on top of my head to nest in my hair. Asa long-time dinosaur admirer, I had known aboutthe remarkable tuatara long before I developed aninterest in reptiles. The tuatara (genus Sphenodon)is a lizard-like creature that has been around sincedinosaurs roamed the earth and it has no othersurviving relatives. Few of them, however, existtoday. The tuataras at the San Diego Zoo aremembers of the highly endangered Brothers Islandpopulation (S. guentheri) and are the only oneskept anywhere outside of New Zealand. We wentoutside to meet the Galápagos Tortoises; Johnknew them all by name and knew which individu-als were grumpy and which would come gallopingover to take the sweet potatoes we brought astreats.

My favorites, of course, were the FijianBanded Iguanas, tiny, brilliantly colored creatureswith alert eyes and calm temperaments. I particu-larly enjoyed holding the babies and a three-leggedfemale who had lost a limb to infection. I was a lit-tle surprised to learn that her name was Ilene (I lean) and it left me wondering whether the I. delicatissima were named Hors d’oeuvre andEntrée (John has promised to let me name someof the next Brachylophus babies that hatch).

For days, even after returning home, I couldstill close my eyes and see so many beautiful faces,sense the weight of plump, healthy bodies, and feelwonderfully textured skin. Although much workremains to preserve all of the endangered animalsof the world, we can only applaud the work beingdone on behalf of the reptiles by the dedicatedstaff at the San Diego Zoo.

ReferencesAlberts, A. and J. Lemm. 1997. Guided by nature: Conservation

research and captive husbandry of the Cuban iguana. Reptiles5(8):76–87.

Lazell, J. 1997. The Stout Iguana of the British Virgin Islands.Iguana Times 6(4):75–80.

Lemm, J. 2000. Conservation of the Lesser Antillean Iguana,Iguana delicatissima. Vivarium 11(2):22–25.

Ed. Note: The Cyclura pinguis housed at CRES laid eggs in July 2001. After 90 days incubation at 30° C, fourhatchlings emerged. This represents the first captivebreeding of this species in a zoological facility.

Alberts, A.C. 1999. Conservation spotlight: developingrecovery strategies for West Indian rock iguanas.Endangered Species Update 16:107–110.

Alberts, A.C. (ed.). 2000. West Indian Iguanas: StatusSurvey and Conservation Action Plan. IUCN/SSC WestIndian Iguana Specialist Group, IUCN-The WorldConservation Union, Gland, Switzerland.

Banbury, B.L., Y.M. Ramos, R. Powell, and J.S.Parmerlee, Jr. 2000. The Cyclura of Parque NacionalIsla Cabritos. Journal of the International IguanaSociety (Iguana Times) 8(2):3–7.

Bendon, J. 1998. It take two to tango on Booby Cay.Journal of the International Iguana Society (IguanaTimes) 7(3):37–42.

Berovides, V. 2001. Usos potenciales de la iguana cubana(Cyclura nubila nubila). Revista Biología 15:3–8.

Crother, B.I. (ed.). 1999. Caribbean Amphibians andReptiles. Academic Press, San Diego, California.

Ehrig, R.W. 1998. Progress for Cyclura rileyi cristata.Journal of the International Iguana Society (IguanaTimes) 7:27–28.

Glor, R.E., R. Powell, and J.S. Parmerlee, Jr. 1998.Cyclura ricordii. Catalogue of American Amphibiansand Reptiles (657):1–3.

Glor, R.E., R. Powell, and J.S. Parmerlee, Jr. 2000.Cyclura cornuta. Catalogue of American Amphibiansand Reptiles (709):1–6.

González, A., V. Berovides, and M.A. Castañeira. 2001.Aspectos de morfometría, abundancia, y alimentaciónde la iguana cubana (Cyclura nubila nubila) en elArchipeélago de los Canarreos, Cuba. Revista Biología15:98–104.

Hartley, L.M., R.E. Glor, A.L. Sproston, R. Powell, andJ.S. Parmerlee, Jr. 2000. Germination rates of seedsconsumed by two species of rock iguanas (Cycluraspp.) in the Dominican Republic. Caribbean Journal ofScience 36:149–151.

Knapp, C. 1998. Morphologic characters of herbivorouslizards. Journal of the International Iguana Society(Iguana Times) 7(1):11–17.

Knapp, C. 1998. Vanishing iguanas. Journal of theInternational Iguana Society (Iguana Times)7(3):27–30.

Knapp, C.R. 1999. Population biology of a translocatediguana (Cyclura) in the Bahamas. Unpubl. M.Sc. Thesis,University of Florida, Gainesville.

Knapp, C.R. 2000. Home range and intraspecific interac-tions of a translocated iguana population (Cycluracychlura inornata Barbour and Noble). CaribbeanJournal of Science 36:250–257.

Knapp, C., S. Buckner, A. Feldman, and L. Roth. 1999.Status update and empirical field observations of theAndros rock iguana, Cyclura cychlura cychlura.Bahamas Journal of Science 7:2–5.

Malone, C.L., T. Wheeler, J.F. Taylor, and S.K. Davis.2000. Phylogeography of the Caribbean rock iguana(Cyclura): implications for conservation and insights onthe biogeographic history of the West Indies.Molecular Phylogenetics and Evolution 17:269–279.

Martins, E.P. 1998. Estimating ancestral states of acommunicative display: a comparative study of Cyclurarock iguanas. Animal Behavior 55:1685–1706.

Mitchell, N.C. 1999. Effect of introduced ungulates ondensity, dietary preferences, home range, and physicalcondition of the iguana (Cyclura pinguis) on Anegada.Herpetologica 55:7–17.

Mitchell, N. 2000. Anegada iguana (Cyclura pinguis), pp.22–27. In R.P. Reading and B. Miller (eds.), EndangeredAnimals: A Reference Guide to Conflicting Issues.Greenwwod Press, Westport, Connecticut.

Powell, R. 1999. Herpetology of Navassa Island, WestIndies. Caribbean Journal of Science 35:1–13.

Powell, R. 2000. Cyclura onchiopsis. Catalogue ofAmerican Amphibians and Reptiles (710):1–3.

Powell, R. 2000. Horned iguanas of the Caribbean. Reptile & Amphibian Hobbyist 5(12):30–37.

Powell, R. and R.E. Glor 2000. Cyclura stejnegeri.Catalogue of American Amphibians and Reptiles(711):1–4.

Powell, R. and R.W. Henderson. 1999. Addenda to thechecklist of West Indian amphibians and reptiles.Herpetological Review 30:137–139.

Powell, R., J.A. Ottenwalder, S.J. Incháustegui, R.W.Henderson, and R.E. Glor. 2000. Amphibians andreptiles of the Dominican Republic: species of specialconcern. Oryx 34:118–128.

Rodríguez Schettino, L. (ed.). 1999. The Iguanid Lizardsof Cuba. University Press of Florida, Gainesville.

Wasilewski, J. 1998. Booby Cay update. Journal of theInternational Iguana Society (Iguana Times)7(2):29–30.

Recent Literature Pertaining to West Indian Iguanas

From the West Indian Iguana Specialist Group Newsletters 1999 2(2), 2000 3(1), 2000 3(2), 2001 4(1)


IntroductionCalcium plays a key role in many biological

processes, such as muscle contraction, egg shellproduction, blood clotting, enzyme activity, ner-vous system function, hormone release, and cellmembrane permeability, in addition to being anessential component of bones. Calcium in theblood can be bound to other ions such as phos-phate and citrate. It can also exist in the biologi-cally active or free form. The blood concentrationof calcium and its associated ions, as well asabsorption and use by the body organs, is regu-lated by three major factors that interact indepen-dent of calcium intake. These factors are parathy-roid hormone (PTH), calcitonin (CT, anotherhormone), and vitamin D. Parathyroid glands,which are located in the neck and which secretePTH, and the thyroid gland, which secretes CT,are present in all air-breathing vertebrates.

AbsorptionVitamin D3 (cholecalciferol) is converted by

the liver and kidney to its biologically active form,1, 25-dihydroxycholecalciferol (1,25 DHCC). Thisform is necessary for an iguana to properly absorbcalcium from digested food in its intestine. VitaminD3 can be provided in the diet either as a pre-formed molecule or be converted from other mol-ecules in the iguana’s skin in the presence of ultra-violet stimulation. Dietary vitamin D can either beprovided directly in specific food items or as part ofa supplement added to the food prior to feeding.Iguanas utilize vitamin D3 rather than ergocalcif-erol, vitamin D2. Ergocalciferol is derived fromplants and is less expensive to produce than chole-calciferol. Supplements listing vitamin D without areference to the specific type usually contain ergo-calciferol. Some food items such as commercial dogor cat food or monkey biscuits contain excessiveamounts of vitamin D. When these items are fed toan iguana, hypervitaminosis D will occur. Likewise,

if the iguana is over-supplemented with vitamin D,the condition will also occur.

Vitamin D3 can be synthesized in the skin ofan iguana from precursor molecules. Ultravioletlight at wavelengths of 290–320 nm (UVB), shin-ing on the iguana, is necessary for this reaction totake place. Following this reaction, the vitamin D3from the skin is transported in the blood to theliver and kidney where the conversion process intothe biologically active form, 1,25-DHCC, is com-pleted. Necessary wavelengths of ultraviolet radia-tion can be provided either from direct, unfilteredsunlight or a high quality, full-spectrum lightsource. This is the preferred manner for vitaminD3 to be provided because an overdose is less likelyto occur.

Unfortunately, glass or plastic will filter ultra-violet radiation out of sunlight, providing onlyheat and light. Likewise, poor quality or old ultra-violet lights will not provide adequate stimulationfor proper vitamin D3 conversion. Full-spectrum,ultraviolet strip lights will provide a better range ofultraviolet irradiation than incandescent or screw-in type bulbs. Currently, my favorite light is ZooMed’s Iguana Light 5.0 and Reptisun 5.0. Thesehappen to be the same bulb with different packag-ing. Regardless of the brand you choose, the bulbshould be replaced every 6–9 months. Althoughthe bulb may still be producing light, the func-tional spectrum of ultraviolet irradiation will beinsufficient for proper vitamin D3 metabolism.Once the bulb develops a dark band near eachend, the filament necessary to provide the ultravi-olet spectrum has been burnt out. Remember,however, that no bulb can replace natural sunlight.

StorageThe main storage area for calcium in all verte-

brates is in the bone matrix. The amount of cal-cium stored at any given time is a function of theaction of two specific cell groups, the osteoblastsand the osteoclasts. Osteoblast cells are responsi-

Calcium Metabolism in IguanasBruce Bogoslavsky, DVM

Animal Veterinary Hospital of OrlandoOrlando, FL 32809


ble for the resorptionof calcium from bloodand the production of bony matrix.Conversely, osteoclastcells are responsible forthe demineralizationof bone and a releaseof calcium into theblood.

When blood cal-cium levels decrease,the parathyroid glandsecretes PTH. PTHstimulates osteoclastactivity and inhibitsosteoblast activity,which results in therelease of stored cal-cium from the miner-alized bone. PTH alsocauses an increase inphosphorus excretionby the kidneys. Lastly,PTH stimulates thefinal conversion of D3to metabolically useful1,25 DHCC. The endresult of all this activityis to increase the cal-cium absorption fromdigested food in theintestine. Once theblood calcium levelreturns to normal, thethyroid gland secretes CT. This hormone inhibitsthe release of stored calcium from bone bydecreasing osteoclast activity. It also inhibits theexcretion of phosphorus by the kidneys.Phosphorus is significant in that it can bind withfree calcium and render it unavailable for use bythe iguana’s systems.

Cardiac EffectVertebrate cardiac function is also influenced

by calcium ions within the myocardium (heartmuscle). Calcium is rapidly exchanged within themyocardium. As calcium levels increase so doesthe animal’s heart rate. Small elevations of calciumwithin the heart cells will result in increased con-

traction strength. However, as the concentrationincreases, the animal may develop a cardiacrhythm disturbance that may eventually lead to acardiac arrest.

Reproductive EffectBlood protein levels vary depending on the

season, reproductive state, nutritional state, andtemperature. The proteins in the blood that bindand transport calcium are highly influenced by thereproductive cycle. As a consequence of increasedestrogen activity, blood calcium levels will increasein female iguanas as egg follicles are producedwithin the body. Thus the gravid iguana will notdeplete its own calcium stores during reproduction.


ExcretionIn a normal metabolic situation, a dietary excess

of soluble calcium will not be absorbed by the intes-tine and will be excreted in fecal material.Additionally, a small amount of calcium is excretedwith the urates. Likewise, only trace levels of calciumare present in the iguana’s salt gland excretions.

HypercalcemiaExcessive amounts of calcium in an iguana’s

diet will only cause a problem if it is accompaniedby an excessive amount of vitamin D3. This willhappen when an owner is adding a vitamin-min-eral supplement to an already high quality diet. Inaddition, the owner may be providing direct unfil-tered, natural sunlight or full-spectrum, ultravioletlighting. This can result in an excessively highamount of calcium being absorbed from digestedfood within the iguana’s intestines. Ultimately, thiswill cause calcium salts to be deposited in varioussoft tissues. The most commonly affected sitesinclude the aorta, heart muscle, pulmonary air-ways, gastrointestinal tract, and urinary system.

HypocalcemiaHypocalcemia (low blood calcium levels) can

be caused by a number of different factors.Growing iguanas require a quality diet containingan adequate amount of calcium without an excessof phosphorus. Because calcium and phosphoruscompete for the same binding sites in the blood, adiet high in phosphorus will result in low bloodcalcium levels. Sensing this decrease in circulatingblood calcium, the parathyroid gland will secretePTH. PTH stimulates the release of calcium storedin the iguana’s bones. If the blood phosphoruslevel is still elevated at this time, the majority offreed calcium will be unable to find free bindingsites. This results in the excretion of calcium withinstool, leaching it away from bone and leaving theiguana with very soft bones. Some commonly feditems that are excessively high in phosphorus arelettuce, spinach, sprouts, tofu, peas, grapes,banana, mealworms, and crickets.

Oxalic acid in the diet should also be moni-tored as it will bind with calcium and prevent itsabsorption from the intestine. Foods containingoxalic acid such as spinach, rhubarb, cabbage, peas,potatoes, and beet greens should be fed in smallamounts or avoided entirely.

ConclusionCalcium metabolism in iguanas is a complex

process with several variables. Specific wavelengthsof ultraviolet stimulation from either direct, unfil-tered sunlight or a high quality, full spectrum lightsource are necessary to convert vitamin D3 withinthe iguana’s skin. Vitamin D3 must then be trans-ferred first to the liver and then to the kidneysbefore it is finally converted into its biologicallyactive form. Biologically active vitamin D is neces-sary to aid in the absorption of calcium fromdigested food in the iguana’s intestine.

I try to make my clients visualize their iguanain the middle of a triangle with ultraviolet stimula-tion, optimum environmental temperature, and ahigh quality diet on the three corners. An optimalenvironmental temperature is necessary to activateintestinal microbes for proper digestion. A highquality diet, rich in calcium and low in phospho-rus, should be provided. If all of these factors areprovided, an iguana will have solid bones, a goodgrowth rate, a strong cardiovascular system, andwill be a fine specimen for breeding.

ReferencesFrye, F. 1991. Reptile Care. An Atlas of Diseases and Treatments.

T.F.H. Publications Inc., Neptune City, New Jersey.

Frye, F. 1993. A Practical Guide for Feeding Captive Reptiles.Krieger Publishing Co., Malabar, Florida.

Frye, F. and W. Townsend. 1993. Iguanas: A Guide to Their Biologyand Captive Care. Krieger Publishing Co., Malabar, Florida.

Gans, C. and F.H. Pough. 1982. Biology of the Reptilia. Volume 12.Physiology C. Academic Press Inc., London.

Mader, D. 1996. Reptile Medicine and Surgery. W.B. Saunders Co.,Philadelphia, Pennsylvania.

Marcus, L. 1981. Veterinary Biology and Medicine of CaptiveAmphibians and Reptiles. Lea & Febiger, Philadelphia,Pennsylvania.

McDonald, L.E. 1980. Veterinary Endocrinology and Reproduction.Lea & Febiger, Philadelphia, Pennsylvania.


Calcium-related pathology can be causedby several different factors and result in anumber of different conditions, which arecollectively known as Metabolic Bone

Disease or MBD. Some of the most commoncauses of MBD are low dietary calcium levels, anexcess of dietary phosphorus over calcium, exces-sive or deficient intake of vitamin D, and a lack ofultraviolet light in the proper wavelength for vita-min D conversion (UVB).

In the case of cacium/phosphorus ratioimbalance (i.e., an excess of dietary phosphorusover calcium), calcium stored in the bones isdrawn out in order to maintain proper serum calcium levels. The early signs of this condition arethe gradual swelling of limbs and uneven bone

growth, particularly in the jaw (Figure 1). A con-dition known as fibrous osteodystrophy produceswhat appear to be “chubby” limbs, which actuallyconsist of fibrous tissue built up in order to stabi-lize the weakened bone (Figure 2).

Calcium-related Pathologies in Green Iguanas

AJ GutmanWest Hartford, CT


Figure 1Bunny is a ten year old male iguana, who is 4-1/2 ft long and weighs 14 lbs. He was given his name because he had beenfed nothing but shaved carrots for two years before he was adopted and had an overwhelming case of hypervitaminosis A.He was fluorescent orange from head to tail. Urine, feces and shed skin were orange for weeks even after his diet improved.The one lingering sign of his early malnutrition is an overgrown lower jaw. When he is relaxed (most of the time), his tongueprotrudes slightly, giving him a rather comical expression. Photograph by Carole Saucier

Figure 2The muscular appearance ofDante’s legs is caused byfibrous tissue built up inresponse to demineralization.Photograph by CaroleSaucier

Softening of the bone may also occur in thecase of a vitamin D deficiency. If the diet containsinsufficient vitamin D3 or ultraviolet light exposureis insufficient for the vitamin to be synthesizedunder the skin, the result is identical: calcium willnot be absorbed from the intestine, even if it ispresent. Calcium is instead leached from the bonesto maintain serum levels. The joints can becomeswollen and the demineralized bone can be easilyfractured. Another sign of MBD is an inability tolift the trunk (Figure 3), first with the back legsand then with the front, to the point where theanimal is virtually unable to move its body eventhough the legs move vigorously. Spinal deformi-

ties may occur, and the spinal column can break,resulting in paralysis of the rear limbs. Anothervery obvious result is general failure to thrive,resulting in various degrees of stunting (Figure 4).

Hypocalcemia, or low blood calcium levels,will produce fine muscle tremors, which begin inthe digits and progress to the limbs and the tailbase. In severe cases, full seizures and tetany(locked limbs) can result. The animal will eventu-ally become completely flaccid and die.

Hypercalcemia can occur as a consequence ofexcess dietary vitamin D as well as a supply of cal-cium and either direct sunlight and/or artificialUVB radiation. The excess calcium may be


Figure 3Dante is a female ofunknown age who cameto me with very littlehistory. Her bones were sohopelessly demineralizedthat she could virtually betied into a pretzel. She wasunable to lift her body andcould only move bywriggling across a flatsurface. The mostastonishing thing aboutDante is that she hassurvived. After two years,she has shown consider-able improvement in skintone and color, muscletone and mobility. Herfront and back limbsremain bowed and herthighs are thickened byfibrous metaplasia. She isentirely mobile, even ableto climb, despite consider-able nerve damage to allof her digits. Photographby Carole Saucier

deposited in the bone (Figure 5) as well as in thesoft tissues of the body. If the heart muscle isaffected, sudden death may occur.

ReferencesFrye, F. and W. Townsend. 1993. Iguanas: A Guide to Their Biology

and Captive Care. Krieger Publishing Co., Malabar, Florida.

Guyton, A.C. 1981. Textbook of Medical Physiology. W.B. SaundersCo., Philadelphia, Pennsylvania.

Mader, D. 1996. Reptile Medicine and Surgery. W.B. Saunders Co.,Philadelphia, Pennsylvania.

All of the animals featured in this article are permanent residentsof the CT Iguana Sanctuary. Special thanks to Dr. Larry Linnetz ofthe Chippen’s Hill Veterinary Hospital for reviewing this article.


Figure 5Kahlo is an 8 year-oldfemale green iguana whopresented to a veterinaryclinic at age 6 with acalcium deficiency (5.0mg/dl, normal range is 7.5– 10 mg/dl). Her ownerswere advised to give hercrushed Tums as a calciumsupplement. She wasgiven Tums twice a day fortwo years and hervertebral column, from theneck to the base of the tailis fused. X-rays showsubstantial thickening ofthe bone throughout thetrunk as well as in the longbones of the rear limbs.There is no apparent softtissue involvement.Photograph by CaroleSaucier

Figure 4Hemingway was purchased from a pet shop for $6.99,selected from a large group of virtually identical hatchlingsbecause she had 23 toes. One rear foot has one extra toe andthe other has two partially formed feet, one with three toesand one with four. Despite being raised with the same diet andlighting provided for other animals who have grown to fullsize, after 5-1/2 years, Hemingway is only 15 cm SVL andweighs a little over 1 kg. Last year she produced a clutch of 18eggs which were about 1/3 the size of normal green iguanaeggs. She has a healed vertebral fracture and both her frontlegs have been broken at various times. Her lower jaw is alsoslightly splayed but she has no other skeletal deformities.Photograph by Carole Saucier

The pictures on the following pages are ofOsiris, the green iguana adopted by thestudents of Ms. Joyce St. Germaine’sGrade 7 Morning Meeting group at King

Philip Middle School in West Hartford,Connecticut.

Osiris had spent the first eight or nine years ofhis life languishing, like all too many “Iggys,” in a55-gallon aquarium with a marginal diet and inad-equate lighting. Both calls came in to theConnecticut Iguana Sanctuary at roughly the sametime, one from Iggy’s owners asking me to find ahome for him and the other from Ms. St. Germaineabout adopting an iguana for her classroom. Iggywas of “unknown sex” and allegedly fairly calm,although he hadn’t been handled much. I thoughthe’d make a good candidate for the school, but Ilost contact with his owners as the school year wascoming to a close. Iggy had apparently left home tospend the summer outdoors in the natural sunlight.He was actually in pretty fair physical conditionwhen he reappeared in the fall and his new place-ment was still available. Between a leftover cagefrom another adopted iguana, scavenged light fix-tures, and proper ultraviolet lights donated by alocal pet shop, he was well equipped to start school.

The habitat had all been set up and the studentshad diet and care sheets, iguana books, and backissues of the Iguana Times in hand before Iggy evenarrived. The students were tremendously excited butlistened intently when I came in to explain the basicsof green iguana care with the iguana perched on myshoulder. Shyly at first, and then with increasingassurance, the students approached to touch himand ask their questions. Beyond providing propercare for their new “classmate,” I asked that the kidsgive him a new name. The next day, when Ireturned with my Cuban iguana to give a talk aboutconservation, the board in the classroom was cov-ered with potential names. We had a bit of a closecall with “Fluffy,” but “Osiris,” after the EgyptianGod of the Underworld, received the most votes.

In the following weeks, I was pleased with theprogress reports I received on Osiris. The students

would apparently come into the classroom in themorning, prepare Osiris’ food and water and tidyhis habitat even before the teacher arrived. Hebecame quite the conversation piece, attractingstudents from all over the school, even if theyweren’t in any of Ms. St. Germaine’s art classes!The special education students would come and sitby his habitat at lunchtime just to enjoy his com-pany. A list had to be made up so that everyonewould have the opportunity to take him homeover weekends and holidays in his portable habi-tat. Ms. St. Germaine and Osiris brought out won-derful qualities in the students, who proved to beresponsible and well-informed caregivers.

I was delighted with the artwork that Osirishad inspired and impressed with the physicaldetails the students had depicted. I had talkedabout the differences between green iguanas androck iguanas in the genus Cyclura — and all of thepictures display lovely dorsal spines and gularshields.

Sadly, the next chapter of Osiris’ life was to beless happy, and provides a cautionary tale for allpotential iguana owners. For the first time in hislife, Osiris had a good diet and proper lighting; hehad gained some weight and become quite com-fortable with the school routine — and then camebreeding season. Again, most likely for the firsttime in his life, Osiris was healthy enough to expe-rience the hormonal surges of an adult maleiguana. He became irritable, occasionally evenmildly aggressive, and had a poor appetite. At onepoint, while the teacher was handling him, hejumped and bit her finger. Fortunately, she was notbadly hurt, but she was justifiably concerned forthe safety of her students. Together we decided totake the iguana out of the classroom before any-one else had a bad experience. So Osiris is back atthe Iguana Sanctuary awaiting another placement,preferably with a male as his primary caregiver orwith other iguanas with which he can socialize.Osiris’ legacy remains, for the most part, positivefrom the months he shared with the students ofKing Philip Middle School.

Green Iguana Invades West Hartford School

AJ Gutman



Spencer Glantz, Age 12

Michelle Fechtor,Age 12

Craig Wentworth, Age 12

Michelle Fechtor,Age 12

Allegra Levy, Age 11

GregoryDassonville, Age 12Ryan Reitz,

Age 12

Daniel W.Goldbaum-Shortell, Age 13

Victoria Skinner,Age 12

Josh Scheinblum, Age 12

Jonah Small,Age 13

Two species of spiny-tailed iguanas,Ctenosaura bakeri Stejneger, 1901 and C.similis (Gray, 1831) are native to theCaribbean island of Utila, which is situ-

ated off the coast of Honduras. These two speciesutilize different habitats and generally do not comeinto contact with one another. Ctenosaura similisoccurs in open rocky areas and C. bakeri, which isendemic to the island, is largely restricted to man-grove swamps. Due to the limited distribution andthe acute threat to the C. bakeri population fromover-hunting by locals, the “Schutz- undForschungsprojekt Utila-Leguan” (Conservationand Research Project Utila Iguana) was establishedin 1994. This project is sponsored jointly by theZoologischen Gesellschaft Frankfurt (FrankfurtZoological Society) and the SenckenbergischenNaturforschenden Gesellschaft (SenckenbergBiological Research Society) (Köhler 1998a). Acaptive breeding program for C. bakeri was estab-lished on Utila in April 1998 at the IguanaResearch and Breeding Station.

On 11 May 1998, a freshly killed femaleCtenosaura bakeri was brought to the Station bylocal people. The identity of the animal, which ispresently part of the collection at theForschungsinstitut and NaturmuseumSenckenberg (Senckenberg Research and NatureMuseum) (SMF 78870), is unambiguous.Necropsy, which was performed immediately,revealed that she contained ten eggs. Eight of thesewere placed in moist vermiculite in a Jäger incuba-tor at 28–32° C (temperature fluctuations due topower failure were limited). By 27 June, six eggshad spoiled, but the remaining two had increasedin volume. On 12 and 14 August, two apparentlyhealthy young animals hatched from these eggs(identity numbers 0510 and 0520). They mea-sured 55 mm from snout to vent (SVL), 181 mmtotal length (TL), and weighed 4 g. Both had twoclaws removed for permanent identification.

When they were placed in natural sunlight inan outdoor enclosure, their coloration obviouslydiffered from that of normal Ctenosaura bakerihatchlings. The latter are a nearly patternless grey-brown (Köhler 1998b), whereas these two speci-mens had vivid markings with clear crossbands andocelli and greenish coloration on the anterior dor-sal half, as is typical for C. similis hatchlings. On 11October, at the age of two months, the followingcoloration was noted using the “Naturalists ColorGuide” by Smithe (1975–1981) as a color refer-ence (code numbers of the colors in brackets):

Hatchling no. 0510 (217 mm TL; 70 mmSVL): dorsal surface of head and neck lime green(159); body sayal brown (223 C) with 5 dark(sepia 119) crossbands and light (cream color 59)ocelli; tail sayal brown (223 C) with dark (sepia119) crossbands. Hatchling no. 0520 (199 mmTL; 65 mm SVL) varied only in the shade of greenon the dorsal surface of the head and neck, whichwas registered as parrot green (160).

The scalation of the two hatchlings display aconfiguration that appears to be intermediatebetween those of Ctenosaura bakeri and C. similis.Hatchling no. 0510 has, without exception, twocomplete rows of enlarged spiked scales betweenthe whorls (typical of C. similis) on the right sideof the tail, whereas only one row is present ininterspaces 9–12 on the left side (typical of C. bak-eri). Consequently, the halves of the whorls do notcorrespond to one another along the median, acondition that does not occur in typical C. similisor C. bakeri. Hatchling no. 0520 has two com-plete rows of enlarged scales between the whorls;however, in interspaces 3–13, the foremost row isgreatly reduced. Scales on the anterodorsal side ofthe upper thighs are enlarged but not spiky, there-fore intermediate between C. bakeri (in whichthese scales are enlarged and spiky) and C. similis(in which these scales are not enlarged). Also, thenumber of dorsal spines, at 59 (no. 0510) and 64

Natural Hybridization between Ctenosaura bakeri and Ctenosaura similis

on Utila, Honduras*Gunther Köhler and Elke Blinn


(no. 1520), is intermediate between the values forC. bakeri (40–53, mean 44.6) and C. similis(61–96, mean 76.6) (data from Köhler 1995a, b).

An investigation of phylogenetic systematicsbased on morphological and genetic (RAPD fin-gerprinting) characters (Köhler 1995a) has shownthat the thirteen species of the genus Ctenosauracan be divided into three monophyletic groups at

the subgeneric level. Ctenosaura bakeri and C. sim-ilis belong to different subgroups. Ctenosaura bak-eri is more closely related to C. melanosterna, C.oedirhina, and C. palearis, whereas C. similis formsa tight cluster with C. acanthura, C. hemilopha,and C. pectinata. This is the first reportedhybridization between two species in the subfam-ily Iguaninae. Although we have been performing

field studies on Utila for manyyears, no further instances ofhybridization between C. bakeriand C. similis are known to haveoccurred. From this, we assumethat crosses between these speciesare rare, due primarily to theirobvious ecological separation.Both hybrids will be raised at theStation in order to determine ifthey are fertile.

Literature CitedKöhler, G. 1995a. Zur Systematik und

Ökologie der Schwarzleguan (GattungCtenosaura). Diss., Univ. Frankfurt,Frankfurt a/M.

Köhler, G. 1995b. Freilanduntersuchungenzur Morphologie und Ökologie vonCtenosaura bakeri und C. oedirhina aufden Islas de la Bahia, Honduras, mitBemerkungen zur Schutzproblematik.Salamandra 31:93–106.

Köhler, G. 1998a. Das Schutz- undForschungsprojekt Utila Schwarzleguan.Natur und Museum 128:44-49.

Köhler, G. 1998b. Schutz- undForschungsproject Utila Schwarzleguan:Die Nachzucht von Ctenosaura bakeriStejneger, 1901 im ex-situ-Zuchtprogram. Salamandra 34:227–238.

Smithe, F.B. 1975–1981. Naturalist’s ColorGuide. Part 1. Color Guide. 182 ColorSwatches. Amer. Mus. Nat. Hist., NewYork.

* Reprinted with permission from Salamandra36, 2000. Original title: NatürlicheBastardierung zwischen Ctenosaura bakeriund Ctenosaura similis auf Utila, Honduras.Translation by AJ Gutman.


Hybrid between C. bakeri and C. similis at two months of age.Photograph by Gunther Köhler

Wildlife Smug-glers Arrested inGrand CaymanGerman nationals JoachimSchmidt, Harald Endig, andJürgen Geisler were detainedby Customs officials afterchecking in at GrandCayman’s internationalairport on 30th January2001. They were attempt-ing to depart for Berlin viaMiami with luggage packedwith live reptiles, plants,and other wildlife from the Bahamas and GrandCayman. Schmidt pre-sented a local exportpermit, which was appar-ently forged, to an official.

The three came undersuspicion when theNational Trust for theCayman Islands was alertedto Schmidt’s presence onthe island by a localresident who rememberedconcern over collecting bythe same individual in themid 1990’s. After inquiriesand further reports, theTrust learned that the threemen were posing asacademic researchers whilecollecting live reptiles inconsiderable numbers, buthad made no recentcontact with local officialsand had not been issuedwith export permits.

As the Trust, the Depart-ments of Environment andAgriculture, and localenforcement agencies beganto assess the situation, alocal security firm, Intelsec,agreed to donate servicesand placed the men undercontinuous surveillance.Evidence rapidly mountedthat this was an illegalanimal smuggling opera-tion. This assessment wasreinforced by informationprovided to the Trust byTraffic International onSchmidt’s record ofinternational trading inendangered species.

In a closely coordinatedoperation involvingIntelsec, the Trust,

Department of Agriculture,Customs, Immigration,Department of Environ-ment, and Civil Aviation,all possible routes for illegalexport were monitored andthe men were followeduntil they checked in at theairport. Bags seized byCustoms after they werechecked for internationaltransfer contained 930endemic Grand Caymananole lizards, 140 Curly-tailed lizards from theBahamas, and 112 Curly-tailed lizards from GrandCayman. In smallernumbers were otherreptiles and amphibians,marine life, terrestrialinvertebrates, and acollection of bromeliads,cacti, and ferns. CITESrestricted material includedfour endemic GroundBoas, and 13 endemicBanana Orchids, all fromGrand Cayman.

Communication with theBahamas Governmentrevealed that no exportpermit had been issued forthe reptiles collected in theBahamas, nor had anycorresponding importpermit for these animalsbeen issued in GrandCayman.

On 2nd February inSummary Court, Schmidt,Endig, and Geisler wereremanded in police custodyon a series of smugglingcharges. The three werereturned to court on 7thFebruary, when Schmidtalso was charged withforgery: several chargesunder environmentallegislation may soon beadded. A trial date isexpected to be set shortly.

Once photographed bypolice as legal evidence, thewildlife was identified anddocumented by the Trustworking with the Depart-ments of Environment andAgriculture. Zoologyprofessor Sandy Echter-nacht from the Universityof Tennessee was flown

down by the CaymanIslands government toassist in identification of thereptiles from the Bahamas.By the evening of 3rdFebruary, all the GrandCayman wildlife had beenreleased back to the wild.

For the full list of seizedwildlife, and photographsincluding the suspectspreparing lizard nooses whilein Grand Cayman, visit theTrust’s web site at www.cay-mannationaltrust.orgNational Trust for the

Cayman Islands17 Courts Road,P.O. Box 31116 SMB, Grand CaymanCayman Islands, West Indies

8th February, 2001

Cyclura cornutaand C. ricordii inthe DominicanRepublicThe Parque ZoologicóNacional of the DominicanRepublic, The ToledoZoological Gardens, andthe Indianapolis Zoo arecollaboratively coordinat-ing a conservation projectdesigned to collect baselinevalues for blood chemistryand vitamin D levels inwild populations ofRicord’s iguanas, Cycluraricordii, and Rhinocerosiguanas, Cyclura cornuta.Blood samples werecollected from thepopulations of both speciesat Parque Nacional IslaCabritos, an island in LagoEnriquillo near the Haitianborder. A comparablestudy is being done by theDepartamento de RecursosNaturales y Ambientales of Puerto Rico and theToledo Zoological Gardenson Isla Mona, Puerto Rico, for the Mona Islandiguana, Cyclura stejnegeri.

Both Cyclura cornuta andC. ricordii occupy theapproximately 24 km2 IslaCabritos, which sits in themiddle of the hypersalineLago Enriquillo. The

habitat is typical ofsubtropical dry Antilleanforest, with several speciesof cacti and trees such as Bursera, Consolea,Guaiacum, and Prosopis.

Cyclura cornuta are fareasier to snare than thewary C. ricordii, whichbolt for their burrows atthe first sight of a human.Fifteen C. cornuta andnine C. ricordii wereprocessed during a recenttrip. Animals were sexed,weighed, and measured. In addition to a generalhealth examination, fecalsamples were taken todetermine bacterial speciesinhabiting the lowergastrointestinal tract.Ed. Note: Please see The Cycluraof Parque Nacional in IguanaTimes 8(2).

Source: ISG Newsletter 4(1),Spring 2001

InternationalIguanaFoundationEstablishedFirst introduced at theIguana Specialist Group’sannual meeting in 2000, the International IguanaFoundation has now beenincorporated as a Texas not-for-profit corporation. TheBoard of Directors andseveral guests from zoologi-cal societies and wildlifeconservation organizationsacross the U.S. held theirinaugural meeting at the FortWorth Zoo in August 2001.

The IIF will effectivelyreplace the iguana conser-vation funds administeredthrough the Fort WorthZoo that have helpedoperate the recoveryprograms in both Jamaicaand Anegada (British VirginIslands). The Foundationwill provide a peer reviewprocess to evaluate fundingrequests from a greaterarray of iguana projects.With $42,000 in commit-ments from 13 organiza-tions, the Foundation will

I G U A N A N E W S B R I E F S42 Iguana Times Volume 9, Numbers 1 and 2

be in a position to encour-age long-term planning.Additional corporations andzoos in Europe, Australia,and North America havebeen identified as possiblecontributing members.

The IIF will take itsfunding priorities fromthose identified by theIUCN/SSC IguanaSpecialist Group. Themission of the IIF is topromote the conservationof all species of iguanas.Source: ISG Newsletter 4(2) Fall 2001

Morris AnimalFoundationGrant AwardedIn July 2001, the Fort WorthZoo received a $46,440grant from the MorrisAnimal Foundation entitled“Health Assessment of Free-Ranging and HeadstartedWest Indian Iguanas.”

This grant will providefunds to:

• Establish baseline healthparameters for five speciesof free-ranging iguanas.

• Perform health screeningon headstarted iguanasprior to release into thewild.

• Provide training andtechnology for localveterinarians and biologiststo carry on this work.

The proposal is a collabora-tive effort involving theFort Worth Zoo, theWildlife ConservationSociety, the San DiegoZoo, the Indianapolis Zoo,and the Toledo Zoo. Thegoal of the project is tocompile and publish acomparative physiologicaldatabase for use byresearchers on variousspecies of Cyclura.Rick HudsonFort Worth Zoo

Source: ISG Newsletter 4(2) Fall 2001

The Impact ofHurricane Floydon the Bahamianrock iguana,Cyclura rileyirileyiLess than 700 endemicCyclura rileyi rileyi remainin a small number oflocations on and near SanSalvador Island. The iguanapopulation continues todecline due to predation ofeggs and young by feralrats, competition with acactus-eating moth, andsmuggling. In September1999, Hurricane Floydlashed San Salvador with155 mph winds and asubstantial storm surge.Adult iguana populationswere generally unaffected,but much of the nestinghabitat on offshore cayswas swept away and mostof the 1999 hatchlingswere destroyed. Some ofthe cay populations mayhave been extirpated by thestorm. Long-term damageto vegetation due to soilloss should be monitoredand immediate nestinghabitat restoration needs tobe considered.R.L. Carter & W.K. HayesDepartment of Natural SciencesLoma Linda University


IguanadelicatissimaMartiniqueIguana delicatissimapopulations on the variousislands of Martiniqueremain healthy. Measureshave been taken to expandsome of the nesting siteswhere late-arriving femaleshad previously beenobserved excavating theeggs of other females.Populations of introducedIguana iguana are beingcontrolled to preventproblems with competitionand hybridization.

Petite Terre(Guadeloupe)A particularly intense dryseason in the area has hadan adverse affect on theIguana delicatissimapopulation. In one area,only two iguanas were seenalive where thirty adults areusually found. In anotherarea of one hectare that washistorically inhabited bytwenty or more iguanas, nolive animals were seen —but 25 carcasses werefound. The mortality maybe due to starvation oroverheating as all theanimals found dead weremummified. The trees wereleafless. In two and a halfhours, 321 dead adultiguanas were counted invarious vegetation types. Anestimated 2500 adults mayhave died during this dryspell. This does not includeanimals that could havedied at the beginning of thedrought and were eaten byhermit crabs. This estimatealso does not include thoseindividuals expected to diebecause they are unable towalk and feed. Such animalswere found hanging in thetrees, barely alive. The adultloss may therefore be ashigh as 4000 animals. Withsurviving adults weakenedand unable to lay normalclutches, the impact oneffective population sizemay be substantial.Unfortunately, no collect-ing permits were issued forthe dead individuals so thatstudies might be performed.

Saint-BarthélemyA small remnant populationof Iguana delicatissimasurvives. The dry season wasresponsible for the death ofseveral goats that competedwith iguanas for food.Michel BreuilParis Museum of Natural History


Booby CayUpdateThe first trip to Booby Cayin two years was plannedfor November 2001, toallow Joe Wasilewski, JohnBendon, and two otherinvestigators to continueresearch on the endemicCyclura carinata bartschi.Unfortunately, due to ahurricane, the mail boatdidn’t arrive on Mayaguanaand no gasoline wasavailable to power the smallboat needed to get to thecay. John Bendon stayed onMayaguana until the gasarrived and took a day-tripto Booby Cay to assessdamage from the wind andstorm surge. All appearedto be well and severalpreviously marked iguanaswere sighted.

Since the previous trip in1999, a letter has beenwritten by Mr. JamesKnowles of the Ministry ofAgriculture in Nassau to thepolice on Mayaguana askingthem to remove theremaining goats from thecay. The letter was deliveredand funds should be availableto assist the police. A returntrip by the Booby Cay teamis planned for March 2002and the researchers hope tofind the island free of goats.Annual trips are proposed inorder to continue the iguanacensus and ensure that thecay remains free of the goatsthat compete with iguanasfor the limited vegetation.An application for a grant tocover the cost of investiga-tions for three years is beingmade to the newly formedIIF (International IguanaFoundation). In addition,further funds should haveaccrued from the sale of T-shirts and artwork (by J.Bendon) to facilitate futureresearch.John Bendon

I G U A N A N E W S B R I E F S43Iguana TimesMarch/June 2002

Iguana Times Back IssuesVolume 2, Number 1 (January 1993):Fiji Banded Iguanas; Roatán Island Spiny-TailedIguana; Salmonellosis in Reptiles; The CaptiveHusbandry and Propagation of the Cuban RockIguana, Cyclura nubila: Part 4. Breeding; IguanaRescue Group Update; Lizard Letters; IguanaNewsbriefs; News of the Society. Not available at the present time. May be reprinted in the future.

Volume 2, Number 2 (May 1993):The Mating Behavior of Iguana iguana (Part 1); A Visit with Mao; News of the Society; AlbertSchwartz; Iguana Newsbriefs

Volume 2, Number 3 (August 1993):The Mating Behavior of Iguana iguana (Part 2); The Captive Husbandry and Propagation of theCuban rock iguana, Cyclura nubila: Part 5. RaisingHatchlings; Iguana Rescue Group Update; FirstBreeding of Rhinoceros Iguana, Cylcura cornuta,at the National Zoo; Iguana Newsbriefs

Volume 2, Number 4 (December 1993):Evolution of Nesting Patterns in Iguanine Lizards;Iguanas as Pets?; News of the Society; IguanaNewsbriefs

Volume 3, Number 1 (March 1994): Herpetoculture and Conservation; Smuggling...Bahamian Iguanas; Cesarean Section in a Cyclura;Full Spectrum Lighting; Respiratory Diseases iniguanas; Lizard Letters; Iguana Newsbriefs; Treasurer’s Report; First full-color cover!

Volume 3, Number 2 (June 1994): Muy Dificil: Ctenosaura similis and defensor; The Indianapolis Zoo's Cyclura Program; IISVegetation Studies on San Salvador; IguanaSmuggling, A Sign of the Times; Iguana Newsbriefs

Volume 3, Number 3 (October 1994):Adaptations to Herbivory in Iguanine Lizards;Herpetoculture Today: One Person's Thoughts;Ecology, Status, and Conservation of the Utila Spiny-tailed Iguana, Ctenosaura bakeri; Book Review:Iguanas: A Guide to Their Biology and Captive Care;Treasurer’s Report; Iguana Newsbriefs

Volume 3, Number 4 (December 1994): The Paleate Spiny-tailed Iguana, Ctenosaura palearisstejnegeri: Distribution and Life History; AReintroduction Program for the Iguanas ofGuantanamo; A Trip to Mona Island; Reflections on Mona Island;Just My Opinion: A Commentary on Zoos and thePrivate Sector; Lizard Letters; Iguana Newsbriefs

Volume 4, Number 1 (March 1995): DivingDragons of the Galapagos; Living with Tee Beau:Sharing Your Life and Home with a 26-year-old

Rhino Iguana; Salmonella; Lizard Letters; Treasurer’sReport; Iguana Newsbriefs

Volume 4, Number 2 (June 1995): Population Status and Conservation of the Endangered San Salvador Rock Iguana, Cyclura r.rileyi; Chuckwallas; Crisis in the Galapagos; Iguana Newsbriefs

Volume 4, Number 3 (September 1995):Green Iguanas: Emerald Gems of the Jungle; What do Wild Iguanas Eat?; Need a Home for anUnwanted Iguana?; John G. Shedd Aquarium StudiesPossible Decline of Exuma Rock Iguanas in Bahamas;Reptile-associated Salmonellosis: Selected Cases;Lizard Letters; Iguana Newsbriefs

Volume 4, Number 4 (December 1995): A Study of the Mona Rock Iguana, Cyclura cornutastejnegeri, Nesting Sites on Mona Island, Puerto Rico;IIS Conference in San Salvador: Review and Report;Lizard Letters; Iguana Newsbriefs

Volume 5, Number 1 (March 1996): Psychosocialization of the Green Iguana: How toBetter Handle Your Pet; Green Iguanas are SocialBeings; International Sweep Targets ReptileSmugglers; Illegal Trade in Reptiles: Traffic Protectedby Legal Void; Abstracts of Scientific Presentations from the 3rd Annual IIS Conference in San Salvador;Iguana: Survival of the Tastiest; Book Reviews; Lizard Letters; Iguana Newsbriefs

Volume 5, Number 2 (June 1996): Lost in Time: Galapagos Land Iguanas; NorthernExposure: C. c. cornuta; Greasy Lizard Stuff; Iguanas, Salmonella and Herpetoculture: A Conflictof Interest... and Conscience?; Swampa Goes toKindergarten to Help its Survival; Utila Iguana GetsHelping Hand from Foreign Friends; Lizard Letters;Treasurer’s Report; Iguana Newsbriefs

Volume 5, Number 3 (October 1996): Lost and Found: Hope for the Jamaican Iguana;Genetic Studies of the Jamaican Iguana; CycluraForest Habitat; IIS Cyclura Island Habitat ClassificationSystem; Iguana Newsbriefs; News of the Society

Volume 5, Number 4 (December 1996): Any Hope for Grand Cayman's Blue Iguana?;Scalation Renderings of Cyclura nubila lewisi and C. nubila; Surviving Atlantis: The Molecular Evolutionof the Galapagos Iguanas; Iguana Report from Japan;IIS Conservation Award: Edwin Duffus; Lizard Letters;News of the Society

Volume 6, Number 1 (Spring 1997):Mayaguana Blues: Booby Cay iguana; Utila IguanaConservation and Research Project; Video Reviews;Inside the Labyrinth: Why Iguana Numbers Do NotAdd Up; Mood Swings for Iguanas; Youth in Science:The Green Iguana in Captivity; Iguana Newsbriefs

Volume 6, Number 2 (Summer 1997): Status of the Sandy Cay Rock Iguana, Cyclura rileyicristata; Decline of the Sandy Cay Iguana; Encounterwith Mama Iguana: A meeting with Dr. DagmarWerner; A Delicate Situation: Iguana delicatissima atthe Jersey Wildlife Preservation Trust; Youth inScience: Pook’s Hill Green Iguana Project; LizardLetters; Iguana Newsbriefs; News of the Society

Volume 6, Number 3 (Fall 1997): Searchingfor Iguana delicatissima; An Overview on theEvolution of the Family Iguanidae; Cuban and Rhino Iguana Egg Laying; Lizard Letters; News of the Society

Volume 6, Number 4 (Winter 1997): The Stout Iguana of the British Virgin Islands: Cyclura pinguis; Moon Over Mayaguana: Return to Booby Cay; Lizard Letters; Iguana Newsbriefs;News of the Society

Volume 7, Number 1 (Spring 1998): Galapagos Land Iguanas: Surviving in Peril; Morphologic Characters of Herbivorous Lizards;Feeding Behavior of a Free-Ranging Iguana iguanain the Lower Florida Keys;

Volume 7, Number 2 (Summer 1998):Progress for Cyclura rileyi cristata; Booby Cay Update; Iguana Rescue: Spot, Socks, Stanley andE.T.; Meet the Board (I.I.S.); Obituary: Captain RonHarrod; Iguanas on the Web; Lizard Letters; Iguana Newsbriefs

Volume 7, Number 3 (Fall 1998): Vanishing Iguanas (Cyclura cychlura figginsi); Newsof the Society: Searching for the Gwaya-Maga inBelize; Update: It Takes Two to Tango on Booby Cay;Lizard Letters

Volume 7, Number 4 (Winter 1998): Galopagos Marine Iguanas: A Living Heritage; Can a Spiney-tailed Iguana Survive in L.A.?; Iguana Newsbriefs; Lizard Letters

Volume 8, Number 1 (Spring 2000): The Status of the Lesser Antillean Iguana on SintEustatius; Where is the Beast of Andros?; The BigIguana Contest; Lizard Letters; Iguana Newsbriefs

Volume 8, Number 2 (Summer 2000): The Cyclura of Parque Nacional: Isla Cabritos; Cyclura Behavior During a Hurricane; JamaicanIguana Recovery Program; A Nature Sanctuary on Utila; Iguana Newsbriefs

Volume 8, Number 3 (Fall 2000): North Andros Island Report; Care of Iguana Eggs;Ctenosaurs of West Hartford; In Caribbean,Endangered Iguanas Get Their Day; IguanaNewsbriefs

Volume 8, Number 4 (Winter 2000): Fijian Banded Iguanas at the San Diego Zoo; Andros Island Iguana Update: 2001; Update: Iguana delicatissima; The Queen of Andros; Iguana Nutrition; Iguana Newsbriefs

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I.I.S. Bookstore

As a service to our members, a limitednumber of publications will be distributedthrough the I.I.S. Bookstore. The follow-ing publications are now available:

Green Iguana, The Ultimate Owner’s Manual,by James W. Hatfield. 1996. $28.00 (includingpostage); $35.00 (nonmembers). Covers just abouteverything from birth to death of an iguana. 600+ pp.Limitied quantities.

The Green Iguana Manual, by Philippe de Vosjoli.1992. $7.00 (including postage); $8.75 (non-members).

Send check or money order (payable to International Iguana Society) to:

I.I.S. BookstoreP.O. Box 366188Bonita Springs, FL 34136

Phot

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esy

of J

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B. Cyclura nubila nubila

A.Mona Island

Iguana

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The NewestInternational Iguana Society

T-Shirt The second in a series featuring various species of your favorite iguanas !

C.

Send check or money order to:International Iguana Society, Inc.P.O. Box 366188Bonita Springs, FL 34136

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The second shirt in the series, (picturedhere) features the ever-popular green iguana, Iguana iguana.

Illustration by John Bendon.

Adult male Ctenosaura nolascensis from Isla San PedroNolasco on a rock overlooking the Gulf of California.Photograph: L. Lee Grismer

Documents

IT 9.1-2 B&W pages · 2017-01-15 · Iguana Times THE JOURNAL OF THE INTERNATIONAL IGUANA SOCIETY g T VOLUME 9, NUMBERS 1 AND 2 MARCH/JUNE 2002 $12.00 A large male spiny-tailed iguana,