IRCF Reptiles & AmphibiAns...IRCF ReptIles & AmphIbIAns • Vol 18, no 1 • mAR 2011 3 basking in the alligator snapping Turtle, Macrochelys temminckii (Testudines: chelydridae) John

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Hylarana tipanan was described in 2000, but the experts who describe biodi-versity are disappearing (see commentary on p. 39).

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this juvenile female Giant musk turtle (Staurotypus salvinii) was recently discovered in miami, miami-Dade County, Florida (see article on p. 55).

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melanistic sombrero Ameivas (Ameiva corvina) occur only on the small isolated island of sombrero, the northernmost of the lesser Antilles. this individual had just excavated a beetle grub (see travelogue on p. 42).

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texas horned lizards (Phrynosoma cornutum) are popular with texans who often bring them home. they regularly show up at the south plains Wildlife Rehabilitation Center in lubbock, texas (see article on p. 18).

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A distressed male Sphaerodactylus leucaster with the vertically elliptical pupil it shares with other nocturnal geckos (see article on p. 12).

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back cover: stuart brookerAn adult male saint lucia Iguana (Iguana cf. iguana) from near the nesting site at louvet estate on saint lucia’s northeastern coast. this geneti-cally unique population might be threatened by invasive non-native Green Iguanas (I. iguana) that are breeding in the dry and mesic forests around soufrière in the southwest of the island. the most pressing concern is potential hybridization between individuals in the two populations, although com-petition between them could also pose a problem if they came into contact. see the article on p. 24.

Front cover: barry brownteiid lizards (racerunners and whiptails) are typi-cally not territorial, although males will fight for access to females and for other reasons we do not fully understand. these two male Curaçao Whiptails (Cnemidophorus murinus murinus) were so intent on each other that they tolerated a very close approach by the photographer. look for an article by Gerard van buurt about the teiid liz-ards of Aruba, bonaire, Curaçao, and the nearby paraguaná peninsula of Venezuela in an upcom-ing issue of Reptiles & Amphibians.

IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 1tAbleoFContents IRCF REPTILES & AMPHIBIANS • VOL15, NO 4 • DEC 2008 189TABLE OF CONTENTS

T A B L E O F C O N T E N T S

F E A T U R E A R T I C L E S

Chasing Bullsnakes (Pituophis catenifer sayi) in Wisconsin: On the Road to Understanding the Ecology and Conservation of the Midwest’s Giant Serpent ...................... Joshua M. Kapfer 190

The Shared History of Treeboas (Corallus grenadensis) and Humans on Grenada: A Hypothetical Excursion ............................................................................................................................Robert W. Henderson 198

R E S E A R C H A R T I C L E S

The Texas Horned Lizard in Central and Western Texas ....................... Emily Henry, Jason Brewer, Krista Mougey, and Gad Perry 204 The Knight Anole (Anolis equestris) in Florida

.............................................Brian J. Camposano, Kenneth L. Krysko, Kevin M. Enge, Ellen M. Donlan, and Michael Granatosky 212

C O N S E R V A T I O N A L E R T

World’s Mammals in Crisis ............................................................................................................................................................. 220 More Than Mammals ...................................................................................................................................................................... 223 The “Dow Jones Index” of Biodiversity ........................................................................................................................................... 225

H U S B A N D R Y

Captive Care of the Central Netted Dragon ....................................................................................................... Shannon Plummer 226

P R O F I L E

Kraig Adler: A Lifetime Promoting Herpetology ................................................................................................ Michael L. Treglia 234

C O M M E N T A R Y

The Turtles Have Been Watching Me ........................................................................................................................ Eric Gangloff 238

B O O K R E V I E W

Threatened Amphibians of the World edited by S.N. Stuart, M. Hoffmann, J.S. Chanson, N.A. Cox, R. Berridge, P. Ramani, and B.E. Young .............................................................................................................. Robert Powell 243

CONSERVATION RESEARCH REPORTS: Summaries of Published Conservation Research Reports ................................. 245 NATURAL HISTORY RESEARCH REPORTS: Summaries of Published Reports on Natural History ................................. 247 NEWBRIEFS ...................................................................................................................................................................................... 248 EDITORIAL INFORMATION ..................................................................................................................................................... 251 FOCUS ON CONSERVATION: A Project You Can Support ............................................................................................... 252

Front Cover. Shannon Plummer.Totat et velleseque audant mo estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus aut dolor apicto invere pe dolum fugiatis maionsequat eumque moditia erere nonsedis ma sectiatur ma derrovitae voluptam, as quos accullabo.

Back Cover. Michael KernTotat et velleseque audant mo

estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus

aut dolor apicto invere pe dolum fugiatis maionsequat eumque

moditia erere nonsedis ma sectia-tur ma derrovitae voluptam, as

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REPTILES & AMPHIBIANSC O N S E R V AT I O N A N D N AT U R A L H I S T O R Y

T a b l e o f c o n T e n T s

F E A T U R E A R T I C L E S

baskingintheAlligatorsnappingturtle,Macrochelys temminckii(testudines:Chelydridae)................................................. ....................................................................................................... John L. Carr, Samuel R. Holcomb, and Mitchell J. Ray 2

sympatrybetweentwoWide-rangingsalamanderspecies............................................................................Jennifer Deitloff 6

notesonActivitypatternsofFivespeciesofSphaerodactylus(squamata:sphaerodactylidae)fromtheDominicanRepublic.............................................Daniel P. Scantlebury, Julienne Ng, Miguel Landestoy, Anthony Geneva, and Richard E. Glor 12

herpetofaunaAdmittedtothesouthplainsWildlifeRehabilitationCenter(lubbock,texas):Atwo-decadeperspective......................................................................................................E. Kathleen McGaughey, Mark Wallace, and Gad Perry 18

nativeandAlienIguanasonsaintlucia,WestIndies................................................ Matthew N. Morton and Ulrike Krauss 24

ReadingtheAshes:ArsonDecimatesatropicalWetland,butAllowsnewobservationsofaneotropicalmudturtle..................................................................................................................................... Jeremy E. Guinn and Palani Luger 34

C O M M E N T A R Y

themassextinctionofscientistsWhostudyspecies..................................................................................... Craig McClain 39

T R A v E L O g U E

sombrero:lizardsAmongtheRuins............................................................................................................ Elizabeth Wright 42

I N T R O D U C E D S P E C I E S

thetentacledsnake, Erpeton tentaculatumlacépède1800(homalopsidae),inFlorida........................................................ .........................................................................................................................Joshua D. Holbrook and Kenneth L. Krysko 52

CubanbrownAnoles(Anolissagrei)insaintlucia............................................. Matthew N. Morton and Christopher A. Cox 52

AFailed(?)IntroductionoflizardsandFrogsonst.Vincent.......................................................................... Amos Glasgow 54

thepacificCoastGiantmuskturtle,Staurotypus salviniiGray1864(Kinosternidae),anewnon-indigenousspeciesinFlorida......................................... Dustin C. Smith, Kenneth L. Krysko, Tracy A. Sorensen, and Maria N. Sider 55

v ConseRVAtIonReseARChRepoRts:summariesofpublishedReportsonConservation..................................................57

v nAtURAlhIstoRYReseARChRepoRts:summariesofpublishedReportsonnaturalhistory........................................57

v neWsbRIeFs..................................................................................................................................................................................58

v Advertisements...................................................................................................................................................................................61

v eDItoRIAlInFoRmAtIon.......................................................................................................................................................63

v FoCUsonConseRVAtIon:AprojectYouCansupport.......................................................................................................64

theGrandCaymanblue-fannedAnole(Anolis conspersus)istheonlynativeanoleonGrandCayman.however,introducedCubanbrownAnoles(A. sagrei)haveforcedA. conspersustousehigherperchesinopenhabitatswherebothspeciesoccur.

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AjuvenileMacrochelys temminckii(alsoillustratedinFig.2)baskingonafloatingrailroadtieat0727h.notetheleechoccupyingaspacebetweenthevertebralandrightlateralkeels.

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basking in the alligator snapping Turtle, Macrochelys temminckii (Testudines:

chelydridae)Johnl.Carr1,samuelR.holcomb1,2,andmitchellJ.Ray1,3

1Departmentofbiologyandmuseumofnaturalhistory,Universityoflouisianaatmonroemonroe,louisiana71209,UsA([email protected])

2louisianaDepartmentofWildlifeandFisheries,p.o.box98000,batonRouge,louisiana70898,UsA([email protected])33321bluebirdRidge,newbraunfels,texas78130,UsA([email protected])

photographsbytheseniorauthor.

macrochelys temminckii is the largest freshwater turtle in northAmerica,andisoftenconsideredoneofthemostaquaticspecies

aswell(pritchard1967).onlyadultfemalesareknowntoregularlyleavethewater,andthenonlytonest(ernstandbarbour1972,Zappalorti1976,pritchard1989).baskinginthisspecieswaslongthoughttobenon-existentorextremelyrare(brattstromandCollins1972,ernstandbarbour1972).Althoughsomeauthorshavesuggestedthataquaticbask-ingoccursinthisspecies(Zappalorti1976,howeyandDinkelacker2009),nopublishedreportsdescribesuchbehavior.however,severalpublicationsdescribeaerialbaskinginM. temminckii,buteachliteraturereportcon-sistsofasingleobservation(ewert1976,shelbyandJensen2002,Farretal.2005,selmanetal.2009,thomas2009,selmanandWillig2010).ofthesixreportedinstancesofbasking,onewasahatchling(42mm,shelbyandJensen2002),threewerejuveniles(~100–200mmCl;ewert1976,thomas2009,selmanandWillig2010),andtwowereofadultsizes(~350–400mm,Farretal2005;~400–500mm,selmanetal.2009).oneinstanceofbaskingwasinferredfromcapturingaspecimeninabaskingtrap(shelbyandJensen2002).Intheotherfivecasesinwhichbaskingwasactuallyobserved,threeoftheturtlesappearedtohavebeenbaskingforsometime,astheanimalsweredescribedasdry(ewert1976,Farretal.2005,thomas2009),butthemaximumreportedobservationperiodlastednolongerthanabout12min(Farretal.2005).Allsixobservationswereassociatedwithlotichabitats(creeksorrivers),withbaskingsubstratesincludingafallentreetrunk,snags,andriverbanks.Wherereported,turtleswererestingonthebaskingsubstrateinclinedatanglesestimatedat30–60°,withtheheadpointedupslope(ewert1976,Farretal.2005,selmanetal.2009).Inonecase,theanimal’srestinganglewasnotnoted,butthesnagtoascendtothebaskingspotwasnearlyvertical(90°)andinvolvedclimbingoveraninterveningobstacle(selmanandWillig2010).thefiverecordedtimesforbaskingbehaviorwerebetween1000and1700h. Duringthe2009fieldseason,weobservedtwoinstancesofaerialbaskingandoneofaquaticbasking,allinvolvingjuvenileM. temminckii.observationsweremadeatblackbayoulakenationalWildlifeRefuge,11.5kmnorthofmonroeinouachitaparish,louisiana.thecenterpieceoftherefugeisalargelake(~845ha),whichisabackswamp(Wang1950)thathasbeendammedbyarailroadline,andnowhasthewaterlevelcon-trolledwithastop-logstructure.Airtemperatureswereobtainedfromtheneareststationinouachitaparishfortheappropriatedateandtime(WeatherUnderground,http://www.wunderground.com). Whileradio-trackingonblackbayoulakeatapproximately1400hon9march2009,ajuvenileM.temminckii(carapacelength[Cl]=174.3mm,weight=1.04kg)wasobservedwithitsheadexposedabovethewater.theafternoonwassunny,withverylittlecloudcover.Uponcloserinspec-tion,theturtlewasfoundtobefloatingaboveabedofsubmergedaquatic

vegetationcomposedprimarilyofEgeria densa (brazilianWaterweed) andCeratophyllum demersum (Coontail).theheadandkeelsofthecarapacewereexposedabovethewater,whereitremainedforaperiodof~5min,withnoindicationthatitwasforaging.theturtlewascapturedbyhandandbroughttothelabforexamination.ItwasidentifiedbyapIttagasa“head-started”juvenilefromthe2004year-classthatwereleasedon12June2008.Watertemperatureatthetimeofcapturewas21°Candairtemperaturewasapproximately27°C.thisobservationmatcheswellthe

AllIGAtoRsnAppInGtURtles

Fig. 1.JuvenileMacrochelys temminckii(A)baskingontheendofarailroadtieintheshadeofaWaterelm(Planera aquatica).thesameindividual(b)immediatelyaftercapture.notethelargeleechattachedtothecarapace.


descriptionofaquaticbaskingbymollandlegler(1971).theydescribedaquaticbaskingasbehaviorinwhich“turtles...weremotionlessatthesur-facewithlimbsfullyextendedandpartofthecarapaceoutofthewater,”andwentontosaythatthemostcommonsitesforaquaticbaskingofpanamaniansliders(Trachemys venusta)“wereoverbedsof[asubmergedaquaticplant]ornearothermatsofvegetation.” Inthecourseofterrestrialnestingsurveysalongtherailroadcausewayinthenorthwesterncornerofthelake,weobservedajuvenile(Cl=190.2mm,weight=1.52kg)on20April2009at0900h.Itwasrestingonanoldrailroadtieamongriprapatthemarginofthelake,29cmabovegroundlevel.thewetturtlewassittingaskewonthebankendoftherailroadtiewithalargeleechontherightsideofitscarapace(Fig.1).therailroadtiewasinclinedat~12°intothewaterfromthebank.basedontheanimal’spositionandthewettrailitleftonthetie,theturtlehadtohaveclimbedthebankandthentheendoftherailroadtie(~80–90°)toreachitsloca-tionatopthehigh,out-of-waterend.theturtlewaspositionedinsuchawaythatitwasrestingatananglenearhorizontal.thelefthindlegwasdanglingintheair,buttheright-sidelimbswereextendedandincontactwiththesubstrate;theheadandneckwerealsoextendedandrestingonthewood.thedigitswerenotmaximallyabducted(spreadapart).thisturtlewasobservedforjustafewminutesbeforebeingcaptured.Whenapproached,theanimalquicklydoveintothewateronthelake-sideoftherailroadtie.Atthetimeofthisobservationconditionswereclearandsunny,althoughthebaskingsitewasfullyshadedbyasmallWaterelm(Planera aquatica).Airtemperatureatthetimeofcapturewasapproximately17°C. WefoundanotherbaskingM. temminckiiduringaterrestrialnestingsurveyalongthelakemarginon29April2009.At0720h,ajuvenile(cara-

pacelength~140mm)wasobservedonarailroadtiefloating~3mfromtheshoreline(Fig.2).Ascanbeseeninthephotograph,thetailextendsstraightbehindthebody,therighthindlegisminimallyextendedwithoutabducteddigits,therightforelimbismorefullyextended,andallareincontactwiththesubstrate.theheadandneckarefullyextendedandtheanimalappearsalert,possiblywiththeproximalportionoftheneckrestingonthesubstrate.theweatheredrailroadtiewashorizontal,providinga0°baskingsurface,andwaslocatedintheearlymorningshadows,notexposedtothesun.Aleechwasnotedonthecarapace,andtheturtlewasphoto-graphedandobserveduntil0730h,atwhichtimetheobserver(JlC)leftthesite.Atapproximately0807h,whentheobserverreturnedtothebask-ingsite,theindividualwasseeninthesameposition,andthecarapacewasdrier,indicatingthatbaskinghadcontinuedinthemeantime.Inaddition,theleechthathadbeenobservedonthecarapacemoveduntilitwasoutofsight(Fig.2insert),whichfitswiththeideathatthedryingofaturtledur-ingbaskingwouldtendtocauseleechestoreleasetheirhold(pope1939,Cagle1950).theemptypupariumandanewlyemergedadultdamselfly(odonata:Zygoptera)arealsovisibleinoneofourphotos,sotheturtlehadremainedinthesamepositionlongenoughtoserveasasubstrateforemer-gencebythisaquaticinsect.thefirsttolastobservationsofthisindividualspanned58min.Airtemperaturewasapproximately22°C. ourobservationsincludethefirstspecificinstanceofaquaticbask-ingreportedforthespecies,andtwoadditionalinstancesofaerialbask-ing.obbardandbrooks(1979)notedapreponderanceofaerialbaskinginChelydra,buttheincidenceofaquaticbaskingwasashighas16%ofallbaskingobservationsforradio-telemeteredindividualsintheirpopulation.thesearetheonlyobservationsofbaskingM. temminckiithatwehave

Fig. 2.AsecondjuvenileMacrochelys temminckiibaskingonafloatingrailroadtie.notethattheleechismovingacrossthevertebralkeelat0816h.Intheinsert,theleechisnolongervisibleonthecarapaceat0817h.

CARR,holComb,AnDRAY


madeduring15yearsoffieldworkatthissitebytheseniorauthor(1996–2010),soweareinclinedtoagreewiththenotionthatbaskingoccursonlyrarelyinthisspecies(ewert1976).obbardandbrooks(1979)notedamongthereasonsthatbaskingmighthavebeeninfrequentlyreportedinChelydra serpentinapriortotheirstudywasthatindividualsalmostalwaysbaskaloneonaparticularobject,asopposedtoemydids,whicharefre-quentlyfoundinlargenumbersonthesameobjectorevenstackedononeanother.AllreportsofaerialbaskingMacrochelysareofsingleindividuals,thusiftheanimalsarewary,anindividualcouldveryeasilyslipintothewaterunnoticed.Inaddition,withrespecttoM.temminckii,thedarkcol-orationofthecarapaceandsoftpartswouldtendtobewellcamouflagedonmostlogsorotherwoodystructures,andtheroughtextureoftheshellandsoftpartswouldtendtodisrupttheoutlineoftheturtle,bothofwhichwouldcomplicateseeinganindividualbeforeitslippedintothewater. AllreportsofaerialbaskinginM. temminckiiinvolvesingleindi-vidualsonabankorwoodsubstrate.InC. serpentina(itsclosestnorthAmericanrelative),obbardandbrooks(1979)reportedthat99%of233aerialbaskingobservationsinvolvedasingleturtleonabaskingobject.thepostureassumedbythetwoaerialbaskingM. temminckiithatweobservedwassimilartowhatewert(1976)describedforChelydra,inwhichthefeetremainincontactwiththebaskingobject,ratherthanbeingfullyextendedinmid-airwiththedigitsmaximallyabductedasiscommonlyseenwiththehindlimbsinemydids(Cagle1950,Auth1975).obbardandbrooks(1979)reportedseeinglimbsextendedwiththewebbingspreadinChelydra, buttheydidnotquantifytheincidenceofthebehavior;however,intheirphoto,thelefthindlimbisnotelevatedabovethesubstratewiththedigitsmaximallyabducted.obbardandbrooks(1979)alsonotedthatonlyabout5%oftheirChelydrabaskingobservationsinvolvedanon-shorebaskingsubstrate.WithrespecttoMacrochelys,fouroftheaerialbaskingobservationshaveinvolvedbaskingsubstratesonthebankoralogattachedtothebank,andfourinvolvedfree-floatingorfixedoffshoreobjects. Withatotalofninebaskingobservationsforthespeciesnow,com-parisonwithfactorsrelatedtotheappearanceofthebehaviormaystarttobemade.someexplanationsforbaskingareapplicableonlytoadultturtles.ofthenineknowninstancesofbaskinginM. temminckii,themajorityofindividualswerejuveniles(7of9,includingour3observations),soanyassociationwithmaturationofovarianfolliclesinfemalesorwarmingbybaskinginthespringtoenhancematingbehavior(obbardandbrooks1979)couldaccountfornomorethanasmallfractionofthereportedcases.Anumberofgeneralexplanationsforbaskingbehaviorareavailable,includingthermoregulation(boyer1965,mollandlegler1971,obbardandbrooks1979),vitamin-Dsynthesis(pritchardandGreenhood1968),dryingoftheskinandshellsurfacethatcouldreduceectoparasiteandepi-zoophyteload(pope1939,Cagle1950,boyer1965,shelbyandJensen2002),andasaresponsetoillnessorinjury(selmanandQualls2009).thecaseofaquaticbaskingwouldseemclearlyrelatedtothermoregula-tionsinceitwasexposedtofullmiddaysunlightandistheearliestspringdaterecordedforbaskingwhenwatertemperatureswererelativelylow.ourothertwoobservationsbothinvolvedturtlesintheshade,ewert’s(1976)observationwasunderalightlyovercastsky,andthomas(2009)reportedtheindividualhesawwasinpartialsunlight.Althoughmostindividualsthathavebeenobservedwereexposedtosunnyskiesasonewouldexpectofaturtlebaskingforthermoregulation,indirectultravioletlightassoci-atedwithvitamin-Dsynthesiscouldstillreachaturtleunderacloudyskyorintheshade,butmuchlesseffectivelythanexposuretodirectsunlight.bothoftheaerialbaskingindividualsweobservedhadlargeleechesonthecarapace,andforoneofthetwo,wewereabletodocumentthattheleechreleaseditsinitialholdandmovedovertimeastheturtle’sshelldried.thiswassimilartoobservationsofGraptemysbyselmanetal.(2008)and

selmanandQualls(2009);however,thomas(2009)noted100+leecheswerestillattachedtothejuvenilehecapturedwhilebasking.

AcknowledgmentsthisresearchwasperformedunderlouisianastatescientificCollectingpermitlnhp–09–059andU.s.FishandWildlifeservicespecialUsepermit42651–09–03.FundingwasprovidedbythelouisianaDepartmentofWildlifeandFisheriesandtheU.s.FishandWildlifeservice,DivisionofFederalAid,throughthestateWildlifeGrantsprogram.headstartingwasundertakenbythenatchitochesnationalFishhatchery.WewouldliketothankthestaffofblackbayoulakenationalWildlifeRefuge,andKarenKilpatrickatthenatchitochesnationalFishhatcheryfortheircoopera-tion.WethankD.ligonforhelpfulcommentsonthemanuscript.

Literature CitedAuth,D.l.1975.behavioralecologyofbasking in theYellow-belliedturtle,

Chrysemys scripta scripta (schoepff).Bulletin of the Florida State Museum, Biological Sciences20:1–45.

boyer,D.R.1965.ecologyofthebaskinghabitinturtles.Ecology46:99–118.

brattstrom,b.h.andR.Collins.1972.thermoregulation.International Turtle and Tortoise Society Journal6(5):15–19.

Cagle,F.R.1950.thelifehistoryofthesliderturtle,Pseudemys scripta troostii(holbrook).Ecological Monographs20:31–54.

ernst,C.h.andR.W.barbour.1972.Turtles of the United States.UniversitypressofKentucky,lexington.

ewert,m.A.1976.nests,nestingandaerialbaskingofMacroclemysundernatu-ralconditions,andcomparisonswithChelydra(testudines:Chelydridae).Herpetologica32:150–156.

Farr,W.,p.Crump,andJ.Caraviotis.2005.Macrochelys temminckii(Alligatorsnappingturtle).Aerialbasking.Herpetological Review36:168.

howey,C.A.F.ands.A.Dinkelacker.2009.habitatselectionoftheAlligatorsnappingturtle(Macrochelys temminckii)inArkansas.Journal of Herpetology43:589–596.

moll,e.o.andJ.m.legler.1971.thelifehistoryofaneotropicalsliderturtle,Pseudemys scripta(schoepff),inpanama.Bulletin of the Los Angeles County Museum of Natural History, Science (11):1–102.

obbard,m.e.andR.J.brooks.1979.FactorsaffectingbaskinginanorthernpopulationoftheCommonsnappingturtle,Chelydra serpentina.Canadian Journal of Zoology57:435–440.

pope,C.h.1939.Turtles of the United States and Canada.AlfredA.Knopf,newYork.

pritchard,p.C.h.1967.Living Turtles of the World.t.F.h.publications,JerseyCity,newJersey.

pritchard,p.C.h.1989.The Alligator Snapping Turtle: Biology and Conservation.milwaukeepublicmuseum,milwaukee,Wisconsin.

pritchard,p.C.h.andW.F.Greenhood.1968.thesunandtheturtle.International Turtle and Tortoise Society Journal2:20–25.

selman,W.,b.Drescher,andC.Qualls.2009.Macrochelys temminckii(Alligatorsnappingturtle).Adultbaskingbehavior.Herpetological Review40:79.

selman,W.andC.Qualls.2009.Graptemys flavimaculata(Yellow-blotchedmapturtle).baskingandparasiteremoval.Herpetological Review40:78–79.

selman,W.,D.strong,andC.Qualls.2008.Graptemys gibbonsi(pascagoulamapturtle).baskingandparasiteremoval.Herpetological Review39:216.

selman,W.andm.Willig.2010.Macrochelys temminckii (Alligatorsnappingturtle).Aerialbaskingandclimbingability.Herpetological Review41:486.

shelby,J.A.andJ.b.Jensen.2002.Macrochelys temminckii(Alligatorsnappingturtle).Aerialbasking.Herpetological Review33:304.

thomas,t.m.2009.Macrochelys temminckii(Alligatorsnappingturtle).Aerialbasking.Herpetological Review40:336.

Wang,K.1952.Geologyofouachitaparish.Geological Bulletin(28):126.

Zappalorti,R.t.1976.The Amateur Zoologist’s Guide to Turtles and Crocodilians.stackpolebooks,harrisburg,pennsylvania.

AllIGAtoRsnAppInGtURtles


Plethodon cinereushasaverylargegeographicrangespanningmostofthenortheasternUnitedstates.

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sympatry between Two Wide-ranging salamander species

JenniferDeitloff

Departmentofecology,evolution,andorganismalbiology,IowastateUniversity,Ames,Iowa50014(Currentaddress:Departmentofbiologicalsciences,AuburnUniversity,Auburn,Alabama36849;[email protected])

photographsbytheauthorexceptwhereindicated.

Abstract.—Understandinghowcloselyrelatedspeciescoexistremainsacentralproblemincommunityecology.habitatcharacteristics,pres-enceofpredatorsandcompetitors,andfrequencyofdisturbanceallaffectthegeographicdistributionofaspecies.Plethodon cinereusandP. electromorphus aretwocloselyrelatedandecologicallysimilarspeciesthathavealargegeographicoverlapintheirdistributions.Inthisstudy,Ihadtwomaingoals.First,Iwantedtodeterminehowcommonlysympatriclocationsoccurredbetweenthesetwospecies.toaddressthisgoal,Iconductedlandscape-scalesurveysinareasthatshouldcontainsympatriclocationsbasedontheirdistributions.mysecondgoalwastodetermineiftherelativenumberofindividualsofeachspecieswithinsympatriclocationswasstableovertimebymonitoringfoursympatriclocationsoverthreeyears.Inaddition,floodingoccurredduringoneyearofthestudy,impactingtwoofthefourmonitoredsympatriclocations;therefore,Iwantedtoexaminewhetherthisdisturbanceaffectedthesalamanderpopulationsatthoselocations.thisfloodingeventprovidedarareoppor-tunitytostudytheimpactoffloodingonterrestrialsalamanders,atopicaboutwhichlittleisknown.Ifoundthatsympatriclocationsoccurredlessfrequentlythanexpectedbychanceandthatsomesympatricsitesseemtoberelativelystablewhileatothers,therelativepopulationsizeofP. cinereuswasincreasing—andthatthisshiftmightbepositivelyaffectedbyflooding.WhilethisstudydemonstratedthatP. cinereusandP. electromorphus donotoccurinsympatryasfrequentlyasexpectedbychance,thecauseforthisresultisunknown.themostlikelyfactoristhatthetwospeciesexhibitsubtlehabitatdifferencesthathavenotyetbeendemonstrated.theinteractionsbetweenP. cinereusandP. electromorphusarecomplex,andthepatternsobservedherecouldhavebeengeneratedbyanynumberofmechanisms.Furtherresearchonthissystemcouldclarifysomeoftheseremainingquestions.

Understandinghowcloselyrelatedspeciescoexistremainsacentralproblemincommunityecology.theamountofoverlapbetweenthe

geographicrangesofcloselyrelatedspeciesisthoughttodepend,inpart,ontheextenttowhichsimilarspeciessharethesamehabitatrequirements.Closelyrelatedspeciestendtooverlapmorethanexpectedbychance,sug-gestingthatspecies’distributionsaremoreconstrainedbysharednicherequirementsandcommonancestrythanbycompetition(harveyandpagel1991,letcheretal.1994).Conversely,nichesimilaritymayleadtointensecompetitionandeventualexclusionresultinginrareco-occurrenceofthetwospecies(bowersandbrown1982).thedistributionofaspeciescouldbeaffectedbyanynumberofmechanismsthatcouldincludehabitatcharacteristicsandthepresenceofpredatorsorcompetitors.

Interspecificcompetitionandterritorialaggressionseemtobeprev-alentwithin thegenusPlethodon (e.g., Jaeger1970,nishikawa1985,Anthonyetal.1997,marshalletal.2004).Insomeinstances,interspecificcompetitionhaslimitedthegeographicrangesofcompetingspecies(Jaeger1970,hairston1980,Arifetal.2007)andhasledtoavarietyofevolu-tionaryconsequencesbetweensympatricspecies,suchascharacterdisplace-ment(AdamsandRohlf2000),competitiveexclusion(Jaeger1971,myersandAdams2008),andalpha-selection—intenseinterspecificaggression(nishikawa1987,Deitloffetal.2009).Further,theseeffectsappeartohaveshapedspeciesdistributionsatabroadgeographicscale,becausepatternsofcommunitycompositionareconsistentwithwhatispredictedfromcom-petitive-basedmodelsofcommunitydynamics(Adams2007).

WIDe-RAnGInGsAlAmAnDeRspeCIes

Fig. 1.AdultPlethodon cinereus. Fig. 2.AdultPlethodon electromorphus.

8 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 DeItloFF

Plethodon cinereus(p.6andfig.1)hasaverylargegeographicrangespanningmostofthenortheasternUnitedstates;and,whilethegeographicrangeofP. electromorphus(Fig.2)ismuchsmaller,thisspecieshasarela-tivelylargegeographicrangecomparedtomanyotherspeciesinthegenusPlethodon(petranka1998).Inohio,thegeographicoverlapoftherangesofthesetwospeciesextendsthroughoutmostofthestate;but,whenviewedatafinerscale,thegeographicoverlapismoreaccuratelydescribedasinter-digitation(seeDeitloffetal.2008,figs.1and2).Inotherwords,allopatriclocationsmaybemorecommonlyfoundthansympatriclocations.Assug-gestedinDeitloffetal.(2008),thisdifferencebetweenthecoarse-andfine-scaleviewsofthegeographicdistributionsinthesetwospeciescouldleadtodifferingconclusionsabouttheextentofsympatry.therefore,Iaddressedtwogoalsinthisstudy.First,Iexaminedtheextentofsympatriclocationsbyconductinglandscape-scalesurveys(ratherthanusingpastdistributionmaps).second,Imonitoredfoursympatricsitesoverthreeyearstodeter-minewhethertherelativepopulationsizeofeitherspecieschangedovertime(i.e.,whethertheconditionofsympatrywasstableortransient). Duringtheperiodwhenthefoursympatricsitesweremonitored,oneoftheselocationswasstronglyaffectedbyaflood,anotherlocationwasweaklyaffectedbythesameflood,andtwolocationswerephysicallyundisturbedduringthestudy.thisdisturbanceprovidedanaturalexperi-menttoaddresstherelativeimpactoffloodingonthesepopulationsovertime.Floodingisacommondisturbanceofsalamanderassemblagesinbothaquaticandterrestrialsystems;however,moststudieshavefocusedonaquaticandsemi-aquaticspecies.Insomeinstances,individualsthatsurvivefloodingcanexhibitindicationsofcompetitiverelease(petrankaandsih1986);however,otherspeciesrequirealongertimeperiodforrecovery

(swansonetal.1998).theimpactoffloodsonterrestrialsalamandersislargelyunknown.

Materials and MethodsSpecies Co-occurrence.—Iconductedlandscape-scalesurveysofspeciespresenceduringspring2007andfall2007.toconducttheselandscape-scalesurveys,IsearchedforlocationswithinthedistributionaloverlapofP. cinereusandP. electromorphus.Iidentifiedlocationsonlywithincoun-tieswherebothspecieshavebeenfound(sympatriccounties)accordingtodistributionmapsofohio(pfingstonandmatson2003).Ichose21sympatriccountiesfromnortheasternohio(fromallpossiblesympatriccounties)becausethisareacontainedaconcentrationoftownships(sub-divisionsofcounties)wherebothspecieshavebeenfound(pfingstonandmatson2003).Withineachofthese21sympatriccounties,Iidentifiedlocationsthatcouldcontainsalamandersbyusingtopographicmapstodeterminewhichareascontainedwoodedhabitat.Afterarrivingataloca-tion,iftheareawasindeedsuitablehabitat,Isearchedforsalamanders.Ateachoftheselocations,IrecordedthenumberofindividualsofP. cinereusand/orP. electromorphusatthatlocation.eachlocationwassearchedforaminimumof20minutestodeterminethepresenceofsalamanders.Ifanysalamanderswerefound,theareawassearchedexhaustively(i.e.,allcoverobjectswereoverturnedandtheboundariesofeachsiteweredelineatedbyrivers,roads,ornon-forestedprivateproperty).Iadaptedthedistribu-tionmapsofpfingsonandmatson(2003)usingtheselocationsaswellasthelocationsmentionedbelow(sitesA,b,C,andD),whichalsowerelocatedwithinsympatriccounties(mapisdescribedinresults).locationswhereIfoundnosalamandersmighthavecontainedsalamanders;however,theseareaswerenotincludedintheanalysisdescribedbelowandtherefore

Fig. 3.siteA,wherefloodingsweptmostofthecoverobjectsfromtheslopesoftheravineintothestream.


didnotinfluenceresults.Iamconfidentthatlocationsdescribedasbeingallopatricwereindeedallopatricbecause:(1)eachlocationwassearchedexhaustively,and(2)whensympatriclocationswereidentified,minimalsearcheffortfoundbothspecies(searchtimeaslowas5minandtwototalsalamandersconfirmedthepresenceofbothspecies).Also,whenanaly-seswerecalculatedexcludingsiteswithfewerthanfivetotalsalamanders,resultswerethesame.notethatIfocusedonlyoncountieswherebothspecieshadbeendocumented,and,thus,hadthebestchanceoffindingsympatriclocationsiftheyexisted. totestthehypothesisthatP. cinereusandP. electromorphusco-occurlessfrequentlythanexpectedbychance,Iusedthecollectiondataobtainedfromthelandscape-scalesurveystoconstructapresence/absencematrix.thismatrixwasusedtodeterminewhetherco-occurrencebetweenP. cinereus andP. electromorphuswasbestdescribedassegregation,aggrega-tion,orrandom.IusedtheC-scoreindex(stoneandRoberts1990)asameasureofco-occurrence.thisindexiscalculatedas:

Cij=(ri–S)(rj–S)

wheresisthenumberofsharedsites(sitescontainingbothspecies)andriandrjarethenumberofoccurrences(rowtotals)forspeciesiandj.IftheC-scorewasgreaterthanexpectedbychance,speciessegregate;iftheC-scorewas smaller thanexpectedbychance, speciesaggregate. IusedecosimVersion7.0(Gotelliandentsminger2007)tocomparetheobservedC-scoretotheaverageC-scoregeneratedfrom10,000randomlyconstructedassemblages.Iusedamodelwhererowvalueswereretainedsoeachspeciesonlyoccurredasoftenasintheobservedmatrix(fixedsum)andwheresites(columns)areequallylikelytoberepresented(equiprob-

able),eliminatingobserveddifferencesinspeciesrichnessofsitesinthenullassemblages.

Flood.—Ilocatedfoursympatricsitesinohioduringspring2004.siteAwas~3mieoflisboninelkRuntownship,ColumbianaCounty(40°46’04”n,80°43’33”W);sitebwas10minoflisbon insalemtownship,ColumbianaCounty(40°49’48”n,80°49’19”W);siteCwas12mineofnewarkinperrytownship,lickingCounty(40°10’01”n,82°15’28”W);andsiteDwas9minofnewarkinWashingtontownship,lickingCounty(40°11’27”n,82°26’38”W).thesefoursitesweremoni-toredduringsevencollectingtripsfromspring2004tofall2007.springcollectingtripsin2004,2005,and2007occurredinearlymay,andfallcollectingtripsin2004,2005,2006,and2007occurredinlateseptemberandearlyoctober.thesefoursitesweresimilarincompositioninthattheyalloccurredintemperatedeciduousforestineast-centralohio.DuringlateAugustandseptember2004,severeandfrequentrainsresultedinflood-ingofsitesAandb.WhensiteAwasvisitedon25september2004,Iobservedthatmostoftherocks(usedascoverobjectsbysalamanders)andleaflitter(usedforforagingandrefugiaduringdryerconditions)hadbeensweptintoaravinelocatedwithinthesite(Fig.3).salamanderhabitatatsitebwasalsodisturbedbytheflood,butnotasseverely,andmanycoverobjectsremainedontheslope.sitesCandDwerenotfloodedduringthestudyperiod. Duringvisitstoeachsite,IcollectedadultP. cinereusandP. elec-tromorphusandrecordedthenumberofindividualsofeachspecies.Duringfall2004,sitesbandCweresearcheduntilapproximately25salamandersofeachspecieswerecollectedforotherresearch.therefore,thiscollectingseasonwasnotincludedintheanalyses


Fig. 4.photographofalocationwithallopatricPlethodon cinereus.thislocationwas8minWofnewarkinmcKeantownship,lickingCounty(40°08’06”n,82°32’05”W).


andisreportedonlybecausethiswasthecollectionthatoccurredimmediatelyafterthefloodandsiteAcontainedonlyonesalaman-der.Allcollectingwasconductedat0700–1100h,andallsalaman-derswerecapturedbyhand.Allsitesduringeachcollectionseasonweresearchedexhaustively(allcoverobjectswereoverturnedandtheboundariesofeachsiteweredelineatedbyrivers,roads,ornon-forestedprivateproperty).therefore,theareaateachsitethatwassearchedwasthesameforeachcollectionseason.Unfortunately,timespentcollectingsalamanderswasnotrecordedforthefirstcollectingseason(spring2004).threepeoplecollectedsalamandersduringthisseasonandeachsitewassearchedforapproximately1–2h(3–6col-lectingh/site).Forallothercollectingseasons,twopeoplecollectedsalamanders,andcollectioneffortpersitewassimilar(siteA:3heachcollectionseason;siteb:2–4h;siteC:2.5–4h;siteD:2h).thegoalofthispaperwastoestimatetherelativenumberofindividualsofP. cinereusandP. electromorphus,soactualpopulationsizeswerenotestimated.Instead,thenumberofindividualsofeachspeciesencoun-teredwasusedtocalculatetheproportionofP. cinereusateachsiteforeachseason,andthismeasureisreferredtoas“relativepopulationsize.”todetermineifthisproportionchangedsignificantlyovertime,Icomparedtheproportionfoundduringthesecondthroughseventhcollectingseasons(post-flood)totheproportionfoundduringthefirstcollectingseason(pre-flood)usingchi-squaretestswitha=0.05.

ResultsSpecies Co-occurrence.—Ifound20allopatriclocationsforP. cinereus (Fig.4),nineallopatriclocationsforP. electromorphus(twolocationswerewithinthesametownship),and12sympatriclocations;Ididnotfindanysalamandersin27townships.Fortheadaptedmap(Fig.5),eachtown-shipwascodedascontainingallopatricP. cinereus,allopatricP. electromor-

phus,bothspecies(sympatric), orneitherspecies.Plethodon cinereusandP. electromorphusco-occurredlessfrequentlythanwasexpectedbychance(C-score=180;p=0.0008).therefore,thedistributionsofP. cinereusandP. electromorphusinregardtothepresenceoftheotherspecieswouldbebestdescribedassegregation,asopposedtoaggregationorrandom.

Flood.—overthecourseofthisstudy,P. cinereuswasmorecommonthanP. electromorphus atallmonitoredsympatricsites.Atbothfloodsites(Aandb),IrecordedatemporalpatterninwhichP. cinereussignificantlyincreasedinrelativepopulationsizecomparedtoP. electromorphus(table1).Furthermore,infall2004(immediatelyaftertheflood),IfoundonlyoneadultP. cinereusandnoP. electromorphusatsiteA.Atthenon-floodedsites(CandD),relativepopulationsizesofP. cinereusandP. electromorphus didnotchangesignificantly(table1).morespecifically,atsiteC,anon-significantincreaseinP. cinereusoccurreduntilfall2006,whentherelativepopulationsizesweresimilartothefirstcollectingseason.WhilethechangewasnotsignificantatsiteD,noP. electromorphuswerefoundduringthefinalcollectingseason.

DiscussionthefirstgoalofthisstudywastoexaminetheextentofsympatrybetweenP. cinereusandP. electromorphusinohio,andIaddressedthisgoalbycon-ductinglandscape-scalesurveys.IfoundthatallopatricsitesofP. cinereusaremorecommonthansympatricsitesandallopatricsitesofP. electro-morphus,thatco-occurrencewaslessthanexpectedbychance,andthat,atsympatricsites,P. cinereusismoreabundant.thesecondgoalofthisstudywastomonitorsympatriclocationstodetermineiftheconditionofsym-patrywasstableovertime.Ifoundthatatonelocation,localextinctionofP. electromorphusmighthaveoccurred(siteD);atonelocation,therelativenumberofeachspeciesremainedapproximatelythesame(siteC);andattwolocations,therelativepopulationsizeofP. cinereusincreasedovertime(sitesAandb),butthesiteremainedsympatric. FloodingoccurredatthetwositeswhereP. cinereusincreased,sug-gestingthatfloodingcouldaffectrelativepopulationsizesofeachspecies.Furthermore,P. cinereusreturnedmorequicklyafterthefloodtositeAthanP. electromorphus.Atsiteb,wherefloodingoccurredbutdidnotdestroytheentireterrestrialhabitat,thecommunitychangedsignificantly,butthischangeoccurredoveralongertimeperiod.thismajordisturbanceappearedtoaffectthepopulationofP. electromorphus morethanthepopu-lationofP. cinereus.evidencesuggeststhatfloodinginfluencestherelativepopulationsizeofthesespeciesthroughoneofthree,non-mutuallyexclu-sivemechanisms:(1)enhancedcompetitiveexclusionofP. electromorphusbyP. cinereus,(2)achangeofenvironmentalcharacteristicsatthefloodedsites,makingthesesitesunsuitableforP. electromorphus,or(3)aslowerrateofreestablishmentforP. electromorphus relativetoP. cinereus. tomyknowledge,thisisthefirststudythatexaminedtheeffectsoffloodingonterrestrialsalamanders.naturaldisturbanceevents,suchasflooding,providerareopportunitiestostudypopulationdynamicsthatcannotbeunderstoodwithlaboratoryexperiments.Floodingcandisruptpopulationsofterrestrialsalamandersinmultipleways,includingdamag-ingthephysicalenvironmentbyremovingcoverobjectsthatarerequiredfortheirsurvival,decreasingpopulationsizesofpotentialcompetitorsandpredators,ortemporarilyremovingallindividuals. Inthissystem,localdistributionpatternsrevealthatcomplexinterac-tionsoccurbetweenthesetwospeciesandpossiblywiththeenvironmentorotherspecies(suchaspredators)notstudied.Competitionoftenplayssomeroleintheinteractionsofspecies,buttheimportanceofitseffectsinacom-munityarecontext-dependentandotherfactorsareimportantaswell(telloetal.2008),Forexample,disturbancehistory(hydeandsimons2001)andpredation(Connell1975)cansuppressdensitiestochronicallylowlevels,sothatsharedresourcesneverbecomelimiting—whichwouldpromoteco-occurrence.Unfortunately,theeffectofpredatorsoncommunitiesofPlethodonisunderstudiedandshouldbeexamined.Afullunderstandingof

Fig. 5.ResultsoflandscapesurveysofPlethodon cinereus andP. electromorphusinohioshowingsurveyedtownshipsoutlinedinblack(mapsadaptedfromohioDepartmentoftransportation2007).townshipswithsympatriclocationsarefilledwithblack,allopatricP. cinereuslocationswithlightgray,andallopatricP. electromorphuslocationswithdarkgray.townshipswhereIfoundnosalamandersarewhite.

DeItloFF


theroleofcompetition,predation,andenvironmentalconditionsinregulat-ingpopulationsrequiresbroadspatialandtemporalstudies(petrankaandsih1986)andwasbeyondthescopeofthisinvestigation.Futurestudiesshouldleadtoanunderstandingoftherelativeinfluenceofcompetition,predation,anddisturbanceinthecommunitydynamicsofterrestrialsalamanders.

AcknowledgementsIthankD.C.Adams,C.berns,K.barrett,J.o.Church,s.Graham,C.Guyer,V.Johnson,D.laurencio,e.m.myers,C.m.Romagosa,andD.A.steenforreadingdraftsofthiswork,andC.D.Anthony,J.o.Church,C.m.hickerson,e.m.myers,J.erickson,andG.Riceforhelpincollect-ingsalamanders.salamanderswerecollectedunderohioDepartmentofnaturalResources,DivisionofWildlifepermits122(2004),13(2005),144(2006),and299(2007),andwerehousedinaccordancewithIowa

stateUniversityCommitteeonAnimalCarepolicies(IACUC3-04-5618-D).thisresearchwassupportedbyaUsDAIFAFsmultidisciplinaryGraduateeducationtrainingGrant(2001-52100-11506).

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Table 1.numberofindividuals(n)ofeachspeciescollectedduringsam-plingperiodsandtheproportionoftotalindividualsthatwereP. cinereus(%pc).AtsiteAduringfall2006and2007,theproportionofP. cinereustoP. electromorphusissignificantlygreaterduringthefirstcollectionseasonindicatingtheimpactofthefloodonrelativepopulationsizes.Inaddition,atsitebwhereminorfloodingoccurred,theproportionofP. cinereustoP. electromorphusissignificantlygreaterduringfall2006and2007.p-values(p)fromchi-squaretestsareprovided.

P. P. electro- cinereus morphus (n) (n) %Pc P

Site A: Flood Site

spring2004 34 8 0.81 0—

Fall2004 1 0 0— 0—

Fall2005 32 4 0.89 0.23

Fall2006 33 1 0.97 0.02*

spring2007 15 1 0.94 0.19

Fall2007 40 3 0.93 0.04*

Site B: Weak flooding

spring2004 23 9 0.72 0—

Fall2004** 25 22 0.53 0—

spring2005 27 5 0.84 0.12

Fall2006 58 2 0.97 <0.0001*

spring2007 22 5 0.81 0.27

Fall2007 17 1 0.94 0.03*

Site C: No flooding

spring2004 49 13 0.79 0—

Fall2004** 25 25 0.50 0—

spring2005 28 3 0.90 0.12

Fall2006 82 28 0.75 0.25

Site D: No flooding

spring2004 25 3 0.89 0—

Fall2004 8 1 0.89 0.97

spring2007 7 0 1.00 0.36

*IndicatessignificantdifferencesbetweenthepercentofPlethodon cinereusduringfirstcollectingseasonandsubsequentcollectingseasons.

**thenumberofsalamanderscollectedwasintentionallyapproximately25foreachspecies.thesecollectingseasonswereomittedfromanalyses.


maleSphaerodactylus thompsonifromalocalityofftheAlcoaRoad,northofCaboRojo.thisindividualisonalimestonerockthattypifiesthepreferredsubstrateofthisspeciesinthearea.

sCAntlebURYetAl.


DwarfgeckosofthegenusSphaerodactylus(“sphaeros”)areoneofthedominantcomponentsoftheWestIndianherpetofaunainterms

ofspeciesrichnessandsheerlocalabundance—88WestIndianspeciesandnumeroussubspeciesarepresentlyrecognized(hendersonandpowell2009)andsomepopulationsapproachdensitiesof60,000/ha(Roddaetal.2001).Indeed,theyaresuchaubiquitousfeatureofCaribbeanherpetologythatsphaerosandrummightwellbestaplesofCaribbeanislands.thesegeckosoccupythefullscopeofavailablehabitatsintheWestIndies—coastalregionsincludingmangrovethickets,cactus-coveredxericbadlands,andmontanecloudforests.

sphaerosaretinygeckos,andincludethesmallestamniotespecies(hedgesandthomas2001).overall,thegenusisconsideredtobediur-nallyactive,althoughnocturnalactivityisreportedforafewspecies.mostspeciesliveasecretive,semi-fossoriallifestylethat,coupledwiththeirmin-isculesize,rendersobservationchallenging(e.g.,hendersonandpowell2009).thisdifficultyhasnodoubtcontributedtothefactthatwecur-rentlyknownexttonothingabouttheirbehavior,ecology,distributions,andfunctionalmorphology.thisdearthofknowledge,however,makessphaerosveryrewardinganimalstostudy.Almosteverydayoffieldworkspentonthesegeckosgeneratesnewdiscoveries.Inthispaper,wediscuss

notes on activity Patterns of five species of Sphaerodactylus (squamata: sphaerodactylidae)

from the Dominican RepublicDanielp.scantlebury1,Julienneng1,miguellandestoy2,AnthonyJ.Geneva1,andRicharde.Glor1

1Departmentofbiology,UniversityofRochester,Rochester,newYork14627-0211,UsA([email protected])2ministeriodemedioAmbienteyRecursosnaturales,santoDomingo,DominicanRepublic;sociedadornitologicadelahispaniola,santoDomingo,

DominicanRepublic([email protected])

photographsbytheseniorauthorexceptwherenoted.

SPhaerodaCtyluSIntheDomInICAnRepUblIC

maleSphaerodactylus ladaefromparquenacionalFrancisCaamaño.

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esto

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somerecentobservationsofnocturnalorcrepuscularactivitypatternsinfivepoorlyknownspeciesfromtheDominicanRepublic. nocturnalityistheancestrallifestyleforgeckos,althoughdiurnal-ityhasevolvedmultipletimes.Infact,thefamilysphaerodactylidaecon-tainsamixtureofdiurnalandnocturnalspecies.thegeneraAristelliger,Saurodactylus,Euleptes, andTeratoscincus arepredominatelynocturnal(althoughsomespeciesofAristelligerareknowntobaskonexposedsur-facesduringtheday;hendersonandpowell2009).everyothergenusinthefamilyislargelydiurnal,althoughnocturnalorcrepuscularspeciesareknownfromGonatodes,Pristurus,andSphaerodactylus.sphaeroswithrecordednocturnalactivitypatternsincludeS. inaguae,fromxericcoastalareas in thebahamas,S. roosevelti, fromxeric forestsof southwesternpuertoRico,S. savageifromthexericsoutheasterncoastoftheDominicanRepublic,andS. sputatorfromthenorthernlesserAntilles(referencesinschwartzandhenderson1991,hendersonandpowell2009). Inspectionofwherethesespeciesfallwithinpublishedphylogenies(hass1991,Gambleetal.2008)suggeststhatnocturnalityhasevolvedmultipletimeswithinSphaerodactylus.however,thedifferencesbetweendiurnalandnocturnallifestylesmaybeexaggeratedbyhumanperception.becausesphaerosarelargelysemi-fossorial,theymaybe“activeinaper-petualtwilight”(hendersonandpowell2009)regardlessofthetimeofday.therefore,evendiurnallyactivesphaerosmightpossesstheocularmorphol-ogymoretypicalofnocturnallizards.Inanycase,solittleisknownaboutthesegeckosthatabasicunderstandingofactivitypatternsiscriticalfor

accuratesurveywork.Indeed,formostspecies,nopublisheddataaddressactivitypatterns.

Sphaerodactylus leucaster and S. thompsoniSphaerodactylus leucaster andS. thompsoniarepoorlyknownxerophilicgeckosfromhispaniola.DuringrecentresearchtripstotheDominicanRepublic,wecollectedindividualsofthesespeciesunderdeadAgaveplantsandrocks,respectively,duringtheday.shortlyaftercollectingtheseani-mals,wenotedthepresenceofverticallyellipticalpupils,whichstandsincontrasttotheroundpupilcharacteristicofmostspeciesinthegenus,andismoretypicalofnocturnalgeckos.Consideringtheoppressiveheatofthehabitat(exposedsurfacetemperaturesoftensurpass60°Cduringtheheatoftheday),wedecidedtosurveyhabitatsatnighttoseeifthesegeckoswereactive,andperhapsmoreabundantthandaytimesurveysimplied. onsuccessivenights,weobservedmultipleanimalsofbothspeciesmovingaroundonexposedsurfaces.ForS. leucaster,weobservedatotalof20individuals(alladultsorsub-adults)overthecourseoftwoseparatenights(21August2009,14January2010)onakarst-strewnhillsidesouthofmonteRío,Azuaprovince.Wefoundindividualsofbothsexesonthegroundoronthesidesofrocksrevealedbydirectencountersorreflectiveeye-shines.oncedisturbed,geckoswouldrunforthenearestcover,whichwaseitherdeadAgaveorkarsttalus.ForS. thompsoni,weobserved>30ani-malsduringmultiplenightsofsearching(August2009andJanuary2010)andatmultiplelocalitiesalongtheAlcoaRoadbetweenCaboRojoandlasmercedes.thegeckoswereonkarstrocksat2000–2300h,andincludedbothsexesandallageclasses.Animalswerefrequentlyfoundonverticalrockfaces,andwererarelyfarfromcrevices.Anumberofanimalsalsowerefoundbyflippingsmalltaluspiles.Insuchinstances,geckoswereoftenontheundersidesoftherocks.Wefoundmoreanimalsinthesepilesatnightthanwedidwithsimilareffortduringtheday.ourfieldobservationsarerecapitulatedforbothspeciesbycaptivespecimens;animalsemergeafterthelightsareturnedoffandremainactiveforseveralhours. WecollectedmicrohabitattemperaturedataforsomeindividualsofS. thompsonibyusinganinfraredtemperatureguntorecordthesurfacetemperatureatthesiteofencounter.theaveragesurfacetemperaturewas31.9±1.5°C(n=6).Wealsorecordedthetemperatureofactive,butundisturbed,tarantulas(Phormictopus cancerides),whichwehavewitnessedkillingSphaerodactylusladae.theaveragetarantulatemperaturewas29.1±0.8°C(n=3).Althoughthesamplesizeissmall,astudent’sttestindicatedthatthedifferenceinaveragetemperaturesofthesespecieswassignificant(t =2.97,df=7,p=0.02).theselimitedandadmittedlyverypreliminarydatasuggestthatinteractionsbetweentarantulasandgeckosmightbeminimizedbydifferentthermalpreferences.

sCAntlebURYetAl.

maleSphaerodactylus leucasterfromthekarsthillsofmonteRío,southofAzua.thislizardwasactiveatnight.

Juvenile femaleSphaerodactylus ladaecollectedatnight fromparquenacionalFrancisCaamaño.

FemaleSphaerodactylus cryphiusfromanoldrockquarrybetweenDuvergéandJimaní.thescaleisinmm.thisgeckowascollectedasanadultunderneathadeadAgaveduringtheafternooninAugust2008;sheisstillaliveasof27January2011.


Sphaerodactylus ladaeSphaerodactylus ladaeisknownonlyfromtheDominicanRepublic,whereitinhabitsthescrubbyfoothillsofthesierrademartínGarcíaandthesur-roundinglowlandstowardthecityofAzua.WehavecollectedthisspeciesfromnearRancheríaandatparquenacionalFrancisCaamaño.toourknowledge,thelattersiterepresentsthemostnorthernandeasternrecordsforthisspecies.Geckoswereabundantatbothsites.AtRanchería,wecol-lectedanimalsduringthedayfrombeneathdeadandrottingAgaveplants,completelyoblivioustowhatwewoulddiscoverlateratparquenacionalFrancisCaamaño.InJanuary2010,oneofus(ml)visitedtheparktopho-tographtheareaatsunsetandconductapreliminarysurveyoftheareaatnight.thistripresultedinthecollectionofthreeadultS. ladae,whichweremovingaround,fullyexposed,justaftersunset.WerevisitedthesiteoneyearlaterinJanuary2011at2200–2300handfoundfiveanimalsactivelymovingaroundinshallowleaflittercollectedonaboulder-strewnhillside.Allwerefemaleandincludedtwoadultsandthreejuveniles(Dpssexedthembasedonpatternandtheabsenceofanenlargedescutcheonpatch).Weobservedbutfailedtosecureanadditionalthreeanimals.thehabitatatthissiteconsistsofkarstlimestoneboulderswithashortscrubcanopy.theexposedgroundbetweenlargerocksconsistsofshallowleaflitteratopgravel.Weobservedallofthegeckoscompletelyexposedonthesepatchesofleaflitter. We also collected three S. leucaster at parque nacional FrancisCaamaño,wheretheyoccursympatricallywithS. ladae.oneadultfemaleandtwojuvenileswereactiveat2200–2300hinexposedpatchesofleaflitteradjacenttolargelimestonerocks.sympatrybetweenS. ladaeandS. leucasterisnoteworthybecausethesespeciesaresimilarinsize,activitypat-tern,andappearance(maleshaveapalegraybodywithastreaked,brightyellowhead;femalesareadullpurple-gray,withbolddorsolaterallines).

Sphaerodactylus cryphiusAfinalxerophilicspecieswehaveobservedwhileactiveatnightisS. cry-phius.thisobservation,however,islimitedtoasingleindividualobservedafter2200hcrawlinginleaflitternearthebaseofalimestonecliffatanoldrockquarrybetweenDuvergéandJimaní.Additionalworkisnecessarytoclarifyifthisisanisolatedincidentortypicalofthisspecies.

Sphaerodactylus cochranaeSphaerodactylus cochranaeisknownonlyfromthesouthernshoreofthebahíadesamaná.thisbeautifulandrarelyseenspecieshasbeencol-lectedbyonlyasmallhandfulofherpetologists.on11september2009at1800–1845h(justarounddusk),weobservedfourgeckoswithina50-mradiusonleaflitternearthecoastinparquenacionalloshaitises.threeofthesightingswerebetween1830and1845h,suggestingthattheywerejustbecomingactive.Alternatively,theanimalswereendingactivityandseekingroostingspots;however,thisseemsunlikely,consideringthatwespentmuchofthedayinsuitablehabitatanddidnotcollectanyindi-viduals.Additionally,previousinformalnighttimesurveysinloshaitisesconductedin1996–1998byoneofus(ReG)recoveredactiveS. cochranaeinandaroundbromeliads.

DiscussionoursuccessinfindingandcollectingS. ladae, S. leucaster, andS. thompsoni increasedatnight.Furthermore,wesubstantiallyreducedourimpactonthehabitatduringnocturnalsearchesbecausewedidnothavetodisturbdeadAgavesandrockpiles.ConsideringtheclosephylogeneticrelationshipofS. leucasterandS. thompsoni,wehypothesizethattherelatedspeciesS. asterulus, S. rhabdotus,andS. shrevei,whichalsoinhabitxerichabitats,are

SPhaerodaCtyluSIntheDomInICAnRepUblIC

FemaleSphaerodactylus cochranaeperchedonthetopleafofabromeliadthatiscommonatlocalitieswherethisspecieslives.thesegeckoshavebeenseencrawlingintheaxilsofthisplantatnight.

riC

ha

rd

e. G

Lor

16 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 sCAntlebURYetAl.

alsonocturnalandmightoccuringreaterabundancethandaytimesurveysimply.similarly,S. schuberti,thepresumedrelativeofS. ladaefromthesierradeneiba(thomasandhedges1998),mightbenocturnalaswell.Sphaerodactylus schubertiisonlyknownfromthetypelocality,butthisareaisverysimilartohabitatsinwhichwehavecollectedS. ladae.similarly,therangesofthesespeciesmightwellhavebeenunderestimated. ImaginingageckoassmallasS. cryphiussurvivinginbrutallyhotanddryhabitatsisdifficult.nocturnalbehaviorcouldconceivablyallowthis

speciestoforagewhensurfacetemperatureshavecooledsubstantiallyfromdaytimehighs.however,wemustnotethatweneverobservedS. plummeriactiveatnightduringsurveysforS. thompsoni.Sphaerodactylus plummeriisapproximatelythesamesizeasS. cryphiusandlivesinsimilarhabitatsonthesouthernpartofthebarahonapeninsula.however,wereadilycol-lectedS. plummeriduringthedayatthesamesitesatwhichwecollectedS. thompsonithenightbefore.Althoughmoreresearchisnecessary,thisishighlysuggestiveoftemporalnichepartitioning.

AfisheyeviewoftypicalhabitatofSphaerodactylus thompsoniandS. plummeri.notethehighpercentageofexposedsurfacesandxerophyticvegetation.

CoastalleaflitterinparquenacionalloshaitiseswherewecollectedSphaerodactylus cochranae.theorangeflaginthecenteroftheimagedenotestheexactplacewhereageckowasfirstseen.

AfisheyeviewofthehabitatwherewecollectedSphaerodactylus cryphiusandS. rhab-dotusduringtheday.thishabitat,withamixtureofscrubvegetation,deadAgave,andlimestonerocksistypicaloflocalitieswherewehavecollectedS. ladaeandS. leucaster.

IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 17SPhaerodaCtyluSIntheDomInICAnRepUblIC

nocturnalityhasevolvedmultipletimeswithinSphaerodactylus,andnoc-turnalsphaerosarelargelyrestrictedtohotanddryhabitats.Althoughamorerigorouscomparativeanalysisisnecessary(andforthcoming),wehypothesizethatnocturnalityisanadaptationtoxericconditionsasitwouldalmostcer-tainlyreduceevaporativewaterlossbyrestrictingactivitytocoolerperiods.suchtransitionsbetweenactivitypatternsmaybefacilitatedifsphaerosarealreadypre-adaptedtolow-lightconditionsbecauseoftheirsemi-fossorialnature.Futureworkshouldinvestigatetheocularmorphologyofspeciesinacomparativeframework,aswellasquantifyactivityacrossspeciesintermsofambientlightconditions—insteadofemphasizingtimeofday.theobserva-tionthatS. leucasterandS. thompsonipossessverticallyellipticalpupils,typical

ofmostnocturnalgeckos,whereasS. ladaehasroundpupils,suggeststhatmultiplepathscouldfacilitateseeinginthenight.

AcknowledgmentsWehumblythankthesociedadornitologicadelahispaniola,andinpar-ticularJorgebroccaforassistinguswithsecuringpermitsandfieldarrange-mentswhiletravelingintheDominicanRepublic.J.Yoelhernándezpro-videdinvaluableassistanceatparquenacionalFrancisCaamaño.Gabrielde lossantossantanaconfirmed the identityof the tarantulas foundalongsideS. thompsoni.WealsothanktheministeriodemedioAmbienteyRecursosnaturalesinsantoDomingoforprovidinguswithpermitsandassistanceinthefield.thequalityofthismanuscriptbenefitedfromtheinsightfulcommentsoftworeviewers.

Literature CitedGamble,t.,A.m.bauer,e.Greenbaum,andt.R.Jackman.2008.evidence

forGondwananvicarianceinanancientcladeofgeckolizards.Journal of Biogeography35:88–104.

hass,C.A.1991.evolutionandbiogeographyofWestIndianSphaerodactylus(sauria,Gekkonidae)–amolecularapproach.Journal of Zoology225:525–561.

hedges,s.b.andR.thomas.2001.Atthelowersizelimitinamniotevertebrates:AnewdiminutivelizardfromtheWestIndies.Caribbean Journal of Science37:168–173.

henderson,R.W.andR.powell.2009.Natural History of West Indian Reptiles and Amphibians.UniversitypressofFlorida,Gainesville.

Rodda,G.h.,G.perry,R.J.Rondeau,andJ.lazell.2001.thedensestterrestrialvertebrate.Journal of Tropical Ecology17:331–338.

schwartz,A.andR.W.henderson.1991.Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural History.UniversityofFloridapress,Gainesville.

thomas,R. ands.b.hedges.1998.Anewgecko from thesierradeneibaofhispaniola(squamata:Gekkonidae:Sphaerodactylus).Herpetologica54:333–336.

JuvenileSphaerodactylus schuberticollectedatthetypelocalitynearlaDescubiertainAugust2009.

maleSphaerodactylus plummericollectedatthetypelocalityalongtheAlcoaRoadinAugust2009.

18 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 mCGAUGheY,WAllACe,AnDpeRRY

Urbanboxturtles,suchasthisornateboxturtle(Terrapene ornata ornata)inlubbock,frequentlycrossroadsandsometimesgethitbycars.


Herpetofauna admitted to the south Plains Wildlife Rehabilitation center (lubbock, Texas):

a Two-decade Perspectivee.KathleenmcGaughey,markWallace,andGadperry

DepartmentofnaturalResourcemanagement,box42125,texastechUniversity,lubbock,texas79409-2125,UsA

AllphotographsbyGadperryexceptwhereindicated.

morehumansthaneverliveinurbanareas,creatingcomplexenviron-mentalchallengesincludinginteractionswithnativespeciesandthe

introductionofspeciesnotnativetotheregionorevencontinent(Vitouseketal.1997,oldenetal.2006,mitchelletal.2008,Adamsandlindsey2009).Wildliferehabilitationcentersofferavenueforanimalsthathavesufferedfromtheinevitableencountersthatoccurbetweenhumansandanimals,whetherwild(e.g.,animalsinjuredbycarsorpets)orcaptive(e.g.,escapedpets;Karesh1995).theexactnumberofcertifiedwildliferehabili-tatorsintheUnitedstatesisunknown(nationalWildlifeRehabilitatorsAssociation2008),butestimatedat500–1,000(southeasternoutdoors2009,theWildlifeRehabilitationInformationDirectory2008).

thesouthplainsWildlifeRehabilitationCenter(spWRC),locatedinlubbock,texasandservingalargeareainthesouthernplains,rehabili-tatesandreleasesinjured,sick,displaced,ororphanedwildlife(southplainsWildlifeRehabilitationCenter1988).eachyear,spWRCacceptsavarietyofanimals,manyofwhicharenon-nativetotheregion.theadmissionsrecordskeptbythecenterprovideauniquerecordoftheinteractionsofresidentswithreptilesandamphibians,avaluablerecordofthepropagulepressurecontributingtothegrowinginvasiveherpetofauna(e.g.,Kraus2009,powelletal.2011),andhowthesechangeovertime.herewereportthenumbersofnativeandnon-nativeamphibiansandreptilesadmittedtothespWRCovertwodecadesandanalyzebothtaxonomicandtemporalpatterns.ouraimistoassessthemagnitudeofhuman-herpetofaunalinter-actionsinthismid-sizedcity,whichwillhopefullyofferanexampleofwhatmanyotherurbancentersacrossthenationexperience.thesedatacanbeusedtobettermanagesuchinteractionsatacommunitylevelandassistinpreparingarealisticresponseplantonon-nativespeciesarrivals.

MethodsWeexaminedallspWRCadmissionrecordsfortheyears1991–2009.ForeachwerecordedspeciesidentifiedbyspWRCstaff(untilrecently,iden-tificationswereprovidedbynon-herpetologistsandcouldnotbeindepen-dentlyverifiedbyus),arrivaldate,reasontheanimalwasbroughtin,whereitwasfound,injuriessustained(ifany),andultimatedisposition(died,hadtobeeuthanized,orwasreleased).Wecategorizedeachrecordasbelongingtoaspeciesnativeornon-nativetotheregion.Recordsvariedindetail,andnotalldatawereavailableforeachindividualanimal.

soUthplAInsWIlDlIFeRehAbIlItAtIonCenteR

thesehatchlingornateboxturtleshatchedattherehabilitationcenterinlate2010.theywilllikelybereleasednextspringinanappropriatehabitatintown—thisspeciesiscommonandpopularinsidethelubbockcitylimits.

A Red-eared slider (Trachemys scripta elegans) and baby ornate box turtles(Terrapene ornata ornata;atleft)arekeptinsideduringthewinteratthesouthplainsWildlifeRehabilitationCenter.Duringthespringandsummer,thisareanormallyhousesmanymoreturtles.


ResultsAlmost700individualsbelongingtoatleast43identifiedspecieswereadmit-tedtothespWRCbetween1991and2009.themajorityofthese,626indi-viduals(616reptilesand10amphibiansbelongingtoatleast19species)werenative,andanother52(48reptilesand4amphibians)belongingtoatleast24specieswerenon-native(Fig.1).byfar,mostadmittedindividualswerenativechelonians,primarilyornateboxturtles(Terrapene ornata ornata;n=342)andRed-earedsliders(Trachemys scripta elegans,n=138).Althoughthelatterspeciesisnativetothebroadregion,allorsomeofthesemighthaveoriginatedinthepettrade.othernativeturtlesincludedCommonsnappingturtles(Chelydra serpentina,n=20)andYellowmudturtles(Kinosternon flavescens flavescens,n=15).themostcommonlizardwasthetexashornedlizard(Phrynosoma cornutum,n=36),with1–2individualseachofeasternCollaredlizard(Crotaphytus collaris),lesserearlesslizard(Holbrookia maculata maculata),andanunidentifiedskink(whichmayormaynotbenative;andthereforewasnotincludedwiththenon-natives).themostcommonsnakeswerebullsnakes(Pituophis catenifer sayi,n=15)andCheckeredGartersnakes(Thamnophis marcianus,n=13),with1–2indi-vidualseachofakingsnakespecies(Lampropeltissp.)(which,liketheskink,mayormaynotbenative),GreatplainsRatsnake(Elaphe emoryi),WesternDiamondback(Crotalus atrox),Westernhognose(Heterodon nasicus),andWesternRattlesnake(Crotalus viridis).Amongnativeamphibiansthemostcommonwasthebarredtigersalamander(Ambystoma tigrinum mavortium,n=5),withsmallernumbersofunidentifiedspadefoots,toads,and“frogs”(whichalsomayormaynotbenative)completingthelist. turtlesalsodominatedamongnon-natives,withCommonmuskturtles(Sternotherus odoratus,n=9),Desertboxturtles(Terrapene ornata luteola,n=6), andAfricanspurredtortoises (Centrochelys [formerlyGeochelone]sulcata,n=3)beingthemostcommon,followedbyoneortwoeachoftheDeserttortoise(Gopherus agassizii),texastortoise(Gopherus berlandieri), Red-footed tortoise (Chelonoidis carbonaria), Alligatorsnappingturtle(Macroclemys temminckii),horsfield’stortoise(Testudo horsfieldii),easternboxturtle(Terrapene carolina),sabinemapturtle(Graptemys ouachitensis sabinensis),andWesternspinysoftshell(Apalone spi-nifera hartwegi).Identifiednon-nativearrivalsalsoincluded1–3individualseachofunknowngeckos(presumednon-nativebecausenospeciesisnativetotheregion),GreenIguana(Iguana iguana),monitorlizard(Varanussp.),stripedplateaulizard(Sceloporus virgatus),boaConstrictor(Boa con-strictor),easternCornsnake(Pantherophis guttatus),texasIndigosnake(Drymarchon corais erebennus),plainsGartersnake(Thamnophis radix),bullfrog(Lithobates catesbeianus),Americantoad(Anaxyrus americanus),andWesternspadefoot(Spea hammondii).perhapsmostsurprisingwasthenumberofcrocodilians,mostofwhichwereAmericanAlligators(Alligator mississippiensis,n=5).herpetologicaladmissionsgraduallyincreasedduringthe1990sandremainedroughlystableduringthedecadethatfollowed(Fig.2).theproportionofnon-nativesrangedfrom0–30%oftotaladmissionsforagivenyear,averaging7.6%.turtlesdominatedadmissionsduringtheentirestudyperiod(Fig.3).theirproportionofallherpetologicaladmis-sionsrangedfrom70–100%(includingthreeyearsinwhichallrecordedadmissionswereturtles),averagingabout85%. oftheknownreasonsforadmittanceofnatives,216individualswerereported“injured”(themostcommoninjurybeing“hitbyacar”;othersincludedcrackedshell,impaledwithafishinghook,attackedbyadogorcat,anddisease);57werefoundinorneartheroad,yard,home,orpark;18weresurrenderedpets;andonewasbroughtinforinjuringahuman.Amongnon-natives,knownreasonsforadmittanceweregenerallysimilar:11werefoundintheyard,road,orhome;sevenwereinjured;andthreewereconfiscatedbyauthorities(texasparksandWildlifeorAnimalControl). oftheknowndispositionsfornatives,361individualswerereleased(locationusuallyunspecified),54wereeuthanized,23died,threewerereturnedtothefinder,andtwoweretransferredtoalocalpetshop.Unfortunately,dis-positionsofnon-nativesfollowedasimilarpattern,with21being“released,”seventransferredtoalocalsciencemuseumorazoo,threedead,andtwoeuthanized.

DiscussionnativereptilesandamphibiansmadeupthemajorityofallherpetofaunaenteringspWRCin1991–2009.overallnumbersincreasedduringthe1990s,presumablyaresultofthegreaterexposuretheCenterhasachieved

Fig. 3.RelativeimportanceofcheloniansinherpetologicaladmissionsbythesouthplainsWildlifeRehabilitationCenterin1991–2009.

mCGAUGheY,WAllACe,AnDpeRRY

Fig. 1.numbersofamphibiansandreptilesadmittedbythesouthplainsWildlifeRehabilitationCenterovertheentireperiod1991–2009.

Fig. 2.numbersofnativesandnon-nativesadmittedbythesouthplainsWildlifeRehabilitationCenterin1991–2009.


duringthisperiod,followingitsestablishmentin1988.mostanimalswerebroughtinbecauseofaninjuryand~20%eitherdiedorhadtobeeutha-nized.thisleftthespWRCwithasubstantialnumberofnow-healthyani-malsinneedofdisposition;mostofthesewerereleasedintownornearby.

percythepelicanisoneofmanyrehabilitatedanimalsatthecenter.

Achildexaminesastatueofaraptorintheareareservedforpresentationsforthegeneralpublic.educationisabigpartofthemissionofthecenter.

soUthplAInsWIlDlIFeRehAbIlItAtIonCenteR

Citizensdropoffanimalsinthisbuildingattheentrancetothecenter.theyareaskedtoprovideinformationaboutcollectioncircumstancesandencouragedtoleaveadonationtohelpdefraythecostsoftreatinganimals.

Duringthesummer,thispondservesashomeforseveralrehabilitatedturtles,mostlyRed-earedsliders.

Amuralofregionalscenery,somenativevegetation,andthecenter’slogowelcomevisitorsattheroadsideentrance.

22 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 mCGAUGheY,WAllACe,AnDpeRRY

non-nativespecies,mostofthemlikelyformerpets,average~8%ofannualadmissions.thisnumbermaybeagrossunder-estimate,becausemanyoftheRed-earedslidersreceivedbythecentermayhaveoriginatedinthepettradeandlikelyrepresentnon-localgenetics.evenignoringthis,thedataincludeseveralalarmingfeatures.First,thenumberofnon-nativespeciesbeingadmittedisslightlylargerthanthatofnatives.second,largeandpotentiallyaggressivespeciessuchascrocodiliansarecommon.third,atleastsomeofthem,suchasthemonitorlizard(asavannahmonitor,Varanus exanthematicus,capturedinabackyard;G.perry,unpubl.data),theRed-earedsliders,andthebullfrogs,arecapableormaybecapableofsurvivinginthelubbockregion.Fourth,ourdatarepresentafar-from-completelistingofsuchanimalsfoundinthecity.Forexample,alargealbinoburmesepython(Python bivittatus)hadbeencapturedbycityper-sonnelandnotdeliveredtothespWRC(G.perry,unpubl.data).Finally,thedispositionofmanyoftheseanimalsisfarfromsatisfactory.thepythonmentionedabovewas“released”intoacitypark(G.perry,unpubl.data),andnearlyallsurvivingnon-nativesweresimilarly“released”byspWRCstaff.ourdatathusshowbothintroductionpathwaysidentifiedbyKraus(2009),unintentional(escapesofpets)andintentional(poorly-considered“releases”).Forlubbock,theyalsosupportthecontentionofKraus(2009)andothersthatthepet-tradeiscurrentlytheprimarysourceofanimalsestablishinginvasivepopulations.Forexample,manyofthenon-nativetur-tleandtortoisespeciesadmittedtospWRCarethosedescribedbyCeballosandFitzgerald(2004)asbeingtradedintexasforpetsand/orfood. ornateboxturtlescomprisedabouthalfofthecheloniansadmittedtothespWRC.thespeciesiscommoninlubbockandoftenencoun-teredbyresidents,whotendtoviewitfavorably(sosa2009,sosaetal.2010).Unfortunately,theoutcomesfor“released”boxturtlesarerarelyhappy(Cook2004,sosa2009),apatternthatiscommonintranslocatedamphibiansandreptiles(Doddandsiegel1991,siegelandDodd2002,Germanoandbishop2008). otherchelonianswerealsoverycommonamongadmittedherpeto-fauna,perhapsbecausetothepublic,turtlesarethemostcharismaticandnon-threateningofallreptilesandamphibians.Consideringthefearthattheyofteninvoke(morrisandmorris1965),wewerepleasantlysurprisedthatquiteafewsnakesandlizardswerealsoadmitted. our study suggests that rehabilitation centers at other locationsmightalsobereceivingsubstantialnumbersofamphibiansandreptiles,anissuewehopethatfutureresearchwilladdress.Italsoidentifiestwoareasofconcern.First,overtheyears,spWRCstaffmembershavenotbeenproperlyeducatedaboutnon-nativespeciesissuesandtheconsequencesof“releasing”suchanimals.thisisunlikelytobelimitedtoadmissionsof

amphibiansandreptiles.thus,well-meaningpeoplehelpingaddressurbanhuman-wildlifeinteractionsmightbecontributingtofutureproblemswithinvasivespecies.Clearly,opportunitiesforimprovededucationexistatthis,andlikelymanyotherrehabilitationcenters.second,moresolutionsareneededfordispositionofrehabilitatednativeandnon-nativewildlifealike:thelattercertainlyshouldnotbe“released”andtherecordforoutcomesintheformerispoorlydocumentedandgenerallydiscouraging.Forexample,Rodríguezetal.(2010)reportedonthereasonsraptorswerebroughtintoarehabilitationcenterandstatedthatover1,000were“releasedintothewild,”butnotwhathappenedtothempost-release.Inoneofthefewstudiesthatlookedatpost-“release”survival,bennett(1992)reportedthat>90%ofGibbons(Hylobates muelleri)quicklydied.survivaloftranslocatedcarnivorescanalsobepoor(linnelletal.1997).successratesfortranslo-catedamphibiansandreptilesarealsodiscouraging,evenwheretheprocessismuchmorecarefullyconceivedthanthetypical“release”fromareha-bilitationcenter(Doddandsiegel1991,siegelandDodd2002,Germanoandbishop2008).boththelogicandtheethicsofinvestingconsiderableresourcesinnursingasickorinjuredanimalbacktohealth,onlytohaveitdieuponleavingthecenter,aresuspect.

AcknowledgementsWethankthesouthplainsWildlifeRehabilitationCenterforallowingusaccesstotheirwildlifeadmittancerecordsandprovidingtechnicalassis-tance.thisismsno.t-9-1206oftheCollegeofAgriculturalsciencesandnaturalResources,texastechUniversity.

ReferencesAdams,C.e.andK.J.lindsey.2009.Urban Wildlife Management.2nded.CRC

press,bocaRaton,Florida.

bennett,J.1992.Aglutofgibbonsinsarawak—Isrehabilitationtheanswer?Oryx26:157–164.

Ceballos,C.p.andl.A.Fitzgerald.2004.thetradeinnativeandexoticturtlesintexas.Wildlife Society Bulletin32:881–892.

Cook,R.p.2004.Dispersal,homerangeestablishment,survival,andreproductionoftranslocatedeasternboxturtles,Terrapene c. carolina.Applied Herpetology1:197–228.

Dodd,C.K.,Jr.andR.A.siegel.1991.Relocation,repatriation,andtransloca-tionofamphibiansandreptiles:Aretheyconservationstrategiesthatwork?Herpetologica47:336–350.

Germano,J.m.andp.J.bishop.2008.suitabilityofamphibiansandreptilesfortranslocation.Conservation Biology23:7–15.

Karesh,W.b.1995.Wildliferehabilitation:Additionalconsiderationsfordevelop-ingcountries.Journal of Zoo and Wildlife Medicine26:2–9.

thishorsfield’stortoise(Testudo horsfieldii),anAsianspeciescommoninthepettrade,reachedthecenterafteritwasfoundroaming.Itisnowkeptinanoutdoorenclosurebyoneoftheauthorsandisdoingjustfine,suggestingthatitcouldhavesurvivedinlubbock.

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thistortoise,initiallyidentifiedasasulcatatortoise(Centrochelyssulcata),nativetoAfrica,wasfoundroamingalongsideatexashighway.Itisoneofseveralsuchanimalstoreachthecenterinrecentyears.

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Kraus,F.2009.Alien Reptiles and Amphibians: A Scientific Compendium and Analysis.springerseriesinInvasionbiology4.springer,newYork.

linnell, J.D.C., R. Aanes, J.e. swenson, J. odden, and m.e. smith. 1997.translocationofcarnivoresasamethodformanagingproblemanimals:Areview.Biodiversity and Conservation6:1245–1257.

mitchell,J.C.,R.e.Jungbrown,andb.bartholomew(eds.).2008.Urban Herpetology.societyforthestudyofAmphibiansandReptiles,saltlakeCity,Utah.

morris,D.andR.morris.1965.Men and Snakes.mcGraw-hill,newYork.

national Wildlife Rehabilitators Association. 2008. Finding a Rehabilitator.<http://www.nwrawildlife.org/page.asp?ID=214>.

olden,J.D.,n.l.poff,andm.l.mcKinney.2006.Forecastingfaunalandflo-ralhomogenizationassociatedwithhumanpopulationgeographyinnorthAmerica.Biological Conservation127:261–271.

powell,R.,R.W.henderson,m.C.Farmer,m.breuil,A.C.echternacht,G.vanbuurt,C.m.Romagosa,andG.perry.2011.IntroducedamphibiansandreptilesintheGreaterCaribbean:patternsandconservationimplications,pp.63–143.In:A.hailey,b.s.Wilson,andJ.A.horrocks(eds.),Conservation of Caribbean Island Herpetofaunas.Volume1.brill,leiden,thenetherlands.

Rodríguez,b.,A.Rodríguez,F.siverio,andm.siverio.2010.Causesofraptoradmissionstoawildliferehabilitationcenterintenerife(CanaryIslands).Journal of Raptor Research44:30–39.

seigel,R.A.andC.K.Dodd,Jr.2002.translocationsofamphibians:provenman-agementmethodorexperimentaltechnique?Conservation Biology16:552–554.

sosa,J.A.2009.effectsofurbanizationonmovements,activity,andtranslocationsitefidelityofornateboxturtles(Terrapene ornata ornata)inthesouthernhighplainsoftexas.m.s.thesis,texastechUniversity,lubbock.

sosa,A.,o.Reyes,andG.perry.2010.turtlesinthedust:effectsofhands-onscientifictrainingonagroupofbehaviorallyat-riskstudents’knowledgeandempathy.Reptiles & Amphibians17:108–111.

southplainsWildlifeRehabilitationCenter.1988.AboutspWRC.<http://spwrc.org/index.php?option=com_content&task=view&id=17&Itemid=29>.

southeasternoutdoors.2009.WildlifeRehabilitatorsDirectory–UsA.<http://www.southeasternoutdoors.com/wildlife/rehabilitators/directory-us.html>.

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theWildlifeRehabilitationInformationDirectory.2008.howtolocateaWildlifeRehabilitator.<http://www.tc.umn.edu/~devo0028/contact.htm>.

someturtlesarekilledimmediatelywhentheymeetvehicles,butothersarelessseverelywoundedandarebroughttothecenterforrehabilitation.

texashornedlizards(Phrynosoma cornutum)suchasthisonetendtodisappearfromurbancentersbutcanpersistontheoutskirtsoftownandmaybetransportedbyresidentsreturningfromtrips.theyregularlyshowupatthecenter.

thiseasternCornsnake(Pantherophis guttatus),clearlyawell-fedcaptive-bredindi-vidual,wasfoundonthefrontporchofalubbockresidentandbroughttothecen-ter.Itisextremelytameandobviouslyapetthatwasreleasedorallowedtoescape.


saintluciaIguana(Iguanacf.iguana)on“lyennDous”(UmbrellaVine,Ipomoea tiliacea).

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native and alien Iguanas on saint lucia, West Indies

matthewn.morton1andUlrikeKrauss2

1DurrellWildlifeConservationtrust,lesAugrèsmanor,trinity,JerseyJe35bp,ChannelIslands,Greatbritain([email protected])2saintluciaForestryDepartment,ministryofAgriculture,lands,Forestry&Fisheries,GabrielCharlesForestryComplex,Union,

saintlucia,WestIndies([email protected])

Status of Native Saint Lucian Iguanas

thetaxonomyoftheGreenIguana(Iguana iguana)remainsunresolved.maloneandDavis(2004)conductedgeneticanalysesthat“imply

thatatleastthreecrypticspeciesmayexist[withinthetaxoncurrentlyrecognizedasI. iguana] undertheevolutionaryandphylogeneticspeciesconcepts (CentralAmerican,southAmerican, [and]southAmerican[Caribbean]+lesserAntillean).”theyalsoindicatedthattheirdatasup-portedatleasttworadiationsintothelesserAntilles,“firstontosaintluciaandmorerecentlyontosabaandmontserrat.”lazell(1973)dis-missedthecontentionthatI. iguanaisarecentintroduction(e.g.,byCaribpeoples;Underwood1962)totheeasternCaribbean,althoughI. iguanaonmartinique(notconsideredbylazell)wasintroducedinthe1960s(breuil2009).breuil(2002)alsorejectedlazell’sargumentthatI. iguanaisnativetoGuadeloupe,whereitoccurswiththeendemiclesserAntilleanIguana(I. delicatissima).powell(2004)andpowellandhenderson(2005)highlightedtheimportanceofinsularpopulationsclassifiedcurrentlyasI. iguanaandtherisksofassumingtheycompriseasinglespecies. thenativepopulationofI. iguanaonsaintluciaisrestrictedtotheisland’snortheasterncoast,andpossiblyhasbeenrestrictedtotheeasterncoastsincethe19thCentury(tyler1850)1.Aslongasthispopulationissmallandhasarestrictedrange,itremainsaconservationpriorityforthe

ForestryDepartmentoftheministryofAgriculture,lands,Forestry&Fisheries(mAlFF),thestatutorybodyresponsibleforterrestrialbiodiver-sityconservationonsaintlucia.Duringabiophysicalinventoryofsaintlucia’sforestresourcesin2009,thedecisionwasmadetoformallyrefertothepopulationonsaintluciaasIguanacf.iguana—i.e.,similarto,butnotconfirmedas,Iguana iguana(Daltry2009a,morton2009)—andtorefertothispopulationas“thesaintluciaIguana,”withoutimplyinganyresolutionoftheprevailingtaxonomicuncertainty. ConcernsaboutthesurvivalofthesaintluciaIguanawereexpressedbythesaintluciaForestryDepartment(slFD)andJ.Gilardiinthe1990s(Anonymous1998,bendon2003),initiatingworkinthe2000sbytheDurrellWildlifeConservationtrust(Durrell)andslFDtoevaluatethestatusofthepopulationandthethreatsfacingit(morton2007).morerecently,slFDandRareAnimalReliefeffort(RARe)builtuponearlierawareness-raisingefforts(bendon2003)bymeansofa“pride”campaign(narcisse2009,RARe2010).thiscampaignpromotedthesaintluciaIguanaasaflagshipspeciesfortheisland’sendangereddeciduousseasonalforesthabitat,andinparticularthedeciduousseasonalforestsofsaintlucia’snortheasterncorridor.boththepridecampaign’siguanamascotandthenortheasterncorridorweregiventhenameIyanola,aphoneticversionofIouanalao,theAmerindiannameforsaintlucia,meaning“thelandoftheiguana”(Jesse1960).Inthisrespect,theimportanceofthesaintluciaIguanatoconservationeffortsontheislandwouldremainunchangedregardlessofitsfuturetaxonomicstatus.Itissaintlucia’s

1lazell’s(1973)reportofiguanasonmariamajorislandoffthesoutherntipofsaintluciaisinerror(J.lazell,inlitt.,23.VI.2010).

saintluciaIguana(left)andaniguanafrommontserrat(right).notethedifferencesinscalationbetweenthesetwoWestIndianiguanapopulations.

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largestnativeterrestrialanimal,strikinginitsappearance,andaculturalsignifieroflongstanding.Assuch,itisavaluableflagshipspeciesthatrep-resentsagloballythreatenedhabitattypeintheinsularCaribbean(WWFandmcGinley2007),anareathatappearstobethelastremainingstrong-holdofanumberofspeciesandsubspeciesendemictosaintlucia,par-ticularlybirdsandreptiles(Daltry2009a,2009b;morton2009;toussaintetal.2009). IfsaintluciaweretoproduceitsownRedlist,thesaintluciaIguanawouldrequireaRedlistassessmentofCriticallyendangered(IUCn2001)atanationallevel.Ifthispopulationweretobeaccordedspecificorsubspecifictaxonomicstatus,however,itwouldqualifyforaglobalstatusofCriticallyendangered.Ineithercase,thisassessmentisbasedonIUCn’s(2001)criteriab1a,b(i,ii,iii):extentofoccurrenceestimatedtobelessthan100km2(boththeextentofoccurrenceandareaofoccupancyare<50km2);dataindicatingthatthepopulationisseverelyfragmentedorknowntoexistatonlyasinglelocation(onelocation—northeasternsaintlucia);andindicatorsshowingcontinuingdecline,observed,inferred,orprojected,inthe(i)extentofoccurrence,(ii)areaofoccupancy,and(iii)area,extent,and/orqualityofhabitat(effectsoftourismdevelopments,sand-mining,livestockgrazing,andotherdocu-mentedthreatsthatareknowntoreducethequalityandextentofsuitablehabitats).historicalpopulationbaselinesarenotavailableforthesaintluciaIguana,butthisassessmentinfersadeclineinthepopulationbasedonitsveryrestrictedrangeandtheidentificationofanumberofongoingpressuresandthreats,mostnotablythatofintroducedmammalianpreda-torsandthethreatofproposed,large-scaletouristdevelopmentonsaintlucia’snortheasterncoast(morton2007). thecurrentrangeofthesaintluciaIguanacorrespondstooneofthetwoareasontheislandfarthestfrompavedroads(i.e.,leastacces-sibletohumans,theircommensals,anddomesticatedanimals).theotherareaisamountainousregionofsaintlucia’shighestpeaks,fromwhichnoiguanasareknown(perhapsduetoalackofsuitablenestinghabitat).thisdistributionimpliesthathunting(whichstilloccursonasmallscale,primarilyforfood;bendon2003,morton2007)mighthavebeenoneimportanthistoricaldriveroftheinferreddeclineinthesaintluciapopu-lation.Italsounderlinesoneofthethreatsarisingfromproposeddevelop-mentonthenortheasterncoast,overandaboveconversionofhabitat:theimprovementofroadaccessandhencehumanaccessthat,ifnotregulatedinsomeform,canbeexpectedtoincreasehuntingpressureaswellasthedensityofnon-nativemammalianpredatorssuchasmongooses,dogs,cats,andrats.Improvedroadaccessandahigherhumanpopulationinthenortheastmightalsoincreasethelikelihoodthatalieniguanas,nowestab-lishingonsaintlucia,aretransportedintothisarea,eitherdeliberatelyaspetsorfoodorinadvertentlystowedawayingoods.

Introduction and Establishment of Alien Iguanas – Potential ImpactsthequestionofthetaxonomicstatusofthesaintluciaIguanahasagaincometotheforewiththeconfirmationin2008thatnon-saintlucianGreenIguanasarebreedinginthedryandmesicforestsaroundsoufrièreinthesouthwestoftheisland(morton2008,GlobalInvasivespeciesDatabase2010a).themostpressingconcernforbiodiversityconserva-tiononsaintluciaisthelikelihoodofhybridizationbetweennativeandalieniguanas,potentiallycompromisingtheuniquegeneticidentityandlocallyadaptedgenecomplexesof thesaintluciapopulation.Iguana iguana isknowntohybridizewithitscongener,I. delicatissima,oneasternCaribbeanislandstowhichithasbeenintroduced(e.g.,Dayandthorpe1996,breuiletal.2007). Competitionbetweennativeandalienpopulationsisanotherconcern(hendersonandpowell2009).IUCn’sGlobalInvasivespeciesDatabase(2010b)mootsthepossibilitythatcompetitionbetweenintroducedGreenIguanasandtheircongener,I. delicatissima,mightthreatenthosenative

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Iyanola,saintlucia’sprideCampaignmascotforthesaintluciaIguanaandthenamebestowedontheisland’snortheasterncorridor,itslastknownrefuge.

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thedistributionofthesaintluciaIguanaandapopulationofintroducedalienGreenIguanasonsaintlucia,shownaspresencein1x1-kmsquares.thedis-tributionofthesaintluciaIguanawasdeterminedduringfivemonthsofisland-wideface-to-facequestionnairesin2004–05(morton2007).“Falseabsences”wereminimizedbyquestioninglargenumbersofrespondentsaboutiguanasightingsinsquareswherethepresenceofiguanashadbeenindependentlydeterminedpriortothesurvey.Fromthis,wedeterminedtheminimumnumberofrespondentsthathadtobequestionedpersquareinordertoyield95%confidencethatatleastonerespondentwouldreportasightingif,infact,iguanaswerepresent.sightingsdeter-minedas“falsepresences”forthesaintluciaIguana(forquestionnairereturns,forexample,describingcaptiveiguanasorclearlyreferringtootherspecies,notablyCnemidophorus vanzoi)wereexcluded,althoughsomeoftheoutlying1x1-kmsquaresinthedistributionshownherecouldbefalsepositives,wherethepresenceofiguanashasnotbeenconfirmedbysearches.thedistributionofalieniguanaswasdeterminedfromquestionnaireresponsescollectedin2008–09.thislatterquestionnaireisanongoingefforttomaintainanupdateddemarcationofthespreadofthealienpopulation.


populations.meanclutchsizeforthealieniguanais40(n=4clutches),butwithamaximum(todate)of60,whichmightmorecloselyrepresentthetruemeanassamplesizeincreases.bycontrast,meanclutchsizeforsaintluciaIguanas(n=14)is23,suggestingthatthealienscouldout-reproducethenativeiguanas.Competitioncouldalsobeasymmetricalinfavorofthealiensif,asthelimiteddataimply,thealiensattainagreaterbodymass,althoughthismightreflectlowermortalityfrompredatorsinapopulationthatisputativelystillrarecomparedwiththatofthesaintluciapopulation. burton(2004)citedanotherconcernover the introductionofI. iguana ontoGrandCayman,thenativerangeofC. lewisi,namelyconfu-sionofpublicawarenessmessagesfortheconservationofC. lewisi.thisis

asubstantiveconcerngiventhegrowingdiscontentovernegativeimpactsofintroducedGreenIguanasonGrandCayman.AsKryskoetal.(2007)noted,theintroductionofI. iguana toFloridainthe1960sraisedlittlecon-cern,butbythe1990s,thepopulationhadexplodedandpublicdiscontentwithiguanasgrew. othernegativeimpactsofalieniguanas,suchascropdamage(particu-larlytoornamentals),theriskofSalmonellainfections,orpossiblepredationonbirdeggsarealmostcertainlyexaggerated,althoughinvasiveiguanascanbeanuisanceandarelikelytohaveatleastsomeeconomicimpactonvegetation.WhatisclearisahighlevelofpublicdiscontentinplaceswhereinvasiveGreenIguanashavebecomeestablished,inlargepartdueto

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saintluciaIguanas(A,adultmaleandb,adultfemale)andalieniguanasintroducedinsaintlucia(C,adultmaleandD,adultfemale).themorenumerousandprominentnuchaltubercleofthealieniguana,anditslargersubtympanicscale,areclearlyvisible,asisthedifferenceinthecoloroftheiris.

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delayedandinadequateinterventionduringtheearlystagesofinvasion.InpuertoRico,alieniguanasareconsideredanair-strikeriskonrunwaysattheluismuñozmarínInternationalAirport,causingthesuspensionofflightssixtimesinonetwo-monthstudy(engemanetal.2005).sementellietal.(2008)documenteddamagetopublicinfrastructurecausedbyburrowingiguanasintroducedintoFlorida. Atthistime,thealieniguanapopulationonsaintluciaappearstobegeographicallyisolatedfromthenativepopulation,butbydintofsmallpopulationsizeratherthananyimpenetrablegeographicbarriers.evenifthemountainousrain-forestedinteriorofsaintluciadoesprovideabar-rier(wehavefoundnoreportsofsaintluciaIguanasinthisregion),thealienpopulationcouldstilldispersearoundthecoastandcomeintocon-tactwiththenativepopulation.successfulnestingoutsideofcaptivityhasbeenrecordedinthesoufrièrearea,andquestionnairereturnsandfieldobservationssuggestthattheintroducedpopulationhasdispersedatleast2kmfromthesiteoftheinitialintroduction,ahotelontheoutskirtsofsoufrière.mappingcapturesandreportedsightingsimplydispersalhasbeenprimarilyalongtheriverinecorridorsoftheRiversoufrièreandsomeofitstributaries,althoughthisconclusionmightreflectasamplingbias,astheselowerlyingareasaremorereadilyaccessiblebysearchteamsthantheverysteepslopesthatcharacterizemuchofthearea. thealieniguanasareeasilydistinguishedfromthenativesbyhavingalargernumberofmoredenselypacked,andmoreprominentnuchaltuber-clescales,whichare,bycomparison,muchreducedinsizeandnumberinsaintluciaIguanas.Inaddition,thesubtympanicplatethatdistinguishesI. iguana fromI. delicatissima ismuchlargerrelativetothetympanuminthealienpopulation,whereasthescalesimmediatelyanteriortothissubtym-panicplateareproportionatelymuchlargerinthesaintluciaIguana.theeyeofthesaintluciaiguana,atleastinadults,alsoappearstoconsistentlyhaveamuchdarkeriristhanthatofthealien.

Management of the Alien Iguanasthepathwayofintroductiontosaintluciaforalieniguanasappearstohavebeenviathepettrade,asappearstobethecaseforallalienpopulationsofI. iguana introducedintotheCaribbeanandelsewhere(Kraus2009).AfewindividualGreenIguanas(reportsvaryfromfourtosix)wereheldinaprivatecollection,withoutapermit,insoufrièreuntilatleast2002(m.morton,pers.obs.),afterhavingbeenimported,againwithoutapermit,ashatchlingsfromapetshopinCanadaduringthelate1980s(m.bobb,pers.comm.).theoriginofthesehatchlingsisunknown.Althoughtheundesirabilityoftheseanimalsbeingheldinanunsecuredprivatecollec-tioninsaintluciawasnotedin2002,confusionoverlegalinstrumentsapparentlyhinderedtheirconfiscation(tworemainingincaptivityin2009werebelatedlyconfiscatedandeuthanized).Inearly2008,reportsoffree-livingiguanas,adultsandhatchlings,inthesoufrièreareawerecollatedandmapped.Althoughthefoundingindividualswerestillincaptivityin2002(m.morton,pers.obs.),aquestionnaireofsoufrièreresidentscol-lectedreportsofiguanasightingspriorto2000(Krauss2010a),suggestingthattheybredincaptivityandoffspringapparentlyescapedpriorto2002(althoughthishasbeendeniedbytheformerowners). thecurrentgeographicalseparationofsaintluciaandalieniguanasontheislandandtheapparentlyrestricteddistributionofthealienpopula-tioncouldprovideanopportunityforeradicatingthealiens.Itmightalsoofferanopportunitytomitigatesomeoftherisksofmixedmessagesinpublicawarenesscampaignsthatcouldarisefromsimultaneouslytryingtoprotectoneiguanapopulationwhileattemptingtoeradicateanother. simulations of population growth using the population viabil-ityanalysissoftwareVortex(lacy1993,lacyetal.2009)haveledtosomepredictionsregardingthepotentialsuccessoferadicationefforts2.simulatingaworst-casescenarioof60%juvenilemortality(i.e.,thesameasthatestimatedforadultGreenIguanasinpanama;Randandbock1992)andanalienpopulationestablishedsince1990(i.e.,afewyearsafterhatchlingsinitiallywereimported)predictedthatextinctionprob-

abilitiesgreaterthanabout30%areveryunlikely—regardlessofanyhuntingpressurewecouldfeasiblyapply(morton2010).bycontrast,arelativelymodesthuntingpressure(60–180adultsor600–1,000juvenilesremovedperyear)couldraiseextinctionprobabilitiestoalmost100%inabest-casescenarioof90%juvenilemortalityandapopulationestablishedforonlyfouryears(i.e.,accountingonlyforthefirstrecordedappear-anceofhatchlingsinthewild.thisbest-casejuvenilemortalityisslightlylowerthanthatreportedbyharris(1982)andvanDevender(1982),butthosestudiescouldnotdistinguishlossofjuvenilesthroughemigrationanddeath).thesamehuntingpressureispredictedtoincreaseextinctionprobabilitiestoabout80–90%forintermediatescenarios(75%juvenilemortalityand/orapopulationestablishedsince2002,thelastyearwhenalltheknownfounderiguanaswereconfirmedtobeincaptivity).Vortexsimulationsalsopredictedthemostlikelycurrentalieniguanapopulationsizeundertheworst-casescenariotobeinthetensofthousands,whichisinconsistentwiththedifficultyoffindingalieniguanasaroundsoufrière,suggesting,atleastforthemoment,thattheworst-casescenarioisnotthemostaccuraterepresentationofreality.Allscenariospredictexplosivepopulationgrowthifuncheckedbyaggressivemanagementinterventions. In2010,theslFDandDurrellestablishedafour-yearcollaborativepartnershiptoeradicatethealienpopulationofI. iguana(Krauss2010b).todate,however,evenrelativelymodesthuntingpressuresusedinthesimulationsdescribedaboveexceedthealieniguanaremovalratesthatwehavebeenabletoachieveinthefirstyearofthisproject,despiteintensivesearches.Atthistime(late2010),only21adults(includingsub-adults)and132hatchlingshavebeenremoved.Alliguanaremovalshavebeentheresultofintensivevisualsearchessupplementedbyreportsfromlocalresidents,whichledtocapturesbyhandorpole-noose.Capturediguanasareeutha-nizedbylethalinjectionadministeredbytheVeterinaryDepartmentofmAlFF.ourlowcapturerateislikelyduetoacombinationofthecrypticnatureofiguanasatrestintreesandaniguanapopulationputativelyatlowdensities.Attemptstoevaluatedifferentsearchandcaptureapproachesareunderway.

2populationviabilityanalyses(pVAs)havebeencriticizedfornotprovidingvalidpredictionsofextinctionrisk,especiallyoverlongtime-scales(e.g.,>100yrs),astheyassumeconstantecological(andanthropogenic)processesthatareunlikely(boyce2001).theirutilityincomparingtherelativeeffectsofalternativeman-agementscenarios,however,hasbeennoted(boyce2001).AnothercommoncriticismofpVAsisthatallofthemodelparametervaluesarerarelyknownforthespeciesbeingexamined.AsWhite(2000)commented,ifparameterestimatesarelargelyguesses,thenmodelpredictionsarealsoguesses.boyce(2001),how-ever,pointedoutthatobtainingaccurateestimatesforallparametersisunrealisticandshouldnotdetractfromtheheuristicuseofpVAs,forexample,whencom-paringalternativemanagementscenarios.Forthescenariospresentedhere,someparameterestimatesweretakenfrompublishedliteratureonI. iguana.Ageoffirstreproductionwasestimatedattwoyears,whichispessimisticwhencomparedwithZugandRand’s(1987)estimates(butseeandprattetal.1994,citedinRodda2003).Reproductionwasassumedtobepolygynous(referencesreviewedinRodda2003).Adultmortalitywassetat60%usingRandandbock’s(1992)estimates,andjuvenilemortalitywassetat(asaworstcase,andpresumablypes-simistic)thesame.Asabestcase(andprobablyoptimistic),itwassetto90%,asnotedabove.twoparameterswerebasedonunpublisheddatacollectedbyusin2010,albeitusingsmallsamples:sexratioatbirthwasassumedtobe1:1basedonasexratioof37:40fromasampleof77hatchlings.meanclutchsizewasestimatedat40±12.18basedonasampleoffourgravidfemalesin2010.otherparameterswereguesses.percentofadultsbreedingwasestimatedas62.7%formales(givenpolygyny)and100%forfemales.environmentalvariationwassettohavenoeffectonbreeding(i.e.,novariationinreproductivesuccessfromyeartoyear).CarryingcapacitywassetashighasVortexallows(60,000)tomodeluncheckedgrowthinanexpandingpopulation.moredetailsareprovidedinmorton(2010).Althoughalloftheseparametervaluesareuncertain,webelievetheyarebiologicallyplausible(andmorelikelytoerronthesideofpessimism)—andtheydoallowustomodeldifferentmanagementscenarios.Asmoredataaccumulatefromalieniguanacapturesinsaintlucia,wewillbeabletorefineourparameterestimatesandrunfurthersimulations.Fornow,theydemonstratethepotentialutilityofthistoolandallowustosimulateresponsesofapopulationthesizeofwhichisbeyondourmeanstoreliablyestimateatthispoint.


Apilotstudyinseptember2009alsodeployed50tomahawklivetrapsand20releasing-locksnaresinabout0.5km2withinthecoreareaofiguanasightings,butnoiguanaswerecapturedbyeithermethodoverafour-weekperiod.trapsandsnareenclosureswerebaitedwitharomatic

fruit(e.g.,mango,banana)plusfragmentsofredfabric,asiguanasareknowntorespondtoreditemssuchashibiscusflowers(e.g.,Alberts2003).ourlackofsuccesscouldbeattributabletothearborealityoftheiguanas,theputativelowpopulationdensity,and/oranapparentsuper-abundanceoffood(foliage),whichwouldrenderthebaitedtrapslessattractive.Furthertrappingeffortsareprojectedforthenestingseasonin2011,whichispre-sumablyfromFebruarytoearlymay—basedonthetimingofnestinginsaintluciaIguanas(morton2007)andconsistentwiththetimingofalienhatchlingcapturesin2010.Duringthisperiod,femaleiguanaspresumablyspendagreaterproportionoftheirtimeonthegroundand,post-nesting,arelikelytobemoremotivatedtofeedafterhavingsuspendedfeedingwhilegravid(Rodda2003).Wealsoplantopilotandevaluatetheuseofdetec-tordogstolocatealieniguanasin2011.Detectordogshavebeenusedtolocateanotheralieninvasivearborealreptile,thebrowntreesnake(Boiga irregularis)onGuam,bothinairportcargo(Viceandengeman2000)andfree-livingintrees(savidgeetal.2008). Giventheallopatricdistributionofthetwoiguanapopulationsonsaintlucia,webelievethatnohybridizationhasyetoccurred.tissuesam-plesfrombothpopulationshavebeenbankedinordertosearchforallelesuniquetothealienpopulation.thesewillallowustosurveythenativepop-ulationforevidenceofintrogression.Althoughthealieniguanapopulationcurrentlyappearstoberestrictedtoarelativelysmallareainsaintlucia,itisreproducingandwillpresumablyincreaseitsrange.Kryskoetal.(2007)suggestedthatapopulationexplosionofalienI.iguana inFloridafollowedhurricaneAndrewin1992.possibleeffectsofhurricanetomasonsaintluciainoctober2010areunknown.thesoufrièreareawasamongtheworstaffectedbyseverefloodingandextensivelandslides.Alieniguanas,bothadultsandyoungoftheyear,however,havebeencapturedinthecoreareasincetomasstruck.Aswithmanyotherinvasivespecies,reptilianinva-sionsareoftencharacterizedbyanextendedlagperiod,sometimeslastingdecades,whichprecedesanexplosioninnumbersandrange(e.g.,Kryskoetal.2007,Kraus2009).Questionnairereturnsfrom2009and2010sug-gestedthatthealieniguanapopulationinsaintluciamightnowbemov-ing,orabouttomove,beyondthislagperiodintoaperiodofrapidgrowth.effortstoeradicatethealieniguanapopulationmustincludestrategiesforpreservingthesaintluciaIguanainthefaceofongoingthreatsemanatingfromthepossibilityofcontactwiththealieniguanapopulation.

Acknowledgementstheworkonwhichthisarticleisbasedwasfundedbythebalcombetrust,theInternationalIguanaFoundation,and(fortheremovalofalieniguanascomponent)theGlobalenvironmentFacilitythroughtheUnitednationsenvironmentprogramme andmanagedbyCAbI.the comments ofGordonRoddaandCharlesKnappgreatlyimprovedthefinaldraftofthismanuscript.manyslFDstaff,localfieldassistants,andoverseasvolunteerscontractedbyDurrellhavecontributedenormouslytothisworkin2002–2010,especially(forworkonsaintluciaIguanas)Agnesorgrodowczyk,AvrilKelly,bradleyAbraham,Curtismathurin,DenisemcGowan,egilDröge,Gabybesne-Garcia,GregAlexander,hannaho’Kelly,lizCorry,lizzieperret,meaganmcpharlin,neilocculi,andsophieharley;and(forworkonalieniguanas)Alansands,AmyClack,barbaraschaeffer,benjibarca,GraemeDuncan,lenniGriffiths,maryJames,mikeball,nateWood,nickCondie,peterJn.baptiste,Rachelbarwick,RobWilliams,RuthmcKee,stephenlesmond,andtwylaholland.specialthanksareduetoDonaldAnthony,whoinitiatedtheworkonsaintluciaIguanas;Anthony“seako”Johnny,long-timefieldassistantandlocaliguanaexpert;timotheusJn.baptiste,AlwinDornelly,andlyndonJohnfortheirongo-ingcontributionstoallaspectsofprojectmanagement;RichYoungforsageadviceonstudydesignandanalyses;andKarenGrahamforallhersupportandwisdom.Finally,aspecialmentionmustgotoourcolleagueonthealieniguanaremovalwork,lesterJn.baptiste,whowaskilledinalandslideduringhurricanetomasin2010;heisgreatlymissedbyallwhoworkedwithhim.

predictedeffectofhuntingpressureappliedtojuvenilesandadultsinthealieniguanapopulationestablishinginsaintlucia.thegraphsshowthepredictedprob-abilitythatthepopulationwillbeextirpatedifitisexperiencing:(a)low(60%)juvenilemortality,(b)moderate(75%)juvenilemortality,or(c)high(90%)juve-nilemortalityinadditiontothehuntingpressureapplied.thethreelinesineachgraphillustrateuncertaintyoverhowlongthispopulationhasbeenestablishing;i.e.,huntingissimulatedasbeingappliedtoafour-year-oldpopulation(huntingstartsatpoint4Yo),anine-year-oldpopulation(at9Yo),ora21-year-oldpopulation(at21Yo).Forapopulationexperiencinghighjuvenilemortalityinadditiontohunt-ing(c),theprobabilityofextirpationishigh,regardlessofhowlongthepopulationhasbeenestablishing.Formoderateorlowjuvenilemortalityinadditiontohunting(b,a),ahighprobabilityofextirpationexistsonlyfortheyoungerpopulations(firsthuntedat4Yoor9Yo).simulationswerecarriedoutusingVortex(lacyetal.2009);thisfigurewasadaptedfrommorton(2010),whodetailedtheVortex modelparametervaluesusedinthesimulations.

30 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 moRtonAnDKRAUss

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malesaintluciaIguana(Iguanacf.iguana).thisindividualisknowntohavebeenkilledforfoodbyhuntersatGrandAnseestatetheyearafterthisphotographwastaken.

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AburnedwetlandatpaloVerdenationalpark,CostaRica;notetheaggregationofdeadscorpionmudturtles(Kinosternon scorpioides)intheforeground.

GUInnAnDlUGeR

IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 35neWobseRVAtIonsoFAneotRopICAlmUDtURtle

Reading the ashes:arson Decimates a Tropical Wetland,

but allows new observations of a neotropical Mud Turtle

Jeremye.Guinnandpalaniluger

environmentalscienceprogram,sittingbullCollege,Ft.Yates,northDakota58538,UsA

Allphotographsbytheseniorauthor.

DuringanationalscienceFoundation-fundedsummerresearchpro-gramforundergraduates,wewitnessedthelossofmorethan5,000

haofprimesemi-fossorialturtlehabitatnearCatalinastation,paloVerdenationalparkinGuanacaste,CostaRica.Accordingtoparkofficials,thefirewassetintentionallybypoacherstoallowaccesstowildlifefromthetempisqueRiver,whichbordersthepark.naturalwildlandfirescanbedevastating,butalsobeneficialforalocalecosystem;however,human-inducedfiresaregenerallymoredestructiveduetolackofperimetercon-trolsandheavyfuelloads.Wetlandfiresareespeciallydamagingbecauseof

thelossofhabitatandindividualsfromspeciesthatarenotadaptedforfastoverlandmotility.AlthoughCattails(Typhasp.)areexoticinthisregion,theyprovidehabitatforanumberofnativespecies,includingsemi-fossorialturtles.thefireprovidedauniqueopportunitytoviewawetlandcommu-nitythatwewouldotherwisehavebeenhardpressedtoobserve. As the fire still smoldered in the Área de Conservación Arenaltempisque,wemadeapreliminaryhikearoundthesouthernperimeterofthefire.thelandscapewasdominatedbyhundredsofblackVultures(Coragyps atratus)andCrestedCaracara(Polyborus plancus)feastingonthe

AnirruptionofblackVultures(Coragyps atratus)thatarrivedtopickupthepiecesfollowingthefire.

36 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 GUInnAnDlUGeR

remainsofturtlesandotherwildlifekilledbythefire.theputridcombi-nationofsmellsfromburnedflesh,ash,andwetlandsludgehungintheairandclungtoourclothes.thevisualscenewasnomoreappealing;anastonishingabundanceofdeadturtleswasclearlyvisibleevenfrombeyondtheborderofthefirezone.sodramaticwastheeffectthatfrontpagesoflocalandnationalpapersincludedanimageofaburnedturtle(CamposandArroyo2008).

BackgroundpaloVerdenationalparkislocated28kmsouthofbagaces,CostaRicainthetempisquebasin.theparkconsistsofmorethan13,000haof

floodplain,marshes,limestoneridges,andseasonalpoolsinthemiddleofthedriestregioninCostaRica.paloVerdeservestoconnecttotwootherreserves,theDr.RafaellucasRodriguezCaballeroWildlifeRefugeandthelomasbarbudalbiologicalReserve.theswamplandsthatconnectpaloVerdetotheotherreservesarethreatenedbylocalentrepreneurswhohaveinvestedinpermitstocleartheswampforestandplantcrops,whichwouldfurtherincreasetheisolationofthesereserves(baker2004). theregionexperiencesdistinctwetanddryseasons.thelackofpre-cipitationfromroughlynovembertomarchcausestheregiontobecomeverydry.Duringthisperiod,somefarmerswillintentionallystartfiresinordertoclearlandfortheircrops.Inthisregion,iflandisclearedofforest

Remainsofscorpionmudturtles(Kinosternon scorpioides)consumedbyfireatpaloVerdenationalpark.theindividualintheupperrighthadpartiallyburieditselfduringthefire.

IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 37neWobseRVAtIonsoFAneotRopICAlmUDtURtle

bynaturalcauses,itisessentiallyopentofarming.Additionally,poacherssetfirestoimproveaccesstocertaincovetedwildlifespecies. paloVerdeanditssurroundingreserveshave15differenthabitats,includingseveraltypesofswampsandmarshlands.thesehabitatsprovideforawealthofspeciesdiversityrangingfromWhite-tailedDeer(Odocoileus virginianus),White-nosedCoati(Nasua narica),andAmericanCrocodile(Crocodylus acutus)toendangeredspeciesincludingtheGreatCurassow(Crax rubra),Yellow-napedparrot (Amazona auropalliata), andKingVulture(Sarcoramphus papa). prescribedfireisabeneficialmanagementtoolthatcanmaintainorrestoredesired,historicecologicalconditions(e.g.,brockwayandlewis1997,CarterandFoster2004).prescribedburninghasbecomeaprimarytoolofecosystemrestorationintheUnitedstatesbecauseofthebenefitsandhistoricalassociationoffireinsomeecosystems(Johnsonandhale2002).Withsuchanincreaseintheuseofprescribedfires,understandingtheeffectsonwildlifebecameincreasinglyimportant(Fordetal.1999).Unfortunately,littleinformationexistsabouttheeffectsofprescribedfireonamphibiansandreptiles(mcleodandGates1998,Fordetal.1999,Greenburgetal.1994,pilliodetal.2003).Althoughsomeobviousandimmediatedeclinesofamphibianandreptilianpopulationswilloccurdur-ingawetlandfire,vegetativerecoveryandrestructuringmightprovidebet-terhabitatforindividualpopulations—asubjectinneedofmorestudy.

Recording the EffectsWedocumentedturtlesandotherspecieswithinandaroundthefirezonebyutilizingstandardizedtransectsamplingmethods.Wealsoattempted

todetermineturtledensitiesindifferentareasofthefirezonebyusingastratified-randomstudydesign.thefirezonewasclassifiedintothreecategories:(1)Closetotheignitionpoint,(2)closetotheterminusofthefire,and(3)anintermediatearea.Withineachzone,werandomlyselectedthree1,000x1,000-mstudyplots.Withineachplot,wecountedturtleswithinfive300-mtransects. Weexpectedtofindanassemblageofturtlespeciesinthewetland,butthescorpionmudturtle(Kinosternon scorpioides)wasconsideredthemostlikelytooccur.Itrangesatlowelevationsfromsoutherntamaulipas,méxico,southwardtonorthernArgentina,bolivia,andnorthernperú(ernstetal.1992).thisspeciesisomnivorousandwilleatfish,snails,amphibians,insects,algae,andotherplants(Vanzolinietal.1980).Ithasahigh-domedcarapacethatcanmeasureupto17.5cminlength.thecarapacehasthreewell-developedlongitudinalkeelsandisanunmarkedbrowntoblackincolor.theplastronhasasinglemovablehingebetweenthepectoralandabdominalscutes.theplastronmightnotbelargeenoughtocompletelyprotecttheanimalwhenclosed(ernstetal.1992,berryandIverson2001). one subspeciesofscorpionmudturtle, theRed-cheekedmudturtle(K. s. cruentatum),wastheonlyturtleobservedinthefirezoneoradjacentareas.turtlesoccurredatadensityof6.33turtles/transector316.5turtles/ha.thiswassimilar,butslightlygreaterthanthe254turtles/hareportedbymedinaetal.(2007)foranother subspeciesofK. scorpioides(K. s. albogulare).Intemperateregions,Gibbons(1983)reported56K. subrubrum/haandboninetal.(2006)reporteddensitiesforseveralspeciesofmudandmuskturtles,including260turtles/haforK. subrubrum,229

Additionalspecieskilledinthefirezoneincludednortherntamandua(Tamandua mexicana;A),White-lippedpeccary(Tayassu pecari;b),White-nosedCoati(Nasua narica;C),andboaConstrictor(Boa constrictor;D).

A

C

B

D

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turtles/haforSternotherus carinatus,and194turtles/haforS. odoratus.nodensityestimateswereavailableforK. s. cruentatuminotherareas. nosignificantdifferences(one-wayAnoVA,F=0.08,p=0.10)existedinthenumbersofturtlesobservedamongsitecategories,suggestingthatturtlesdidnotmakelargemovementsawayfromtheignitionpoint.mudturtlesmightbeparticularlysusceptibletofiresduetoanadaptationtoseasonaldryweather;theyoftenburrowinthemuduntilthenextrain(ernstetal.1992).thisadaptationcouldbeoneexplanationforwhysomanymudturtleswerekilledinthepaloVerdemarshlandfire.perhapstheydidnottrytoescapethefire,butinsteadsimplytriedtoburrowintothesediments.however,teska(1976)reportedsubstantialmovementsofturtlesinGuanacasteprovinceduringthedryseason. Additionalspecieskilledinthefirezoneincludednortherntamandua(Tamandua mexicana),White-lippedpeccary(Tayassu pecari),White-nosedCoati(Nasua narica),andboaConstrictor(Boa constrictor)alongwithsev-eralunidentifiablesnakespecies.othersources(CamposandArroyo2008)alsolistedtheGrayFox(Urocyon cinereoargenteus).Documentationofsuchmobilespeciesandaccountsfromparkofficialssuggestthatthefirewasextremelyfast-moving.however,thesespeciesmighthavetriedtoclimborburrowinunsuccessfulattemptsatescapingthefire.Interestingly,nocrocodilianswereobservedinthefirezone. While the lossof individualsdue to this actof arson cannotbeoverstated,theinsightintoturtlecommunitiesprovidedbythisresearchincreasesparkmanagers’knowledgeoftheherpetofauna.Accordingtosavage(2002),“verylittleisknownofthelifeofthiscommonspecies”(Red-cheekedscorpionmudturtle).Also,duringourfinaltransect,werecordedtheonlylivingturtlesthatwereobservedduringourentiretimeatpaloVerdenationalpark.threeRed-cheekedmudturtlesmadetheirwayalongthefireline,theirshellsslightlycharred,butotherwiseunscathed.bytheendofourstudy,freshgreensproutswerealreadyeruptingfromtheashes.moll(1990),Gibbonsetal.(1983),andpritchardandtrebbau(1994)suggestedthatscorpionmudturtlescouldrapidlymoveintodif-

ferentareasinresponsetodroughtandhabitatavailability.theoutlookforreestablishingtheturtlepopulationinthisareaappearstobeverypositive.

Literature Citedbaker,C.p.2004.Moon Handbook: Costa Rica.5thed.Avalontravelpublishing,

emeryville,California.

berry,J.F.andJ.b.Iverson.2001. Kinosternon scorpioides.Catalogue of American Amphibians and Reptiles(725):1–11.

bonin,F.,b.Devaux,andA.Dupré.2006.Turtles of the World.translatedbyp.C.h.pritchard.JohnshopkinsUniversitypress,baltimore,maryland.

brockway,D.G.andC.e.lewis.1997.long-termeffectsofdormant-seasonpre-scribedfireonplantcommunitydiversity,structure,andproductivityinalongleafpinewiregrassecosystem.Forest Ecology and Management 96:167–183.

Campos,C.andF.Arroyo.2008.tragediaincalculableenpaloVerde:biologosestimanquerecuperciondeecosistemastardarasiglos.Al Día Periódico,11may2008.

Carter,m.C.andC.D.Foster.2004.prescribedburningandproductivityinsouth-ernpineforests:Areview.Forest Ecology and Management191:93–109.

Costa, h.C., F.b. molina, V.A. sao-pedro, and R.n. Feio. 2010. Reptilia,testudines,Kinosternidae,Kinosternon scorpioides scorpioides (linnaeus,1776):Distributionextension.Check List6:314–315.

ernst,C.h.,R.G.m.Altenburg,andR.W.babour.1992.Turtles of the World.smithsonianInstitutionpress,Washington,D.C.

Ford,W.m.,m.A.menzel,D.W.mcGill,J.laerm,andt.s.mcCay.1999.effectsofacommunityrestorationfireonsmallmammalsandherpetofaunainthesouthernAppalachians.Forest Ecology and Management114:233–243.

Gibbons,J.W.1983.Reproductivecharacteristicsandecologyofthemudturtle,Kinosternon subrubrum(lacépède).Herpetologica39:254–271.

Gibbons,J.W.,J.l.Greene,andJ.D.Congdon.1983.Drought-relatedresponsesofaquaticturtlepopulations.Journal of Herpetology17:242–246.

Greenburg,C.h.,D.G.neary,andl.D.harries.1994.effectofhigh-intensitywildfireandsilviculturaltreatmentsonreptilecommunitiesinsand-pinescrub.Conservation Biology8:1047–1057.

Johnson,A.s.andp.e.hale.2002.thehistoricalfoundationsofprescribedburn-ingforwildlife:Asoutheasternperspective,pp.11–23.In:W.m.Ford,K.R.Russell,andC.e.moorman(eds.),theRoleofFireinnongameWildlifemanagement and Community Restoration: traditional Uses and newDirections.proceedingsofaWorkshop.15september2000,nashville,tennessee,UsA.GeneraltechnicalReportne-288.UsDAForestservicenortheasternResearchstation,newtownsquare,pennsylvania.

medina,G.F.,o.V.C.mora,ando.montenegro.2007.Abundance,populationstructure,andconservationofKinosternon scorpioides albogulareontheCaribbeanIslandofsanAndres,Colombia.Chelonian Conservation and Biology6:163–169.

mcleod,R.F.andJ.e.Gates.1998.Responseofherpetofaunalcommunitiestofor-estcuttingandburningatChesapeakeFarms,maryland.American Midland Naturalist139:163–177.

moll,D.1990.populationsizesandforagingecologyinatropicalfreshwaterstreamturtlecommunity.Journal of Herpetology24:48-53.

pilliod,D.s.,R.b.bury,e.J.Jyde,C.A.pearl,andp.s.Corn.2003.FireandamphibiansinnorthAmerica.Forest Ecology and Management178:163–181.

pritchard,p.C.h.andp.trebbau.1984.The Turtles of Venezuela.societyforthestudyofAmphibiansandReptiles,Ithaca,newYork.

savage,J.m.2002.The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas.UniversityofChicagopress,Chicago,Illinois.

teska,W.R.1976.terrestrialmovementsofthemudturtleKinosternon scorpioidesinCostaRica.Copeia1976:579–580.

Vanzolini,p.e.,A.m.m.Ramos-Costa,andl.J.Vitt.1980.Repteis das Caatingas.AcademiabrasileiradeCiencias,RiodeJaneiro,brasil.

oneofthefewscorpionmudturtlesthatsurvivedthefire,itsshellslightlycharred,butotherwiseunscathed.

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c o M M e n T a R y

Wearecurrentlyinabiodiversitycrisis.Aquarterofallmammalsfaceextinction,and90%ofthelargestoceanfisharegone.species

aregoingextinctatratesequaledonlyfivetimesinthehistoryoflife.butthebiodiversitycrisiswearecurrentlyencounteringisn’tjustalossofspe-cies,it’salsoalossofknowledgeregardingthem.scientistswhoclassify,describe, andexamine the relationshipsbetweenorganismsare them-selvesgoingextinct.themillionsofdollarsspentgloballyontechnologytocatalogspeciesmayactuallybepushingoutthepeoplewerelyupon:taxonomistsandsystematists.We’relikeyoungchildrenfrantictoaddnewbaseballcardstoourcollections,whiletheactualcreatorsofthebaseballcardsthemselvesarevanishing. take,forexample,theaplacophorans,arare,raregroupofinverte-bratescloselyrelatedtooctopuses,squids,snails,andclams.mostofuswillneverseeevenoneoftheapproximately360knownspeciesofsmall(lessthanacoupleofincheslong)aplacophoransthatinhabitoceandepthsgreaterthan50feet—butignoranceofthisgroupisnotlimitedtothepublic.Fewerthantwodozenscientificpapershavebeenpublishedonthegroupsince2005,eventhoughmanynewspeciesawaitdiscoveryanddescription—andmostofthesestudiesweredonebyonescientist,thevenerableAméliescheltemaofWoodsholeoceanographicInstitute.Assheedgesclosertoretirement,shemaysadlybecomethelasttostudyapla-cophorans.If50%ofthespeciesofaplacophoranwentextincttomorrow,wewouldneverknow. Amelie’sstoryistragicallycommon.martinsørensenofthenaturalhistorymuseumofDenmarkisoneoftheveryfewactivekinorhynch,ormud-dragon,taxonomists.martinalsorepresentsoneofonlytwoliv-ingtaxonomistswhohavestudiedgnathostomulids.theother,Wolfgangsterrer,isretired.bothkinorhynchsandgnathostomulidsaresmall,lessthanone-tenthofaninchinlength,anddwellbetweengrainsofsandandmudontheoceanfloor.Fewerthan300speciesaredescribedfrombothofthesephyla—thebroadestclassificationintowhichscientistsgroupanimals—andourknowledgeofthemisbasedalmostentirelyoncol-lectionsfromthewell-exploredeasterncoastoftheUnitedstates,themediterranean,andthewesterncoastofeurope. “evenwithintheseareas,newspeciesappearquiteoften,andwhenIcollectoutside[theseareas],Ialwaysexpecttofindundescribedtaxaexclu-sively,”sørensenwroteinarecente-mailtome.hisnewworkintheeastChinaseahasalreadyuncovered15newspecies.Indeed,themorninghee-mailedme,sørensen,lookingthroughhismicroscope,hadjustdiscov-eredanothernewspecies.“thenumberoftaxonomistsworkingontheseobscuretaxahasalwaysbeenratherlow(whichexplainsourlimitedknowl-edgeaboutthem),butwithinthelast20yearstaxonomyasadisciplinehas

comeunderevenharderpressurewhichhasresultedinafurtherdeclineinthenumberofexperts,”sørensenwrote. thisproblemplagueswell-knowngroups,too.Forexample,nema-todesrepresentmorethan28,000describedspeciesoffreshwater,marine,terrestrial,andparasiticroundworms.ontheseafloor,theyaccountfor85–95%ofallorganisms.however,anewstudyfoundthenumberofscientificpapersdescribingnewnematodespeciesishalfofwhatitwas

The Mass extinction of scientists Who study species1

CraigmcClain2

1Adaptedwithpermissionfrom<www.wired.com/wiredscience/2011/01/extinc-tion-of-taxonomists/>(posted19January2011).

2CraigmcClainisassistantdirectorofscienceforthenationalevolutionarysynthesisCenterandhasconducteddeep-searesearchfor13years,participatingindozensofexpeditionsinthemostremoteregionsofthepacificandAtlanticoceans.mcClainfocusesontheecologicalandevolutionarydriversofmarine-invertebratebiodiversityandbodysize,mostlyindeep-seasystems.heoftenexplorestheconsequencesoffoodlimitationonbiologicalsystems.mcClainisalsothechiefeditoroftheaward-winningscienceblog,Deep-seanews.

thedearthoftaxonomistsandsystematistsisnotasdireforterrestrialvertebratesasforinvertebrates.nevertheless,ongoingnewdiscoveriesofmammals,birds,reptiles,andamphibianspreviouslyunknowntoscienceprovideampleevidencethatwemightbeunderminingoureffortstoprotectbiodiversitybynotsupportingeffortsthatencourageyoungscientiststolearnandimplementtheskillsnecessarytogenerateessentialknowledgeoflife’sdiversitybeforeitdisappears.thisandthefollowingimagesfeaturefourU.s.-basedscientistsactivelyinvolvedinthetaxonomyandsystematicsofamphibiansandreptiles.Janaleep.Caldwell,ofthesamnobleoklahomamuseumofnaturalhistoryandDepartmentofZoology,Universityofoklahoma,inafieldcampinbrazil.Amazoniaandtheneotropicsingeneralhar-boraplethoraofundescribedspecies.CaldwelldescribedthedendrobatidHyloxalus chlorocraspedusin2005.malesserveastheparentalfrog,carryingasmanyastentadpolesontheirbacksbeforereleasingthemintosmallforestpoolsshortlybeforemetamorphosis.

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adecadeago,andathirdofthedecadebeforethat.Anywherebetween10,000and100,000speciesremainundescribed. Whythelossoftaxonomists?Wehavedevaluedtheircontributions,bothmonetarilyandscientifically.someattributethedeclineoftheseresearcherstothereplacementofoutdatedmethodsthatwouldnotmeetthescrutinyofsciencetoday.thesecriticsenvisiontaxonomistsaslone

l.leeGrismer,oflasierraUniversityinRiverside,California,incollaborationwithhissonandanumberofnationalscientists,hasdescribedover70newspeciesinthepastsixyears,allfromsoutheasternAsiaandmanyofthosefromthemalayArchipelago.theaptlynamedCnemaspis psychedelicaisaninsularendemicknowntooccurononlyoneof92poorlyexploredVietnameseislandsthatarebeginningtoshowasurprisingdegreeofendemismanddiversity.thisdiscoveryclearlysuggeststhat,despiterecentwork,theherpetofaunaldiversityoftheregionisbarelyknown.

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Rafebrown(topright),ofthenaturalhistorymuseumandbiodiversityInstitute,UniversityofKansas,withgraduatestudentsinthephilippines.Hylarana(formerlyRana) tipanan (center)wasdescribedbybrownandcolleagues in2000;whenitsIUCnRedliststatuswasassessedin2004,thespecieswasdeterminedtobeVulnerableduetoahighlyfragmenteddistributionattributablelargelytoacontinu-ingdeclineintheextentandqualityofitsforesthabitatonluzoninthephilippines.thediscoveryofa2-m-longlizard(bottom)intheforestsofthephilippinesspeaksclearlytothelackofcurrentknowledgeofthearchipelago’sbiodiversity,evenasitsforestsaredecliningatanalarmingrate.Varanus bitatawaisoneofonlythreeknownspeciesof frugivorous(fruit-eating)monitor lizards intheworld.e.o.Wilson(1992.The Diversity of Life.W.W.norton&Co.,Inc.,newYork)asked:“Williteverbepossibletoassesstheongoinglossofbiodiversity?Icannotimagineascientificproblemofgreaterimmediateimportanceforhumanity.biologistsfinditdifficulttocomeupwithevenanapproximateestimateofthehemorrhagingbecauseweknowsolittleaboutdiversityinthefirstplace.…wedonotknowthevastmajorityofspeciesoforganismswell;wehaveyettoanointsomanyas90per-centofthemwithscientificnames.”

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museumscientistssurroundedbydustywoodcabinetsandbottlesofform-aldehydewherespeciesdescriptionismoreartthanscience—butthispor-trayaloverlooksthesuiteofmoderngeneticmethodsthatthoseinterestedindiscoveryanddescriptionofnewspeciesusewithincreasingfrequency.

thisnewbreedoftaxonomistsincludesChrismahofthesmithsoniannationalmuseumofnaturalhistoryandAdrianGloverofthenaturalhistorymuseuminlondon,whoareamongtheworld’sleadingexpertsonseastarsandmarineworms,respectively.bothdemonstratethatthemostinformativesciencecomesfromsynthesizinggenetictechniqueswithmoreclassicaltaxonomybasedonknowledgeoftheanatomyandnaturalhistoryoforganisms. extinctionoftaxonomistscontinuesdespiteagrowingpooloffundsforbiodiversityprogramsanddatabases.eUnIs,eol,obIs…thelistgoeson.thesedatabaseshavepooledourcollectivebiodiversityknowledge,helpingidentifywhatdrivesbiodiversityandsetconservationpriorities.thankfully(myownresearchhasrelieduponthem),thousandsofhoursandmillionsofdollarshavebeenspentontheseinitiatives.however,manyoftheseprogramsdidnotfinanciallysupporttaxonomistsgeneratingthedatathesedatabasesrequired. Afteradecadeand650milliondollars,theCensusofmarineliferepresentsoneofthelargestinitiativestodocumentbiodiversityonourplanet.Insomeregards,itwasagreatsuccess,supporting2,700scientiststoproduce2,600newscientificpublicationsandthousandsofnewspeciesdescriptions.butastheCensusendsthisyear,noagencyororganizationisofferingtofillthefundingvoidpreviouslyfilledbytheAlfredp.sloanFoundation.perhapsmoreimportantly,theCensus,likemanyinitiatives,didnotprovidelong-termpositionsandappointmentsforthosedoingtaxonomicwork.manybiologydepartmentswithinuniversitiesnolongeremployataxonomist.theremainingpositionsarerelegatedtomuseums. Why?Assørensenexplains,“thedecliningnumberoftaxonomistsandsystematistsisatleasttosomeextentlinkedtothefactthatyoursci-entificproductiontodayshouldbemeasurable.”theunitsofmeasure-mentarecollectedgrantmoneyortheimpactfactorofajournalpaper.taxonomyhasneverbeenconsideredhot,andpuretaxonomicstudiesarerarelyfunded,hewrote.Departmentsneedgrantmoneytooperate. scienceasaninstitutionmayalsobepartlyresponsibleforundercut-tingtaxonomicwork.Althoughacrudemetricfraughtwithseveralissues,wemeasuretheimpactofascientificpaperbyhowmanytimesothersci-entificpapershavecitedit.similarly,wemeasuretheimpactofscientistsbycountingtheircumulativecitations.Unfortunately,taxonomicworkisrarelycited,evenwhenitshouldbe. ontheotherhand,thebrilliantbiodiversitydatabaseswehavecre-atedleadtoaplethoraofscientificpapers.thepaleobiologyDatabase,acomprehensiveonlinecatalogoffossilspecies,hasalreadygeneratedmorethan100publications.buttherequirementforusingthisdatabase,likemostothers,iscitationofthedatabaseitself,notthenearly35,000papersgeneratingtheoriginaldata. thedeclineintaxonomistsmeansthatatsomepointinthefuturewewillbeunabletotrainnewgenerationsoftaxonomists.thisproblemisrecognizedbythenationalscienceFoundation,whichin1994createdaprogramtoenhancetaxonomicresearch.Whilethisinitiativeprovidestraining,itdoesnotcreatejobopportunities.otherproblemsaretakingformtoo.Forexample,in2006,Isetouttoexplorehowbiodiversityandbodysizewerelinkedamonganimals.todosoIneededinformationonthelargest-andsmallest-sizedspeciesforeachgroupofanimals—some-thingsurprisinglynotreadilygarneredfromthepublishedliterature.Ireliedonmyconnectionswithtaxonomistsforguidanceandinformation,butformanygroupsIstruggledtofindacontact.evenforwell-knownanimals,Iwasamazedbyhowfewscientistsstillstudiedthem.mypersonalexperiencehighlightshowprogressinbiologyasawholemaybeimpededifwelosetaxonomy.theproblemwefaceisalossofknowledgenotyetrecordedinthescientificliterature.Inourtechnologicaleffortstoconcen-trateourbiodiversityknowledge,wemayberenderingafieldandbodyofknowledgeobsolete—and,intheprocess,wemaybeunderminingourowneffortstoprotectbiodiversity.

s.blairhedges,inhislaboratoryatpennsylvaniastateUniversity,hasfocusedmostofhisattentionontheWestIndianherpetofauna,whichhasgrownfrom585knownspeciesin1991towellover700today—withasmanyas1,000speciespossible—iftheyarediscoveredanddescribedbeforefallingvictimtodevelopment(mostattributabletothetourismindustry),deforestation,andinvasivespecies.theworld’ssmallestknownspeciesoflizard(Sphaerodactylus ariasae,fromthesouth-westernDominicanRepublic)andthesmallestknownsnake(Leptotyphlops carlae,frombarbados)bothoccurintheregion.

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thedeterioratingrelictsofhumanpresenceonsombrerohavebeeneffectivelyintegratedintothenaturalecosystem.

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thisisaninterestingtimetobealive.humansarerunningoutoflandandconstantly searching fornewopportunities to fueleconomic

growth.Atthesametime,weseeagrowingawarenessofthenaturalenvi-ronment,theresourcesitprovides,andtheharmfuleffectsofhumanpopu-lationgrowthontheearth’sotherspecies.somepeopledenythataprob-lemexistsandsuggestthatweshouldcontinueasweare.othersbelievethatweshouldparkourcars,moveintothewoods,abandontechnology,andbecomeonewithnature.mostofusfallsomewherebetweenthosetwoextremes.Ibelievethatwemustlearntocoexistwithnaturewhilefindingawaytofostereconomicgrowthwithoutdestroyingourplanet.Iamfascinatedbythatgoalandthinkthatthefirststepisinunderstandingwhatisatstake.howcanwecoexistwithnatureifweremainignorantofhercomponents?Ihadanopportunitytowitnessafewimplicationsofthisage-oldparadoxduringarecentvisittosombrero,atinyislandabout60kmnorthofAnguilla. onafinemorninginoctober2010,IwasoneofateamboardingtheaptlynamedOcean Girlz,acatamarancharteredbytheConservationAgencyforatriptosombrero.sombreroisunique.Risingoutoftheocean

sometimeintheearlypliocene,itisabout1.5kmlongandabout0.4kmatitswidestpointandhasneverbeenpartofanotherbodyofland.sombreroisthenorthernmostofthelesserAntilles,andispartofthenationofAnguilla.lifeonthe38haofmarlylimestonearrivedbymigrat-ingfromthemainlandorsmallerislandsonfloatingdebris,byflying,or

sombrero: lizards among the RuinselizabethWright

DepartmentofForestry,Universityofmissouri,Columbia,missouri

AsthenorthernmostofthelesserAntilles,sombreroisalongwayfromeverywhere,sittingonitsownislandbankandneverconnectedtoanyotherlandmass.

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sombreroisoneofthreelesserAntilleanislandsthatharborblackAmeivas.InadditiontoAmeiva corvina(namedafterequallyblackcrows),A. coraxlivesonlittlescrubIslandoffthenortheasterncoastofAnguilla,andA. atratalivesonRedonda,anotherisolated“rock”emergingfromthesea.

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Accessbyladderisnecessaryforscalingthesheerwallsoftheisland,limestonecliffsshooting12mstraightupoutofthewater.

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theanolesofsombreroareverysimilartoAnolis gingivinus,whichisubiquitousontheAnguillabank.Geneticstudieswilldetermineiftheyarethesamespecies.

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beingcarriedbystorms,birds,orhumans.theislandisfrequentlydelugedbygiantwaves,andmightbeentirelysubmergedduringseverehurricanes.suchconditionsmakeithardtogetthereandevenhardertosurvive.theresultingfoundereffect—thesometimesconsiderableimpactthatthefewfoundingindividualscanhaveonlatergenerations—dramatically

increasesthechanceofendemism(Ivie1999).thisiswhyweweregoingtosombrero. thetripwasorganizedbyDr.Robertpowell,aherpetologistatAvilaUniversity,whowantedtolearnmoreaboutthelizardsofsombrero.Dr.Jameslazell,thepresidentoftheConservationAgency,hadvisitedtheislandin1963andhaddiscoveredthatthreespeciesoccurredontheisland(notjusttheonethathadbeenpreviouslyreported):AnanolecurrentlyidentifiedasAnolis gingivinus,thecommonspeciesonthenearbyAnguillabank;aDwarfGecko,whichlazelldescribedasSphaerodactylus“near”spu-tator,thespeciesthatoccursonboththeAnguillaandst.Christopherbanks;andthespectacularblackGroundlizard,Ameiva corvina(lazell1964).In

theDwarfGeckosonsombreroexhibitvariablepatterns,somespotted,somealmostunicolored,andotherswithstripes,asseenonthesetwoadults(A/b)andonejuvenile(C).

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Island Girlz,hereanchoredoffsombrero,madetheexpeditiontotheislandacom-fortableexperience—althoughthecaptainhadnotpreviouslybeentosombrero.

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Abrownbooby(Sula leucogaster),silhouettedagainstthesunlitskyonwhatlookedlikearustyloadingcrane,greetedourarrival.

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Althoughproficientscavengerswhenbirdsorpeopleprovidewelcomemeals,theAmeivasofsombrerousuallyhavetodigandrootforfood.

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the1960s,whenDr.lazellvisitedsombrero,noonewasyetdreamingofthesecretsthatDnAmightonedayreveal,andtissuescollectedduringasurveybyDr.JennyDaltry(1999)weresubsequentlylost.thetimehadcometocollectsomemoresamplessothattherelationshipsandperhapstrueidentitiesoftheselizardscouldfinallybeestablished.ourteamalsoincludedDr.Gadperry,aconservationbiologistattexastechUniversity,Dr.Wenhualu,anentomologistwiththeConservationAgency,andsusanValentine-Cooper,anexperiencedentomologist,who,likeme,hadbeeninvitedtocomealongandhelpcollectsmallarthropods. so,therewewereaboardOcean Girlz,onourwayfromGuanaIslandinthebritishVirginIslands,wherewewereconductinglong-termresearchprojects,tosombrero.Iwasinaweoftheflyingfishastheyexplodedoutofthewaterandsoaredforwhatappearedtobehundredsofmetersacross

thewaves.Asweapproachedtheislandsomethreehoursintoourvoyage,wesawasmallvesselanchoredneartheladderthatprovidesaccesstotheisland.Accessbyladderisnecessaryforscalingthesheerwallsoftheisland,limestonecliffsshooting12mstraightupoutofthewater.Ialmostfellintothewateronmyfirstattempttostepfromthebowofourcatamaran,movingup-and-downwiththewaves,ontotheladder,andwassavedonlybythequickactionofthecrew.mysecondattemptwassuccessfulandIclimbedup,themetalhotundermyhands.

migratoryblackpollWarblers(Dendroica striata)dartedabout,searchingforthesameinsectsweweretryingtocollect(A).thesetinybirds,whichbegantheirvoy-ageinthenortheasternUnitedstates,hadflownoveropenoceanfordays.totallyspent,someindividualsunluckyatmakingtheirfirstlandfallonadesolateislandlikesombrero,wereunabletoscroungeforsparsefoodintimetofendoffexhaus-tionanddeath(b).Althoughunfortunateforthebirds,theAmeivasreadilyscav-engedthecarcasses(C).

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thepowerfulclawsontheforelimbsofAmeivasareideallysuitedforrootinganddiggingforfood.

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theauthor,withaspiratorinhand,searchesforsmallarthropodsamongtheruins.

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theotherboathadcarriedtheimmigrationofficialfromAnguillawhowastomeetusandcheckourpassports.IwasnervousasIapproachedthetopoftheladder,expectingalargeAnguillianofficialwithascowl.WhatIencounteredatthetopoftheladder,however,wasthemostnon-threateningimmigrationofficerIcouldimagine.hewasslenderandworeastrawhat,aloose-fittingbutton-upshirt,andkhakipants,andhewassweatingandsmilingandsettledonarock.Wegavehimourpassports,heverifiedthatwewerewhowesaidwewere,andbeganhislongjourneyhome.Weturnedtotakeinoursurroundings. thelandscapewasdesolate.notatreeinsight,justvineyplantscriss-crossingtherocks.overhead,abrownbooby(Sula leucogaster)wassilhou-ettedagainstthesunlitskyonwhatlookedlikearustyloadingcrane.blackAmeivaswereabundant,scurryingabitwhenwegottooclose.magnificentFrigatebirds(Fregata magnificens)hoveredinpocketsofhotair,theirredthroatsacsflappingalittleinthewind.migratorybirds,mostlyblackpollWarblers(Dendroica striata)dartedabout,searchingforthesameinsectsIwasintendingtocatch. Despiteitsrelativeisolation,humanshavehadaconsiderableimpactonsombrero.oldrustymetalstructuresjuttedupallovertheisland.Giantpitsinthelimestonepepperedtheotherwiseflatterrain,testamentto19th-centuryAmericanandbritishminingoperationsthatquarriedconsider-ablevolumesofphosphates(mineralizedguano)foruseasfertilizerinanefforttoresuscitatethesoilsofthesouthernstatesthathadbeenexhaustedbytheplantationsystem.theyevenblewupwholesectionsofthehat-shapedislanduntilonlythebrimremained(pappalardo2001).however,thiswasnottheonlytimetheislandwasthreatenedbyhumanactivities.In1999and2000,bealAerospaceoftexaswasconsideringsombreroasapotentiallaunchingsiteforrocketscarryingcommercialsatellitesintospace.triggeredbyresistancefrombirdenthusiastswhoclaimedthatsombrerowasanimportantnestingsiteformanyWestIndianspecies,bealcon-ductedanenvironmentalassessmentthatclaimedthatthenestingsitewasnotthatimportantandthatonlyonespeciesoflizardlivedontheisland—andthattherocketswouldnotaffectthem.bealultimatelydecidednotWindowsintheruinsoftheoldlighthousekeeper’squarterslookoutoverthe

desolatelandscapeofsombrerototheAtlanticoceanbeyond.

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Ravagedby19th-centuryminingactivitiesanderodedovertimebyrainandwavesofwaterviolentlywashingovertheisland,thesurfaceofsombreroisamazeofsharpandporous“dog-tooth”limestone.

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tousesombrero,notbecauseofthedestructivenatureoftheprojectbutbecausetheweatherwasnottotheirtaste(pappalardo2001). theisland’sguanodepositswereexhaustedby1890,leaving“dog-tooth”limestonethatissharpandporous,erodedovertimebyrainandwavesofwaterviolentlywashingovertheisland.IwasveryawareofhowexposedmyfeetwereinChacosandals.Despitetheapparentdesolation,

Iknewtheislandwasteemingwithcreatures.“Well,wedon’thavemuchtime,”Dr.powellsaid,awakeningmefrommystateofawe.“let’sstartcollecting.”Drs.powellandperrywentoffinsearchoflizards.“Catch’emifyoucan,ladies!”Dr.perrycalled,referringtolizards,ofcourse,ashetrudgedinthedirectionofthelighthouse. theentomologyteamstartedcollecting.typically,webeatvegetationandcatchfallinginsectsinournetsbeloworsweepthroughgrasseswith

WRIGht

oneofthelargerminingpits,relictsof19th-centuryeffortstoextractmineralizedphosphates(guano)thathadaccumulatedoverthecenturiescourtesyofinnumerablegenerationsofnestingseabirds.

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Duringtheheatoftheday,nestingbrownboobies(Sula leucogaster)liftedtheirbodiesofftheireggstopreventoverheating.squawkingabitifweapproachedtooclosely,theywereotherwiseindifferenttoourpresence.

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Catchingelusivelittlegeckosintherubbleofsombrerowasnotalwayssuccessful.Inthisinstance,however,thebrokentailwillprovideDnAforgeneticstudiestodetermineiftheDwarfGeckosofsombreroare“near”Sphaerodactylus sputator,aslazell(1964)indicated.thelizardescapedandwillregenerateanewtail.

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nets—butsombrerohasnotallvegetation.so,westartedturningoverrocksandpiecesofrustymetal.Wewouldbepreparedwithouraspirators—devicesinvolvingasmallplasticjarequippedwithtubestosuckuplittlecritters.Arubbercorkinthejarhastwocurvedmetaltubesstickingouteitherside,ashortonewithascreenthatleadsintothejarthatisconnectedtoalongrubbertubethatyousuckwithandalongeronethatisplacedneartheinsectyouwant.Yousuckinandtheinsectisvacuumedintothejar(andnotintoyourmouthbecauseofthescreen).Itriedlookingunderrocksallovertheisland,butinsectswerefewandfarbetween—untilweclimbeddownintooneoftheminingpits,whereIfoundsomethingthatIhadnotseen—hugecongregationsoftinymites.perhapstheywereinthepitbecausethetemperaturewaslowerorperhapsthepocketsandcrevicesretainedalittlemoisture. After little success while flipping rocks, I started picking apartsomeofthesmallplantsand,loandbehold,Ifoundwebspinners(orderembioptera).Webspinnersareverycoolprimitiveinsectswithsilkglandsusedtospinthegalleriesinwhichtheylive.Iruthlesslysuckedthemupwithmyaspirator.Ialsolookedthroughanoldboobynest,whereIfoundseveralbeetleandflylarvae.Isuckedthoseup,too—assistedbyanAmeivawithwhomIgladlysharedthetroveofinsectlarvae. Alreadysunburnedandwithlittletimeleftontheisland,Idecidedtoexploresomeofthebuildings.thelighthouse,builtin1868afterashiphadrunintotheislandin1859,wasinruins.evenafterminingoperationswereabandoned,thebritishboardoftrademaintainedthelighthousetoguideshipsthroughtheAnegadapassage.In1960,ahurricanedestroyedthelighthouse,whichwasreplacedandhasgonethroughseveralincarna-tionssince.Itwasautomatedin2002,butleftahistoryclearlyportrayedbythedecayingruins.

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onafood-limitedisland,lizardsmustactivelysearchformeals.thisAmeivawasforagingthroughtheremainsofaboobynestinsearchoffood.

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theabundanceofdown-coveredboobychickstestifiedtothedearthofpredatorsonanisolatedisland.

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theconcretebaseoftheoldlighthouseandtheremainsofthelighthousekeeper’squartersstandside-by-sidewiththenewautomatedlighthouse.

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thissombreroAmeivahadjustexcavatedabeetlegrub.

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Despitetheirghostlyappearance,thebuildingshavebeenintegratedintotheisland’secosystem.AsIenteredwhatwasleftofoneoftheoldstaffhouses,IencounteredtwolargeAmeivas—andonehadsomethingyellowandinsect-likehangingoutofitsmouth.Icreptcloser,determinedtostealtheirsnack.thelizardsscurriedunderapileofplywoodplanks.AsIcare-fullyliftedthem,theAmeivasslippedoutadoorwaywithemptymouths,leavingayellowandbrownscorpionthatlaylifelessandstingerlessontheplank.Icollecteditimmediately. Iwanderedintoanotherbuildingandsawananoleonthewindowsillwithsomethinginitsmouth.Iwasveryconfidentnow,andedgedcloser—buttheanolewasmoreskittishthanthemorerelaxedgroundlizardsanditsprinteddownthewall,acrossthefloor,andunderalargeloosetile.

Itiptoedcloserandslowlystartedtoflipthetile.thelizardranoutbutlefthislunch,alivelycricket.Igrabbeditbeforeithoppedawayandslippeditintotheplasticbagwiththedeceasedscorpion.Ithankedthelizardsandapologizedforstealingtheirlunchesbeforeheadingbacktowheretheoth-ershadgathered. Asthelastspecimenswerebeingcollected,IdecidedthatIneededtocatchmyfirstlizard.Whatbetterplacetodoitthanonanislandliterallycrawlingwithhundredsofimperturbableblacklizards?Icrouchedlowtothegroundandwaitedpatientlyasthelizardsdrewnearer.AssoonasIreachedouttograbone,however,itwoulddashoutoftheway.Dr.perryapproachedandaskedifIwantedalittlehelp.hehandedmeanoose.“Juststickthenooseneartheheadandpullup—prettycomplicated.”Withliz-ardscrawlingaroundmelikesassylittleblackdinosaurs,Ihadonewithinaminute.Dr.perrytaughtmehowtocorrectlyholdalizardbygraspingbothlegsofonesideinthesamehand.Inamedhimbobert.mytwominutesoflizard-catchingblisscametoanabruptendaswedecidedtocallitaday. Whenthecaptainandcrewsawusheadingtotheladder,theypickedusuponebyone.thistimewehadtojumpfromtheladdertothebobbingboat.Again,IalmostfellintryingtoimpersonateIndianaJonesandbarelymadeitontotheboat.Ibeggedthecaptaintoletmetakeadiptocoolmysunburnedbodybeforeweheadedoutandassoonasheagreed,Iwasoutofmyfieldclothes,inmysuit,anddivingoffthesideoftheboat.IhaveneVeRswuminsuchclearwater.WhatIhadassumedwasonlyabout15feetdeepwas,infact,morelikefifty!Itwaslikeswimminginliquidglass. onthetriphome,IthoughtaboutwhatIhadseen,asnapshotofaverycomplexanddelicateecosystem.Iwishedthatwehadhadmoretime,achancetocollectatnight,tomakemoreobservations.I’msureawholebookcouldbewrittenabouttheecologyofsombrero.Despitethemin-ingoperationsandtheproposedrocket-launchingsite,sombrerohashungon.Forthemomentatleast,littleblacklizardsandbigbrownboobieslivelargelyundisturbedbyhumaninterlopers—andmaybethat’showitshouldstay.

AcknowledgementsKarimV.D.hodge,Director,Departmentoftheenvironment,Anguilla,facilitatedourvisittosombreroandissuedpermitsforcollectingspeci-mensandtissuesamples.theConservationAgency,withagrantfromtheFalconwoodFoundation,charteredtheboatthatcarriedustosombrero.thestaffofGuanaIslandmadearrangementsforthecharterandsupportedoureffortsinmanyways.

Literature CitedDaltry,J.C.1999.ConservationassessmentofthesombreroGroundlizard:With

notesontheotherreptilesofsombrero.Flora&FaunaInternational–UKoverseasterritoriesprogramme,Cambridge,UnitedKingdom.

Ivie,m.A.1999.InitialReport,Invertebratebiodiversitysurveyofsombrero,november1999.<http://groups.yahoo.com/group/caribbean-biodiversity/message/521>.

lazell,J.D.,Jr.1964.thereptilesofsombrero,WestIndies.Copeia1964:716–718.

pappalardo,J.2001.loveandrockets.The Dallas Observer.1march2001.

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lackingtrees,typicallythefavoredhabitatofWestIndiananoles,theanolesofsombrerolivedonrockfacesorthedeterioratingruins—theonlyverticalhabitatsavailableontheisland.thismale(A)andfemale(b)wereonpouredconcretewallsnearthelighthouse.

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52 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 IntRoDUCeDspeCIes

I n T R o D u c e D s P e c I e s

thetentacledsnake, Erpeton tentaculatumlacépède1800,isastrictlyaquaticspeciesofAsianorigin(Winokur1977),whichtypicallyoccurs

insloworstagnantbodiesoffreshandbrackishwater(hahn1973),ofteninveryturbidlakes,streams,andricefields(lovic2010).thisspeciesfeedsmostlyonfishes,althoughsmallamountsofvegetationhavealsobeendoc-umentedinthediet(hahn1973).Erpeton tentaculatumpossessesapairoffleshytentaclesontherostrum,whicharelikelyusedasbothsensorystruc-tures(Winokur1977)aswellasanaidincamouflage(smithetal.2002).thisspeciesattainsanadultlengthof50–90cmtotallength(lovic2010). on9December2010at2023h,ajuvenile(280mmsVl,305mmtotallength)Erpeton tentaculatumwasfoundjustoutsidethepropertyofananimalimporter’sfacilityat6450stirlingRoad,hollywood,browardCounty,Florida(26.04591on,80.21976oW,WGs84datum).thissnakewasfoundunderapieceofartificialhumandebrisdirectlyadjacenttoasourceofwater.theindividualappeareddead,likelybecauseofthecold(13oC)evening,butrevivedafterbeingwarmed.thesnakewasdepositedintheFloridamuseumofnaturalhistory,UniversityofFlorida(UF163086).speciesidentificationwasconfirmedbyGregoryWatkins-Colwell,Yalepeabodymuseum. theintroductionpathwayforErpeton tentaculatuminFloridaisthepettrade.theadjacentareassurroundingtheactivefacilityarewellknownforunusualnon-indigenousspeciesandfrequentlysearchedfortheirille-gallyreleasedorescapedanimals.thissnakelikelyhadescapedfromanenclosureattheimporter’sfacilityorwasintentionallyreleased,andrepre-sentsthefirstknownvoucherforthisfamilyandspeciesinFlorida.Furthersearchesbytheauthorsandcolleagueshavefailedtoturnupanymoreindi-vidualsofErpeton tentaculatum,andabreedingpopulationisnotsuspected,norlikelyduetoinsufficienthabitatinthegeneralarea.

AcknowledgmentsWetrulythankJavierborrero(AycockVeterinaryClinic)forpermissiontocollectanimalsonprivatepropertydirectlyadjacenttotheanimalimporter’sfacility;GregoryWatkins-Colwell(peabodymuseumofnaturalhistory)for

speciesconfirmation;andGadperry,Robertpowell,tomChesnes,andananonymousreviewerforprovidinghelpfulcommentsonthisresearchnote.

Literature Citedhahn,D.e.1973.CommentsontentaclestructureandfunctioninErpeton tentacu-

latum(serpentes:Colubridae).Journal of Herpetology7:304–306.

lovic,V.2010.Erpeton tentaculatum (tentaclesnake,Fishingsnake).<www.thai-landsnakes.com>.

smith,t.l.,G.David,e.povel,andK.V.Kardong.2002.predatorystrikeofthetentacledsnake(Erpeton tentaculatum). Journal of Zoology256:233–242.

Winokur,R.m.1977.theintegumentarytentaclesofthesnakeErpeton tentacula-tum:structure,function,evolution.Herpetologica33:247–253.

The Tentacled snake, Erpeton tentaculatum lacépède 1800 (Homalopsidae), in florida

JoshuaD.holbrook1andKennethl.Krysko2

1palmbeachAtlanticUniversity,Westpalmbeach,Florida33401,UsA([email protected])2Floridamuseumofnaturalhistory,UniversityofFlorida,Gainesville,Florida32611,UsA([email protected])

tentacledsnake(Erpeton tentaculatum)(UF163086),collectedon9December2010inhollywood,browardCounty,Florida.

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CubanbrownAnoles(Anolis sagrei)arenativetothebahamaIslands(Caysal,Conception,Crooked,Acklins,Grandbahama,little

bahama,RumCay,andsansalvadorislandbanks),Cubaandassociated

cays,IsladelaJuventud,andlittleCayman,includingmostsatellitesandcayswithevenrudimentaryvegetation.thespeciesalsohasbecomeestab-lishedinJamaica,eithernaturallyorthroughhumanmediation.more

cuban brown anoles (Anolis sagrei) in saint lucia

matthewn.morton1andChristopherA.Cox2

1DurrellWildlifeConservationtrust,lesAugrèsmanor,trinity,JerseyJe35bp,ChannelIslands,Greatbritain([email protected])2CaribbeanenvironmentalhealthInstitute,p.o.box1111,Castries,saintlucia,WestIndies([email protected])

photographsbynathanA.Wood(DurrellWildlifeConservationtrust)


recently introducedpopulationsareknown inGrandCayman,swanIsland,sintmaarten,Grenada,saintVincent,theGrenadines(Canouan),barbados,theAtlanticCoastofméxicoasfarasbelize,theIslasdelabahía(offhonduras),Aruba,hawaii(oahuandCoconutIsland,Kauai),taiwan,thesoutheasternUnitedstates,andCalifornia(orangeCounty;hendersonandpowell2009andreferencestherein,Fläschendräger2010).

At least some introducedpopulationsoriginated in the southeasternUnitedstates. Anindividualofthisspecieswasfirstphotographedandrecordedbyoneofus(CAC)on8June2003athisresidentialpropertyatlatoc,justwestofsaintlucia’scapitol,Castries(photographicvouchermilwaukeepublicmuseummpmp-752).thesinglemalewasfoundinastockpileofcabletelevisionwireonoldtimbersincontactwiththeground.thespeciesisnotknowntohavebeenseenagainuntilreportswerereceivedon5December2010fromasecondresidentialpropertyatlatoc,approxi-mately300mnorthwestofthefirstsightingat14°0’35”n,61°0’13.5”W.thissiteisapproximately1.3kmsouthwestofsaintlucia’smainseacargoport,portCastries.tenA. sagrei werecapturedandphotographedatthissecondsite;threeofthesephotographshavebeendepositedasmuseumvouchers(seefigure).Atthissecondsite,individualswerefoundonandnearconstructionmaterials(primarilylumber)andonfencepostsaroundasmallovergrowngardenplot.endemicA. luciaeandintroducedA. wattsi wereobservedwithinafewmetersofA. sagrei,andintroducedA. extremuswasobservedwithin100m.AsA. luciae canattainalargeradultsizethan A. sagrei (hendersonandpowell2009),negativeeffectsonsaintlucia’snativeanolemightbeminimal,assuggestedforeffectsofA. sagreionendemicA. aeneusandA. richardiionGrenada(Greeneetal.2002). Whetherthenewlyestablishedpopulationremainslargelyrestrictedtoseverelyalteredhabitats,asonGrenada(Greeneetal.2002,Germanoetal.2003),saintVincent(hendersonandpowell2005,treglia2006,tregliaetal.2008,ealesandthorpe2010),andsintmaarten(Fläschendräger2010)remainstobedetermined.theoriginofthepopulationonsaintluciaisunknown,althoughthelocationforoursightingssuggeststhattheintroductionpathwaywaslikelycargo,probablylumber,fromthesouth-ernUnitedstates,againassuggestedfortheintroductionofthisspeciestoGrenada(Greeneetal.2002).

Acknowledgementsthelocationatwhichvouchersweretakenwasfirstbroughttoouratten-tionbyGerardandniallbergasse,whokindlygaveusaccesstothesiteandhelpeduscatchthelizards,asdidbarbaraschaeffer,benjibarca,mikeball,nateWoods,nickCondie,andtwylaholland.mnmisfundedbythebalcombetrustandworksinsaintluciaalongsideDurrell’spro-gramcounterpartsinthesaintluciaForestryDepartment(ministryofAgriculture,lands,Forestry&Fisheries).

Literature Citedeales,J.andR.s.thorpe.2010.Revealingthegeographicoriginofaninvasive

lizard:theproblemofnativepopulationgeneticdiversity.Biological Invasions12:77–86.

Fläschendräger,A.2010.CubanbrownAnoles (Anolis sagrei) inst.maarten.Reptiles & Amphibians17:121-122.

Germano,J.m.,J.m.sander,R.W.henderson,andR.powell.2003.herpetofaunalcommunitiesinGrenada:Acomparisonofalteredsites,withanannotatedchecklistofGrenadianamphibiansandreptiles.Caribbean Journal of Science39:68–76.

Greene,b.t.,D.t.Yorks,J.s.parmerlee,Jr.,R.powell,andR.W.henderson.2002.Discoveryof Anolis sagreiinGrenadawithcommentsonitspotentialimpactonnativeanoles.Caribbean Journal of Science38:270–272.

henderson,R.W.andR.powell.2005.Geographicdistribution:Anolis sagrei.Herpetological Review36:467.


treglia,m.l.2006.Anannotatedchecklistoftheamphibiansandreptilesofst.Vincent,WestIndies.Iguana13:251–262.

treglia,m.l.,A.J.muensch,R.powell,andJ.s.parmerlee,Jr.2008.InvasiveAnolis sagreionst.VincentanditspotentialimpactonperchheightsofAnolis trinitatis.Caribbean Journal of Science44:251–256.

male (Aandb)and female (C)CubanbrownAnoles (Anolis sagrei) fromlatoconst.lucia.photographicvouchershavebeendepositedinthemilwaukeepublicmuseum(mpmp-753).theidentityoftheselizardsandtheindividualphotographedin2003wasconfirmedfromphotographsbyRobertW.henderson(mpm).

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In late october 2010, quarantine officers working in layou, st.Vincent,discoveredtwoCubanbrownAnoles(Anolis sagrei)andthree

GreenhouseFrogs(Eleutherodactylus planirostris)inacontainerofornamen-talplantsshippedbyplantationspiceGrowersinGoulds,miami-DadeCounty,Florida.photographicvouchersforA. sagreiandspecimensofE. planirostrisaredepositedinthemilwaukeepublicmuseum(mpm-p754andmpm33993–5,respectively).RobertpowellconfirmedtheidentityoftheanolesandKennethl.Kryskothatofthefrogsfromphotographs.RobertW.hendersonconfirmedtheidentificationofthefrogsafterexam-iningspecimens. bothspeciesarenativetoCuba,mostbahamianislands,andatleastsomeoftheCaymanIslands.bothhavebeenpresentinFlorida,theoriginofthisandpresumablymostotherintroductionsofbothspecies,foroveracentury(meshakaetal.2004).Anolis sagreihasbeendocumentedfromanumberoflocationsoutsideitsnaturaldistributionandisalreadyestablishedonst.Vincent(hendersonandpowell2005,tregliaetal.2008,ealesandthorpe2010)andelsewhereintheregion(powelletal.2011).Althougheffectsonnativeanoleshavetodatebeenlimited(tregliaetal.2008),largely

becauseoftherelativelyrestricteddistributionofA. sagreionst.Vincent,thatspecieshasexpandeditsrangetothepointwhereeradicationislikelytobeimpossible.however,augmentingestablishedpopulationsorintroducinglizardstonewsites(suchaslayou)shouldbeavoidedinordertopreventmorewidespreadcompetitiveinteractionswithnativecongeners. IntroducedpopulationsofEleutherodactylus planirostrisareestablishedonGreatInaguaIsland(bahamas),Jamaica,andtheCaicosIslands,andhavebeenintroducedtoGrenadaandpossiblyGuadeloupe,butarenotknowntobeestablishedthere(powelletal.2011).IfE. planirostisbecameestablished,itwouldprobablyinteractmostintenselywithE. johnstonei,whichisubiquitousatlowelevationsonst.Vincent,phenomenallyabun-dantinurbanareasanddisturbedhabitats,andisitselfanintroducedspe-cies(e.g.,malleryetal.2007,powellandhenderson2007).AlthoughE. planirostrisisgenerallyconsideredtobemoreterrestrialthanE. john-stonei,thelatterisquiteabundantinterrestrialsituations(underrocksanddebrisinditchesandontheground)onst.Vincent(malleryetal.2007),andfrogsinatleastsomepopulationsofE. planirostrisarerou-tinelyfoundinarborealsituations(e.g.,bromeliads;hendersonandpowell

a failed (?) Introduction of lizards and frogs on st. vincent

AmosGlasgow

ForestryDepartment,st.VincentandtheGrenadines,Campdenpark,st.Vincent

IntRoDUCeDspeCIes

thismaleCubanbrownAnole(Anolis sagrei)ispartofanestablishedpopulationinthemontroseneighborhoodinKingstown,st.VincentandtheGrenadines.Individualsshippedintolayoucouldhavefoundedanadditionalcolonyifnotfortheactionsofthequarantineofficers.

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2009).however,initsnativerangeE. planirostrisreacheselevationsthat,ifexploitedonst.Vincent,couldresultincontactwithendemicPristimantis shrevei,whichislistedasendangeredontheIUCnRedlist(hedgesandpowell2004)andalreadyisthreatenedbycompetitionwithE. johnstoneiandongoingdeclinesinhabitatqualityandextent. Individualsofeitherspeciesareunlikelytohaveescapeddetection,buttheorganizationthatreceivedtheplants(natureCaresVG)hasbeeninstructedbyquarantineofficerstomonitorthemcloselywhilestillon-siteandafteranysalesandrelocations.

AcknowledgementsthequarantineofficersatlayouaretobecommendedfortheirdiligenceindiscoveringtheseinvasiveanimalsandforbringingthemtotheattentionoftheForestryDepartment.IthankRobertW.henderson,Kennethl.Krysko,andRobertpowellforidentifyingspecimens.

Literature Citedeales,J.andR.s.thorpe.2010.Revealingthegeographicoriginofaninvasive

lizard:theproblemofnativepopulationgeneticdiversity.Biological Invasions12:77–86.

hedges,b.andR.powell.2004.Pristimantisshrevei.In:IUCn2010.IUCnRedlistofthreatenedspecies.<www.iucnredlist.org>.

henderson,R.W.andR.powell.2005.Geographicdistribution:Anolis sagrei.Herpetological Review36:467.


mallery, C.s., Jr., m.A. marcum, R. powell, J.s. parmerlee, Jr., and R.W.henderson.2007.herpetofaunalcommunitiesoftheleewardslopesandcoastsofst.Vincent:Acomparisonofsitesvariouslyalteredbyhumanactiv-ity.Applied Herpetology4:313–325.

meshaka,W.e.,Jr.,b.p.butterfield,andJ.b.hauge.2004.The Exotic Amphibians and Reptiles of Florida.KriegerpublishingCo.,malabar,Florida.

powell,R.andR.W.henderson.2007.thest.Vincent(lesserAntilles)herpeto-fauna:Conservationconcerns.Applied Herpetology4:295–312.

powell,R.,R.W.henderson,m.C.Farmer,m.breuil,A.C.echternacht,G.vanbuurt,C.m.Romagosa,andG.perry.2011.IntroducedamphibiansandreptilesintheGreaterCaribbean:patternsandconservationimplications,pp.63–143.In:A.hailey,b.s.Wilson,andJ.A.horrocks(eds.),Conservation of Caribbean Island Herpetofaunas.Volume1.brill,leiden,thenetherlands.

treglia,m.l.,A.J.muensch,R.powell,andJ.s.parmerlee,Jr.2008.InvasiveAnolis sagreionst.VincentanditspotentialimpactonperchheightsofAnolis trinitatis.Caribbean Journal of Science44:251–256.

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theseGreenhouseFrogs(Eleutherodactylus planirostris)werediscoveredinaship-mentofornamentalplantsfromFlorida.

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thepacificCoastGiantmuskturtle,Staurotypus salviniiGray1864,isindigenoustothepacificlowlandsofoaxacaandChiapas,mexico,

rangingeastwardintoelsalvadorandGuatemala(ernstandbarbour1989).thisspeciesfeedsonfishes,invertebrates,andamphibians,andlivesinslow-moving,softbottomlakes,rivers,andotherwaterwayswithaquaticvegetation(ernstandbarbour1989). on2october2010at1400h,anadult(270mmcarapacelength[Cl]) femaleStaurotypus salvinii (Fig. 1; photographic voucherUF160342)wascollectedon thepropertyofZoomiami,miami-DadeCounty,Florida,alongtheperimeterfenceofahoof-stockexhibitadja-cent toa0.85-km-longman-made lake (25.60276°n,80.40123°W,WGs84datum).thisS. salviniiwasplaced in the zoo’squarantinefacilityforafutureexhibit,whereitwaskeptina300-gallonenclosure.on26october2010,theanimalwasradiographedbecauseofunusualbehavioranddiscoveredtocontaintwocalcifiedeggs(Fig.2).on15november2010,thisS. salviniiovipositedthetwoeggsinsubstrateof

sandandleaflitterintheenclosure.theseeggs(10.1g,39.5x20.8mm;9.9g,39.5x19.7mm,respectively)wereremovedandplacedonver-miculitesubstrateandincubatedwithinaGQF1550hatcherIncubator(savannah,Georgia)at28.6°Cwithahumidityofapproximately80%.on2December2010,thetwoeggsappearedtohavefailedtodevelopandwerediscarded. on1november2010at1210h,ajuvenile(107mmCl)femaleStaurotypus salvinii(Fig.3;photographicvoucherUF162276)wascollectedcrossingazooserviceroadbetweenexhibits(25.61079°n,80.40068°W),0.9kmnorthofthefirstindividual.thenearestwatersourcetothisloca-tionisa0.13-km-longman-madelake,whichisnotconnectedtothelarger,0.85-kmlakewherethefirstindividualwasfound.thisS. salviniiwasalsoplacedinthezoo’squarantinefacilityforafutureexhibit,whereitwaskeptinaseparateenclosure. Zoomiamicontainsapproximately300haofpinerockland,approxi-mately133haofwhicharedevelopedfortheinstitution.Althoughmuch

The Pacific coast giant Musk Turtle, Staurotypus salvinii gray 1864 (Kinosternidae),

a new non-indigenous species in floridaDustinC.smith1,Kennethl.Krysko2,tracyA.sorensen1,andmarian.sider1

1Zoomiami,12400sW152ndstreet,miami,Florida33177,UsA([email protected],[email protected],[email protected])2Floridamuseumofnaturalhistory,UniversityofFlorida,Gainesville,Florida32611,UsA([email protected])

photographsbytheseniorauthor.


ofeasternmiami-DadeCountyconsistsofurbandevelopment,propertiesdirectlyadjacenttothezoocompriselargetractsofmostlyundevelopedpinerockland.noneofthesepinerocklandareasareresidential,andforsecuritythezooisborderedbya2.5-m-tallchain-linkfenceandconcreteperimetermoat.Staurotypus salviniihasneverbeenkeptaspartofthezoo’scollection,butisavailablethroughthepettrade,whichislikelytheinva-sionpathway.Althoughthisspecieswasprobablyintentionallyreleasedonthegroundsbyazoovisitor,weacknowledgethatitalsocouldhavebeenreleasedalongperimeterpropertiesbecauseofgapsundergatesandotherpotentiallycompromisedareas.likeinothermiami-DadeCountyparks(e.g.,seeKryskoetal.2010),numerousnon-indigenousamphibianandreptilianspeciesfrequentlyarefoundonornearzoogrounds;thesewerelikelyreleasedbyzoovisitorsand/orpreviousownerswhonolongerwanttocarefortheiranimals.thepresenceofmultipleS. salvinii,includingafemalewitheggsandajuvenilefoundatdifferentsitesonzoogrounds,sug-gestslikelyreproductionandestablishmentofthisspeciesthere.however,beforemakingthisclaimwebelieveevidenceofreproductioninthewildisnecessary.thisisthefirstknowndocumentedintroductionofthisspeciesinFlorida.

AcknowledgmentsWethankJohnb.Iverson,peterA.meylan,paulVanderschouwandJasonbourqueforspeciesconfirmation;J.steveConners,nicoleAtteberry,Gadperry,Robertpowell,andananonymousreviewerforhelpfulcommentsonthispaper.

Literature Citedernst,C.h.andR.W.barbour.1989.Turtles of the World.smithsonianInstitution

press,Washington,D.C.andlondon.

Krysko,K.l.,K.m.enge,e.m.Donlan,e.A.Golden,J.p.burgess,andK.W.larson. 2010. the non-marine herpetofauna of Key biscayne, Florida.Herpetological Conservation and Biology5:132–142.

IntRoDUCeDspeCIes

Fig. 1.Adultfemale(UF160342)Giantmuskturtle,Staurotypus salvinii,frommiami,miami-DadeCounty,Florida.

Fig. 2.Radiographoffemale(UF160342)Giantmuskturtle,Staurotypus salvinii,frommiami,miami-DadeCounty,Florida.

Fig. 3.Juvenilefemale(UF162276)Giantmuskturtle,Staurotypus salvinii,frommiami,miami-DadeCounty,Florida.


Conserving Hellbendershellbenders (Cryptobranchus alleganiensis) areobligateaquaticsalamandersthatareindeclineduetohabitatlossanddisease.twosubspecieshave been described based on morphologicalcharacteristics:C. a. alleganiensis (easternsub-species)andC. a. bishopi(ozarkhellbender).Currentconservationstrategiesincludecaptivepropagationforrestorativereleaseseventhoughinformation regarding the current levels ofgeneticvariabilityandstructurewithinpopula-tionsisnotsufficienttoeffectivelyplanforcon-servationofthegeneticdiversityofthespecies.toinvestigatepatternsofpopulationstructureinthehellbender,Crowhurst et al.(2011.Conservation Genetics,DoI:10.1007/s10592-

010-0170-0, published online 28 December2010)genotyped276hellbenders fromeightmissouriRiverdrainages,representingbothsub-species.theirresultsshowedlowlevelsofwithin-drainagediversitybutstrongpopulationstructureamongrivers,andthreedistinctgeneticclusters.FSTvaluesrangedfrom0.00to0.61andaveraged0.40.theresultsconfirmedpreviousreportsthatC. a. bishopiandC. a. alleganiensisaregeneticallydistinct,butalsorevealedanequidistantrelation-shipbetweentwogroupswithinC. a. bishopiandallpopulationsofC. a. alleganiensis.Currentsub-speciesdelineationsdonotaccuratelyincorporategeneticstructure,andforconservationpurposes,thesethreegroupsshouldbeconsideredevolu-tionarilysignificantunits.

ConseRVAtIonReseARChRepoRts

c o n s e R v a T I o n R e s e a R c H R e P o R T s

Hurricanes and Amphibian Communities in Coastal Wetlands

Isolatedwetlands in the southeasternUnitedstatesaredynamichabitatssubjecttofluctuat-ingenvironmentalconditions.Wetlandslocatednearmarineenvironmentsaresubjecttoaltera-tions inwaterchemistryduetostormsurgesduringhurricanes.GunzburGer et al.(2010.Wetlands Ecological Management18:651–663)evaluatedtheeffectofstormsurgeoverwashonwetlandamphibiancommunities.theysam-pled32wetlandsinnorthwesternFloridaovera45-monthperiodtoassessamphibianspeciesrichnessandwaterchemistry.Duringthisstudy,sevenwetlandswereoverwashedbystormsurgefromhurricaneDennis,whichmadelandfallon10July2005intheFloridapanhandle.specificconductanceacrossallwetlandswas lowpre-storm(<100υs/cm),butincreasedpost-stormattheoverwashedwetlands(mean=7,613υs/cm).Increasedspecificconductancewasstronglycor-relatedwithincreasesinchlorideconcentrations.Amphibianspeciesrichnessshowednocorrela-

tion with specific conductance. one monthpost-stormtheauthorsobservedslightlyfewerspecies in overwashed compared with non-overwashedwetlands,but this trenddidnotcontinuein2006.morespeciesweredetectedacrossallwetlandspre-storm,butnodifferencewasdetectedbetweenoverwashedandnon-over-washedwetlandswhenconsideringallamphib-ianspeciesoradultanuransandlarvalanuransseparately.Amphibianspeciesrichnessdidnotappeartobecorrelatedwithphorthepresenceof fish, although the amphibian communitycompositiondifferedbetweenwetlandswithandwithoutfish.Resultssuggestthatamphib-iancommunitiesinwetlandsinthesoutheast-ernUnitedstatesadjacenttomarinehabitatsareresistanttotheeffectsofstormsurgeoverwash.

Site Fidelity and Extended Activity in Crawfish Frogs

Crawfish Frogs (Lithobates areolatus) basetheirnonbreedingactivitiesinandaroundtheentrancesofcrayfishburrows.thissiteprefer-enceallowsmonitoringofindividualsusingstillandvideoimagingtechniques.hoffman et al.(2010.BioScience60:829–834)usedthreecam-eratypesofferingdifferentcontinuities,scales,andresolutionsofdatatoobserveactivitypat-ternsandnonbreedingbehaviors.theauthorsobservedtwobehaviorsnotpreviouslyreportedforamphibians:(1)Circumdielactivitypatterns,and(2)longperiods(days)ofsurfaceactivity.AlthoughthesebehavioralfindingsareatthistimespecifictoCrawfishFrogs,theymightnotbeunusualactivitypatternsforotherfrogs.theuseofimagingtechniquesthattakeadvantageofthisspecies’dependenceonburrowsanduseofburrowentrancesfacilitatedtheobservationofthesepatternsforthefirsttime.

eastern hellbenders (Cryptobranchus alleganiensis alleganiensis)aregeneticallydistinctfromtwogroupswithinwhathasbeen traditionallyconsidered theozarkhellbender (C. a. bishopi).All three repre-sentevolutionarilysignificantunitsthatshouldbeaddressedseparatelybyeffortstoconservethem.

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newlyemployed imagingtechniquesrevealedcir-cumdielactivityandlongperiodsofsurfaceactivityinCrawfishFrogs(Lithobates areolatus).WhethertheseactivitypatternsareuniquetoCrawfishFrogsisatthistimeunknown.

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speciesnotdetectedpost-storm fromoverwashedwetlandsbutpresent innon-overwashedwetlandswereHyla squirella,Pseudacris ornata(top),Lithobates(Rana)catesbeianus,Lithobates(Rana)clamitans(mid-dle),andPseudobranchus striatus(bottom).twoofthesespecies,L. clamitansandL. catesbeianus,werealsonotdetectedattheoverwashedwetlandspriortothestorm.

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n a T u R a l H I s T o R y R e s e a R c H R e P o R T s

58 IRCFReptIles&AmphIbIAns•Vol18,no1•mAR2011 neWsbRIeFs

Iguana Conservation in Hispaniola and Fiji

the endemic and Critically endangeredRicord’sIguana(Cyclura ricordii),isrestrictedtosouth-centralhispaniola,wherefourdisjunctsubpopulationsremaininanarealessthan100km2.three subpopulationsoccurwithin theJaragua-bahoruco-enriquillobiosphereReserveintheDominicanRepublic(DR).thefourth,thesmallest,andmostthreatenedsubpopulationisfoundinsoutheasternhaiti. Conservationactivities in theDRfocusonhabitatmonitoringandregularpatrollingbymembersoflocalsupportgroupstopreventdisturbances. this integrative, low-cost pro-gramiseffective,raisesawareness,andservesasamodelforotherendangeredspeciesprogramsintheDR.Inseptember,withfundingfromananonymousdonor,theInternationalIguanaFoundation(IIF)andGrupoJaraguapurchased20hectaresintheDRencompassingacriticallyimportant nesting area for Ricord’s IguanascalledFondadelatierranearpedernales.over140hatchednestsweredocumentedinthisareaearlierthisyear.InAnse-a-pitres,haiti,wherethe most fragile subpopulation of Ricord’sIguana exists, the IIF and Grupo Jaraguarecentlyconductedaworkshopforlocalauthor-itieswithaproposal to create amunicipallyprotectedhabitat.Ifsuccessful,thisprotectedareawouldbethefirstofitskindinhaiti.theIIF,UsFWs,macArthurFoundation,Disney,andsociedadespañoladeornitologíasupportworkonRicord’sIguana.

InFiji,twosignificantfindingsemergedfromfieldworkconductedover thepastyearbythenationaltrustofFiji,Universityofthesouthpacific,FijiDepartmentofenvironment,natureFiji-mareqetiViti,tarongaZoo,andtheUsGeologicalsurvey.thefirstfindingwasthediscoveryofsizeablepopulationsofthenewlydescribedFijianbandedIguana(Brachylophus bulabula)ontwogovernment-ownedislands,mokogaiandmakodrago.thesepopulationswillhopefullyserveasthefocusofanassessmentforapotentialnewnationalparkwithinFiji. thesecondfindingwasconfirmationthatGreenIguanas(Iguana iguana),illegallyintro-ducedtoFijiearlierthiscentury,nowoccuronatleastfourislandswithdocumentedbreedingontwoislands.GreenIguanasarehighlyinva-sive,andjuvenilesareverysimilarincolorandshapetothethreatenedendemiciguanasofFiji,asimilarityexpectedtohampercontrolefforts.partialfundingforfieldworkinFijicamefromCepFandtheIIF.

Glenn Gerber and Miguel García Co-Chairs,IguanaspecialistGroup

WithcontributionsfromernstRupp,masaniAccimé,andRobertFisher

IUCnSpecies52:21–22

Bringing Down the Serpent KingAbroken lockonasuitcasemovingthroughKualalumpurInternationalAirportthissum-merledtothediscoveryofnearly100babyboaConstrictors,twovipers,andasouthAmericanturtle.thiswasafairlymodestcacheforawild-lifesmuggler,butthemanwhoclaimedthesuit-casewasnoordinarycriminal.hewasAnsonWongKengliang,theworld’smostnotoriouswildlifetrafficker—and,insteadofaslaponthewrist,whichhemightreasonablyhaveexpected,Wongwasabouttoreceiveasurprisingpunish-ment. Fromthetinymalaysianislandofpenang,inastorefrontnolargerthanyouraveragenailsalon,Wongcommandedoneof theworld’slargest wildlife trafficking syndicates. muchoftheworkofWong’scompany,sungaiRusaWildlife,wasabove-board.helegallywhole-saledtensofthousandsofwildreptilesannu-ally,makinghimthelikelysourceformanyofthesnakes,lizards,turtles,andfrogsonsaleinAmericanpetstores.however,usingaprivatezooasacover,healsoofferedanastoundingarrayofcontraband,includingsnowleopardpelts, panda skins, rhino horns, rare birds,andKomodoDragons.hesmuggledcriticallyendangered wildlife from Australia, China,madagascar,newZealand,southAmerica,andelsewheretomarketslargelyineurope,Japan,andtheUnitedstates.

Wong’slongcareerbeyondthereachofthelawoffersawindowonthe$10–20billion-a-yearillegalwildlifetradeandourbrokensystemtocombatit.Underfundedlawenforcement,governmentcorruption,controversy-shynGos,andafeebleinternationallegalframeworkhaveyieldedfewinroadsagainstwildlifesyndicatesorkingpinslikeAnsonWong.Wong’sarrestandhissentencinginnovember2010provideales-sononhowtochangethat. Wonggot into thebusiness in theearly1980s,sellingexoticanimalstozoosanddeal-ersaroundtheworld.Changesininternationalandmalaysianlaweventuallyledhimtofocusonreptiles,whichhebelievedwerenotaspro-tectedasotherspecies.Wong’stechniquesmir-roredthoseofnarcoticsandothertraffickers.hepaidmulestocarryKomodoDragonshiddeninsidesuitcases,andhidendangeredmalagasytortoisesatthebottomoflegalwildlifeship-ments.purchasingvacationpackagesascover,hesentmenouttopoachrarewildlifefrombreed-ingfacilitiesinnewZealand.themostimpor-tanttechniqueWongandotherlarge-scalesmug-glersemploy,however,isfarlessexoticthanallthat.Instead,ithastodowithpaperwork. the primary treaty governing interna-tionalwildlife trade is theU.n.ConventiononInternationaltradeinendangeredspeciesofWildFaunaandFlora(CItes),whichclas-sifieswildlifeintothreegroups:AnimalslistedinAppendixI,suchastigersandgorillas,aresoclosetodisappearingthattheyarebannedfrominternational commercial trade;Appendix IIanimalsmaybetradedunderapermitsystem;andAppendixIIIanimalsareprotectedbyacountrywitharequest thatothershonortheprotection.smugglersscantheglobeforcoun-trieswithweaklawsorcorruptlawenforcementofficialstaskedwithstampingtheiranimals’doc-umentation,paperthatisasmuchindemandastheanimalsthemselves.suchcountriesbecomewildlifelaunderingpass-throughpoints—ani-malscomeinillegallyandleave“legally.” Fewplaceslaunderasmuchillegalwildlifeaspenang.thiswaslargelytheworkofWong:“Icangetanythingherefromanywhere,”heboasted toanAmericanundercoveragent inmarch1997.“nothingcanbedonetome.Icouldsellapanda—and,nothing.AslongasI’mhere,I’msafe.”thekey,heexplained,waspayingoffgovernmentofficials. Wong’sactivitiesfinallylandedhimontheradarofinternationallawenforcementagenciesintheearly1990s,whenspecialoperations,theeliteundercoverunitoftheU.s.FishandWildlifeservice(UsFWs),madehimthetargetofaninvestigationcalledoperationChameleon.Agentssetupareptileimportingcompanyout-side san Francisco and a retail operation in

n e W s b R I e f s

the smallest and most threatened population ofRicord’sIguana(Cyclura ricordii;top)isinsoutheast-ernhaiti.thediscoveryofFijianbandedIguanas(Brachylophus bulabula;bottom)ontwogovernment-ownedislandsmightservetofacilitateestablishmentofanewnationalpark.

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Reno,nevada,andbegandoingbusinesswithWong.beforelong,theydiscoveredWongnotonlysmuggledrareandendangeredreptiles,butalsocriticallyendangeredbirdsandmammals.hisreachwasglobal. toarrestWong,agentsneededarusetolurehimoutofmalaysia.Alucrativeinterna-tionalblackmarket inbearbile is fueledbydemandasacure-allintraditionalAsianmedi-cine.UsFWsspecialAgentGeorgemorrison,actingundercover,offeredWongapieceofabear-bilesmugglingoperationheclaimedtoberunning,ononecondition:thetwomenhadtomeetinperson.Wongagreed,butbecausehewasalreadywantedintheUnitedstatesonsmugglingcharges,hewouldonlyagreetomeetinmexico. WhenWongsteppedoffaJapanAirlinesflightinmexicoCityon18september1998,he was met by morrison, along with specialAssistantU.s.AttorneyRoberts.Andersonandateamofmexicanfederales,whoarrestedhim.ItwastheculminatingmomentofoperationChameleon,whichhadgrownintooneofthelongestandmostsuccessfulundercoveropera-tionseverundertakenbytheUsFWs,andoneinvolvingauthorities in fourcountries.WongfoughthisextraditionfrommexicototheUnitedstatesfortwoyears,buteventuallygavein. In June 2001, Wong was sentencedin California to 71 months in prison, fined$60,000, andbanned fromexporting to theUnitedstatesforthreeyearsafterhisrelease—butthesentencedidnotstophim.Whilehewasinprison,hiswiferanhiswildlifebusiness,includingsalestotheUnitedstates.Whenhegotoutin2003,Wongreturnedtomalaysia,grewaponytail,andwentbacktowork. Wong’sU.s.convictionhadnodiscern-ibleimpactonhisabilitytooperateinmalaysia.tothecontrary,hisnewplantobuildatigerzooreceivedfundingandlandfromthepenanggovernment. misliah mohamad basir, thewildlife department official directly respon-sibleforpolicingWong,consideredhimtobealegitimatebusinessman,andbelievedtheU.s.authoritieshadframedhim. IntheyearssincetheUsFWs’srevolution-arystingoperationtookdownWong,theglobalwildlife kingpin had grown more powerful,whilethepeoplewhobroughthimtojusticehadfallenonhardtimes.specialoperationsfailedtomakeanothermajorcaseafterWong’s;today,itsbestagentshavegivenupundercoverwork—andtheunit,whichneverconstitutedmorethanahandfulofagents,isallbutdefunct.thingsdidn’tbegin to changeuntil January2010,whenNational GeographicpublishedaprofileofWong,detailinghisgovernmentcon-nectionandhisnewplanstoexploittigers.theoutcrybyboth thepublic and journalists inthemalaysianpresswasimmediate.malaysiannewspapersandtelevisionarestate-controlled,

whichmakesitdifficultforjournaliststocriti-cizethegovernmentdirectly—buttheyarefreetodisclose foreignreportingaboutmalaysia.Inthecourseofthepastyear,theministryofnaturalResourcesandenvironmentannouncedarevampofitswildlifedepartment,promisingto rotate senior officers every three years. Itstrippedthedepartmentofkeypowersandisintheprocessoftransferringmisliah,whoisnowalsounderinvestigationbythemalaysianAnti-CorruptionCommission.Whileinternationalwildlife nGos were cautious about causingtroubleinmalaysia,theyhaveprovidedinvalu-ableadvicetothecountry’sgovernment,includ-ingtheparliament,whichpassedthefirstover-haulofitswildlifelawinnearlyfourdecades. Asaresult,whenWongwascaughtwithasuitcaseofboaConstrictors,hedidn’tgetawaywithit.themalaysiangovernmentrevokedhisbusinesslicenses,shutdownhiszoo,andseizedhisentirecollectionofanimals, includinghisbengaltigers.Innovember,ajudgesentencedhimtofiveyearsinprison,anunprecedentedtermforawildlifetraffickerinmalaysia. the effort to catch Wong offers a fewimportant lessonsonwhat it takes to stopakingpin.twoprinciples float to the surface.First,where long-term,high-volumeinterna-tionalwildlife traffickingexists,oneormoregovernmentofficersareeithercomplicitinthesmugglingor socomplacentas tobe reason-ablyconsideredaccomplices.Aslongasafewcountriesarewillingtobendtherulesandfudgesomepaperwork,itdoesn’treallymatterwhateveryoneelsedoes:Asinglecountry,evenasin-glewildlifeenforcementofficial,canunderminetheentireglobal“system”tocontroltrafficking.

second,thepublicinthekingpin’shomecountryisthebestweaponagainsthim.nosteptomalaysia’sunprecedentedlegalandadmin-istrativereformsthisyearwasmoreimportantthan theoutcry inmalaysia fromconcernedcitizens.Dozensofarticles—manyof themonmalaysiannewspapers’frontpages—finallytoldthestoryofoperationChameleon,Wong’spenangoperations,andthehistoryofpoorman-agementbythecountry’swildlifedepartment,exposingyearsofbadpolicyandofficialvenality. exposureisacriticalingredientforchange.lawenforcement,nGos,andotherswillfindtheirworkmagnifiedandlastingoncethepublicbecomesawareofit.Fullstoriesneedtobetoldinthemedia.IntheUnitedstates,wherewild-lifetraffickingbustsareoftentreatedashumor-ousnewsitems,thatmeansjournalistshavetorealizethatcriminalsyndicatesareoftenbehindthosepeoplestoppedatairportswithexoticani-malshiddenundertheirclothes. ofcourse,nofixisforever.Wildlifesmug-glers, likeanyotherbreedof trafficker,obeythe lawsofsupplyanddemand.As longasamarketforrareandendangeredanimalsexists,someonewillsupplythem.RisingincomesinChina, India, and even in southeasternAsiameanmorecustomersforendangeredwildlife.In2009,over18,000 liveanimalsandmorethan267tonsofdeadanimalsandderivativeproductswereseizedinlawenforcementactionsinsoutheasternAsiaalone—andthatappetitewon’tgoawayjustbecauseWongtemporarilydid.Whethermalaysia’sreformswilltakerootremainstobeseen,asiswhatwillhappenuponWong’srelease.still,somewhereinmalaysiaoranothercountrysomeoneisallbutguaranteed

neWsbRIeFs

ChineseWaterDragons(Physignathus cocincinus),suchasthisjuvenile,escapedfromthecompoundofAnsonWongKengliangonpenangIslandandhavebecomeestablishedinnearbywaterwayswheretheyappeartobedisplacingsomeofthenativeriparianspeciesintheagamidgenusGonocephalus.

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tobewillingtolooktheotherway,andaspiringkingpinsareeagertotakeoverhisbusiness.

Bryan Christy Adaptedfromanarticle.

28December2010

Snake Owners See a Furry Bias in Invasive Species Proposal

manysnake lovershavebeenseethingat theAmericangovernmentsinceearlylastyear,whenitsoughttobantheimportationandinterstatetransportationofninespeciesofforeignsnakes.thefederalFishandWildlifeservicesaidtheanimals,iffreed,posedaseriousrisktonativeecosystemsacrossthesouthernUnitedstates. WhentheFishandWildlifeservicemovedtobantradeinthesnakes,whichincludeboasandspeciesofanacondasandpythons,itarguedthattheymetthelegalcriteriaforbeingbothinjuriousandinvasive.Invasivespecies—fromAsiancarp,whichthreatentheGreatlakes,toZebramussels,whichspreadexponentially—areaseriousenvironmentalconcern,onethatisoftennotdealtwithuntilaspecieshasbecomefirmlyestablished.theFishandWildlifeservicearguesthat,inthecaseofthesnakes,theyaretryingtogetaheadoftheproblem. It is the first time the government hastried to list animals so widely held as pets.RoughlyonemillionAmericansarebelievedtoownsnakesofthetypes listedbytheInteriorDepartment,accordingtotheU.s.AssociationofReptileKeepers,and31,000wereimportedin2008,themostrecentyearforwhichthegovern-menthasdata.tradeinthesespeciesisbigbusi-ness—morethan$100millionannually.thosewithrarecolorscanfetchupwardof$75,000. themove toban the snakeshas setoffaswellofangeramongaggrievedsnakeown-ers andbreeders,whohave themost to losefinancially,aswellasasmatteringofacademicherpetologists,zookeepers,andrepresentatives

of international conservation groups. Whenthe regulations came up for public review,theyfloodedthegovernmentwithobjections.Attheheartoftheirargumentsisacritiqueofthe emerging science of invasive species riskassessment,andtheirresponsehashighlightedthechallengesthatthegovernmentfacesasitincreasinglymovestoprotectnativefloraandfaunanotjustfromcurrentinvasivespeciesbutalsofromfuturethreats. thereptilekeepersgroup,whichclaims12,000 professional breeders and sellers asmembers,hasfiledformalobjectionswiththeInteriorDepartmentandisthreateningalawsuitbasedonwhatitsaysisthegovernment’spoorscientificevidence.AndrewWyatt,thepresidentoftheassociation,arguesthatthegovernmentisnowpromotinganative-species-onlyagendafavored by environmental groups. “this hasimplicationsforeveryanimalinterestoutthere,rightdowntofamilypets,”hesaid,addingthatbysuchstandards,“allamphibiansareinjuriousandcatsanddogscan’tbefarbehind.” thebattlegoesbackto2006,whenthesouth Florida Water management DistrictpetitionedtheFishandWildlifeservicetolistthesnakesunderthelaceyAct,whichwouldmake it a crime to transport them into theUnitedstatesoracrossstateborders.burmesepythons—somethoughttobedumpedbypetownersandsomethatescaped—wereestablish-ingthemselvesacrosstheeverglades,wheretheywereswallowingupeverythingfromendangeredKeylargoWoodRatstoalligators.thepopula-tionhasbeenexpandingnorthwardatroughlythreeandahalftosixmilesayear—Indy500speedsinreptilianterms. Inrecentyears,Floridaofficialshadtakensignificantstepstolimitownershipofinvasivesnakeswithinthestatebutstillwantedmoretobedone.WhatwastoprotecttheevergladesfromasnakeboughtinGeorgiaandcarriedacrossstatelines?tobanthesnakesunderfederallaw,thegovernmentwouldhavetoshowthattheyposedathreattonativeplants,crops,oranimals.WithverylittlescienceavailableabouthowreptilesthatcomefromdistantplaceslikesubtropicalAsiaandAfricamightfareinAmerica,theU.s.Geologicalsurveywasaskedtoassesstherisk. theagencylookedatmanyfactors,includ-ingthedamageintheeverglades.Italsoturnedtoacomputermodeltodeterminewhatpartsofthecountrymighthaveahospitableclimateforthespecies.thescientistslookedatvariablesincludingmeanmonthlytemperaturesandrain-fallatawiderangeofelevationsinthenativehabitatsoftheanimalsandmatchedthemtopatternsintheUnitedstates.

theyestimatedthatsuitableclimatesfortheburmesepythoninparticularmightincludethe11southernmoststatesfromCaliforniatonorthCarolina.Ifglobalwarmingcontinuedapace,thegeologicalsurveyadded,thesnakesmightevenbeathomeinnewYorkCityby2100.thenationalnewsmediagavegleefulattentiontotheprospectofasnakeinvasion—butsoonafter,biologistsattheCityUniversityofnewYorkdid theirownmodeling,usingmorefactorsanddifferentoneslikeprecipita-tionduringthewettestperiodsoftheyear,andcameupwithonlyFloridaandsoutherntexasaspossiblehabitatsforthesnakes.IndependentstudiesofsnakescapturedintheevergladesandtakennorthtoGainesville,Florida,andsouthCarolinafoundthatmostoftheanimalsdiedwhen leftoutside inwinter in those regions.thesefindingswerefurtherbolsteredwhenanunusualcoldsnapintheevergladeslastJanuaryleftalargenumberofburmesepythonsdeadoncanalbanksandlevees. thestudieshavefiredupthesnakeindus-try,whichseesthemasproofthatthegovern-ment is pursuing a hostile and unwarrantedagenda.onebreedersaidthatthegovernmentregulations,whichdonotpreventbreedingandowningbutdoprevent transportationacrossstatelines,wouldruinhisbusinessandthou-sandslikeit.“thereptileindustrywouldsufferacrushingblowoversomethingthatdoesnotmakesense,”hesaid. Dr.elliottJacobson,aprofessorofveteri-narymedicineattheUniversityofFloridaandaspecialistinreptiles,alsoseesthegovernment’sscienceasskewed.hesaidhesuspectedthatthegovernmentwaslesssympathetictoreptilesthantomorecuddlycreatures.“theimpactofferalcats,forexample,onwildlifeismuchgreaterthanwhat theburmesepythonscando,”hesaid,notingthatacateatsmuchmorethanasnakeofthesamesize. however, thomas strickland, assistantsecretaryfor fishandwildlifeandparks, saidthat the government was not going to backdownand that itwould approve the regula-tions by next summer. the science is solid,mr.stricklandsaid,andthegeologicalsurveywillsoonpublishapeer-reviewedanswertoitscritics.likeother invasivespecies,snakesarearealandgrowingproblem,hesaid.“Youarenotdealingwithhamstershere,”hesaid.“Iwasdownintheeverglades,andittookfourpeopletoholda19-footburmesepython.thesethingswreakhavoc.”

leslieKaufmanNew York Times 8January2011

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theU.s.Geologicalsurveyestimatedthatsuitableclimatesfortheburmesepython(Python bivittatus)mightincludethe11southernmostU.s.statesfromCaliforniatonorthCarolina.Ifglobalwarmingcon-tinuedapace,thegeologicalsurveyadded,thesnakesmightevenbeathomeinnewYorkCityby2100.

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NEWPRODUCT

editorsRobert Powell

Executive Editor AvilaUniversity,KansasCity,mo

AJ Gutman Editor

bloomfield,Ct

Gad Perry Associate Editor

texastechUniversity,lubbock,tX

Lilianna Gonzalez Assistant Editor

texastechUniversity,lubbock,tX

Jim McDougalAssistant Editor

Guilford,Ct

Michael Ripca Art Director

Atco,nJ

John Binns Graphics/Photography

InternationalReptileConservationFoundation,tucson,AZ

Sandy Binns Services Coordinator

InternationalReptileConservationFoundation,tucson,AZ

editorial boardAllison C. Alberts

ZoologicalsocietyofsanDiego

Frederic J. BurtonblueIguanaRecoveryprogram

Arthur C. EchternachtUniversityoftennessee

L. Lee GrismerlasierraUniversity

Robert W. Hendersonmilwaukeepublicmuseum

John B. IversonearlhamCollege

Charles R. KnappZoologicalsocietyofsanDiego

Gunther Köhlersenckenbergmuseum

Kenneth L. KryskoFloridastatemuseumof

naturalhistory

Jan RamerIndianapolisZoo

Thomas WiewandtWildhorizons

Copyright©2011bytheInternationalReptileConservationFoundation,Inc.Allrightsreserved.nopartofthisjournalmaybereproducedinanyformorbyanyelectronicormechanicalmeans,includ-inginformationstorageandretrievalsystems,withoutthepublisher’swrittenpermission.IRCF, Reptiles & Amphibians, Conservation and Natural History (Issn1098-6324)ispublishedquarterlybytheInternationalReptileConservationFoundation,anonprofit,tax-exemptorganization,pobox90270,tucson,AZ85752.periodicalpostagepaidattucson,AZ.

sTaTeMenT of PuRPoseThe International Reptile Conservation Foundation works to conserve reptiles and amphibians and the natural habitats and ecosystems that support them.

theInternationalReptileConservationFoundation,Inc.isanon-profit501(c)(3)Californiacorporation.

MeMbeRsHIP InfoRMaTIonIRCF, REPTILES & AMPHIBIANS, CONSERVATION AND NATURAL HISTORY, the Journal of theInternationalReptileConservationFoundation,isdistributedquarterly.

AnnualRates:Individual U.S. Membership ........................................................................................................... $25.00Individual Membership, Digital (Adobe PDF)*.................................................................................$25.00Institutional U.S. Subscription...........................................................................................................$30.00International Membership (including Canada)..................................................................................$55.00International Institutional Subscription.............................................................................................$60.00

Additionalcurrentandbackissuesareavailable.Visitwww.ircf.org/bissuetoorderhardcopiesand/orpdffiles.membersreceiveadiscountonpurchasesofadditionalissues.

*theAdobepDFisoptimizedforwebpublishinganddoesnotprovidethequalityandresolutionofthearchivalprintedversion,especiallynoticeableinphotographsandcomplexgraphics.

JoIn onlIne: www.IRcf.org

MeMbeRsHIP [email protected],ortollfree1-877-472-3674(U.s.only),orwriteto:IRCF,pobox90270,tucson,AZ85752

solIcITaTIonstheIRCFencouragescontributionofarticles,letterstotheeditor,newsitems,andannouncementsforpublicationinREPTILES & AMPHIBIANS.Generalarticlescandealwithanyaspectofreptilianoramphibianbiology,includ-ingconservation,behavior,ecology,physiology,systematics,orhusbandry.submissionofphotographstoaccompanyarticlesisencouraged.Guidelinesforsubmissionsandfiguresareatwww.ircf.org/about/submissions.manuscriptsmaybesubmittedviae-mail([email protected]).Authorsofonepageormoreofprintwillreceiveafreecopyofthejournalinwhichtheircontributionappears,andwillreceiveapDFfileoftheirarticlefordistribution.

DonaTIonsForanydonations,pleaseincludeyourname,address,phonenumber,ande-mailaddress.

aDveRTIsIng PolIcyWeadvertiseonlynon-livingproducts(exceptfeederinsects).Foradvertisingratesandoptionscontactsandybinns,AdvertisingDirector,[email protected],pobox90270,tucson,AZ85752.

2011hasbeendesignatedtheYearoftheturtle.thedesignationrecognizesthesadstateofmanyoftheworld’scheloniansandtheurgentneedtohelpmanyspecies.Ifyouhaven’talready,weencourageyoutochecktheFocus

onConservation(p.64)andbecomeinvolvedinsomecapacityinthe2011Year-of-the-turtleeffortstoconservethesespecialcreatures. onaseparatebutnolessurgentissue,theIntroducedspeciessectioninthis(beginningonp.52)andrecentissuesshowsthatherpetologicalintroductionsareongoing.Documentingthemisessentialforidentifyingthemecha-nismsandoutcomesoftheseeventsandprovidinginformationthatregulatorscanusetoaddressandmoderatetheproblem.thesituationisofparticularconcernonmanyislands,whereendemicspeciesarethreatenedbycompeti-tivelysuperiorcontinentalinvasives.AsCharlesDarwinwroteinThe Origin of Species by means of Natural Selection or, The Preservation of Favoured Races in the Struggle for Life(1872.6thed.):“Asnaturalselectionactsbycompetition,itadaptstheinhabitantsofeachcountryonlyinrelationtothedegreeofperfectionoftheirassociates;sothatweneedfeelnosurpriseattheinhabitantsofanyonecountry,althoughontheordinaryviewsupposedtohavebeenspeciallycreatedandadaptedforthatcountry,beingbeatenandsupplantedbythenaturalisedproductionsfromanotherland.”Weencouragetheresponsibleownershipofamphibiansandreptiles.Ifyoukeepnon-nativespecies,pleasetakeeveryprecautionyourself,andsharewithothersyourconcernsovertheescapeorreleaseofanimalstoplaceswheretheydonotbelong.

theeditorsofReptiles & Amphibians

Editors’ Remarks


turtlesaredisappearingfromtheplanetfasterthananyothergroupofanimals.today,nearly50%ofturtlespeciesareidentifiedasthreat-

enedwithextinction.however,it’snottoolate.theUnitedstateshasmoreendemicturtlespeciesthananywhereonearthandisatrueturtlebiodiversityhotspot.Carefulstewardshipcanpreservetherarespeciesandkeepcommonspeciescommon. thebadnewsisthathumanscausethegreatestharmtoturtlepopu-lations—butthegoodnewsisthatwehavethepowertomakepositivechangestowardturtlesurvival.thegreatestthreatstoturtlepopulationsinclude:(1)habitatlossanddegradation;(2)over-harvestingofwildturtlesforfood,traditionalmedicines,andpets;(3)mortalityfromroads,agricul-turalmachinery,fishingby-catch,andpredators(manyofwhicharenon-native);(4)exoticinvasivespeciesanddiseases;(5)lossofgeneticallyuniquepopulationsduetohybridization(oftenasaconsequenceofturtlesreleasedintoareaswheretheydonotoccurnaturally);and(5)climatechange. throughouttheyear,turtleconservationgroupsinpartnershipwithpartnersinAmphibianandReptileConservation(pARC)willberaisingaware-

nessoftheissuessurroundingturtlesthroughpressreleases,newsletters,photocontests,andrelatedevents.Citizens,naturalresourcemanagers,scientists,andthepet,petfood,andrelatedindustriescanworktogethertoaddressissuesandtohelpensurethelong-termsurvivalofturtlespeciesandpopulations. bookmarkwww.yearoftheturtle.organdcheckitfrequentlyformoreinformationonhowyoucangetinvolved!thewebsiteincludeslinksto2011stateoftheturtleandUsAturtlemappingproject,informationonsubmittingaphotographfortheYear-of-the-turtlephotocontest,andthe2011Yearoftheturtlevideo.Youcanalsosignupformonthlynewslet-tersthatcontain:(1)Adownloadableturtlephotocalendarforeachmonth(ifyouenterthephotocontest,yourphotocouldbeinthecalendar!);(2)informationaboutturtleconservationeffortsandgroups,andhowyoucanhelp;(3)interviewswithturtleexperts,andanswerstosubmittedques-tions;(4)informationonhowyoucanhelpspreadthewordaboutturtles;(5)educationalmaterials;(6)turtleart,poetry,andculturalinformation;andmuchmore!Contactyearoftheturtle2011@gmail.comwithanyques-tions,ortobeapartnerinYearoftheturtle.

2011year of the Turtle

f o c u s o n c o n s e R v a T I o n

thespottedturtle(Clemmys guttata)isendemictotheeasternUnitedstates.ItiscurrentlylistedasVulnerableontheIUCnRedlist.

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Documents

IRCF Reptiles & AmphibiAns...IRCF ReptIles & AmphIbIAns • Vol 18, no 1 • mAR 2011 3 basking in the alligator snapping Turtle, Macrochelys temminckii (Testudines: chelydridae) John