IRC

FReptiles & AmphibiAns

C 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

t2JUN

2010

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sPeCial issue, Part ii: a tribute to Henry s. Fitch

IRCF ReptIles & AmphIbIAns • Vol 17, no 2 • JUn 2010 In thIs IssUe

A heated conflict between two male Grass Frogs (Rana temporaria) and one male Common toad (Bufo bufo). Clutches of frog eggs are visible in the water. see related note on p. 82.

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When Red-sided Garter snakes (Thamnophis sirtalis parietalis) in northerly climes hibernate, hundreds of them aggregate in the same hibernaculum. During the winter, this apparently serves to prevent them from freezing and they also have access to each other for breeding in the spring.

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An adult Red milk snake (Lampropeltis triangulum syspila) constricting and swallowing a large Woodland Vole (Microtus pinetorum). see related note on p. 94.

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back Cover: Jay Fleming

Yellow-bellied sea snakes (Pelamis platurus) are the only sea snake in the eastern pacific ocean, and the only species that is present on the pacific Coast of Central America. see related article on p. 69.

Front Cover: Vipul Ramanuj

A King Cobra (Ophiophagus hannah) eating a hump-nosed pit Viper (Hypnale hynale) at the Agumbe Rainforest Research station in the Western Ghats of India. see related article on p. 95.

IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010 65tAbleoFContents 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

IRC

F

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

special Issue: a Tribute to Henry s. fitch, Part II

R E M E M B R A N C E

What’sinaname?.......................................................................................................................................Marian L. Griffey 66

A R t i C l E s A N d N o t E s

newperspectivesontheecologyandnaturalhistoryoftheYellow-belliedseasnake(Pelamis platurus)inCostaRica:DoesprecipitationInfluenceDistribution?...................................................................................................................... ...............................Harvey B. Lillywhite, Alejandro Solórzano, Coleman M. Sheehy III, Spencer Ingley, and Mahmood Sasa 69

observationsandCommentsontheDietofthemany-bandedKrait(Bungarus multicinctus multicinctus)intaiwan............ ..................................................................... Jean-Jay Mao, Gerrut Norval, Chih-Liang Hsu, Wei-Han Chen, and Ray Ger 73

blocked-flightAggressivebehaviorinsnakes..................................................................................................D. Bruce Means 76

terrestrialburrowinginnestingsoftshellturtles(Apalone mutica and A. spinifera)... Michael V. Plummer and J. Sean Doody 79

ApparentCompetitionbyGrassFrogs(Rana temporaria)andCommontoads(Bufo bufo)forbreedingsites.....Ivan Vergner 82

submergedCallingbyoregonspottedFrogs(Rana pretiosa)RemotefrombreedingAggregations................ Jay Bowerman 84

suburbanizationofaCentraltexasherpetofauna................................................................................ Frederick R. Gehlbach 87

Redmilksnaketakinglargepreylateintheseason.................................................................................. Stephen L. Barten 94

observationsonaWildKingCobra(Ophiophagus hannah),withemphasisonForagingandDiet........................................ ...................................... Dhiraj Bhaisare, Vipul Ramanuj, P. Gowri Shankar, M. Vittala, Matt Goode, and Rom Whitaker 95

t R A v E l o g u E

thebackdoorturtlesoftortuguero....................................................................................................................... Don Moll 103

F E A t u R E A R t i C l E

turtlesintheDust:effectsofhands-onscientifictrainingonaGroupofbehaviorallyAt-riskstudents’Knowledgeandempathy..................................................................................................... J. Alan Sosa, Oscar Reyes, and Gad Perry 108

C o M M E N t A R y

IsWildlifemanagementbusinessorConservation—AQuestionofIdeology..................................................Jesus A. Rivas 112

i N v A s i v E s P E C i E s

IslandInvaders:InvasiveReptilesandAmphibiansintheturksandCaicosIslands.R. Graham Reynolds and Matthew L. Niemiller 116CubanbrownAnoles(Anolissagrei)inst.maarten.................................................................................. Axel Fläschendräger 121KnightAnoles(Anolis equestris)onGrandCaymanIsland...........Mat DaCosta-Cottam, Kristan D. Godbeer, and Tim Austin 122

B o o K R E v i E W

Timber Rattlesnakes in Vermont and New York: Biology, History, and the Fate of an Endangered SpeciesbyJonFurman........... ..................................................................................................................................................................George R. Pisani 123

v nAtURAlhIstoRYReseARChRepoRts:summariesofpublishedReportsonnaturalhistory......................................124v neWsbRIeFs................................................................................................................................................................................125v eDItoRIAlInFoRmAtIon.....................................................................................................................................................127

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evenwhenvisitorswerepresent,Dr.Fitchwasmeticulousaboutrecordingdata.

IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010 67RemembRAnCe

nearly14yearshavepassedsinceIfirstheardthename“Dr.henryFitch,”invariablyspokenintonesofnear-reverence.thename,often

abbreviatedtosimply“Fitch,”isalwaysspokensoftly,andfollowedbythoughtful,staring-at-your-shoessilence—evenmoresosincehisdeath.hisabsenceintheherpetologicalcommunityissorelyfelt,evenbythosewhoknewhimonlyvicariouslythroughthelegacyofhiswork.Forthoseluckyenoughtohavehadacloserrelationshipwiththeman,thescientist,(dareIsay)theLegend,hislossleavesanindeliblewoundontheheart. Whenfirstintroducedtothename,Iwasonlya“spouseof,”partofagroupofperhapsfour,maybesixherpetologistsandwildlifestudents,and

brandnewtothisworldofamphibian-and-reptilelegend,lore,science,andpioneers.likethetipofaniceberg,thetoneandsilence,thatinward-turningfocusandgentlesmilethataccompaniedthename“Fitch,”foretoldofamanwhoIsensedmustbeagiantamongthesemenofscience. WheneverIasked:“WhoisthisDr.henryFitch?”theanswercamebackunvaried:“You’lljusthavetomeethim!”luckily,Idid… Forthisoccasion,Ihadagainjoinedmyhusband,someofhiscol-leagues,andnumerousstudentsforafieldtripwithintheUniversityofKansasFitchnaturalhistoryReservation1.Westoodinjitteryclumps,shufflingourfeetandstaringoffintothegreeningwoods.Wemadehalt-ingconversationwithoneanotheraswewaitedforourguidetoarrive. mymentalimageofaniceberg-sizedGreekheroassumeditstruehumanform.Anaged,frail-looking,stooped-backmancamegingerlytowardus,leaningonhiswell-usedwalkingstick.hestoodinabeamofmorningsunlight,likeanancientturtlebaskinghimselfintowakefulnessandoptimummobility.mycompanionsandIeasedbackabit,yieldinghimasmuchofthesun’senergyaspossible.Yetweremainedcloseenoughthatoursightandhearingcouldstillperceiveeveryparticleofhim. herewasFitch,atlast!thelegend.Afraillittleman? nevertheless,allthatIhadbeenledtobelieveprovedtrue.meetinghimthatday,withnoexchangeofwordsbetweenus,toldmesomethingofthemysterysurroundingthatparticular toneofvoice,thatairofinnercontemplation and silent reverence that surrounded anymentionofthename“Fitch.”Cumulative,spontaneoushonorandrespectrippledthroughandaroundourlittlegrouplikeaninvisiblewave. Idon’trememberwhatDr.Fitchsaidtousthatmorningbeforewesetoffonourhikethroughhisbelovedwoods.myears,Ithink,wentdeafinthemidstofmarvelouswonder.Withthesunbacklightinghisthinbodyandhiswhitehairaglow,dozensofbutterfliessuddenlyappeared,lightingonhisshirt,athisfeet,orhoveringaroundhim.Asheturnedtogo,hedidsowithawarenessofthistenderentourage,movinggentlyandcarefullysoasnottoharmorfrightenhislittlefriends.onceinmotion,however,hiswingedshrouddissipatedasquicklyasithadarrived.WeloadedintotwovansandheadeddeeperintotheReservation. Dr.Fitchwasthefirsttodisembark.hewastednotimeshepherdingpeople,countingheads,oractingthetourguide.oncehisfeettouchedtheearth,hemoved—andhemovedlikeGandhi,stridinginalong-practicedrhythmthattookhimalmostoutofsightbeforewefollowerscouldassembleourthoughts.Acrossthemeadow,upthehill,throughthebriarpatch,overthecrest,intothemixed-oakwoodlands,andskimmingthetalusandlime-stoneboulders,hetraveledwithalltheeaseofafieldmouseonfamiliarterrain. Awkwardly,westraggledintothesmallclearingbeneathacopseoftreeswhereDr.Fitchstoodsmiling,patientlywaiting,observingwithsomeamusementeachperson’sbreathlessarrival.Wegatheredaroundhim,anxioustoquellourhuffingandpuffingandmissnothingofwhatthesoft-spokenmanhadtosay.Wetotteredonthesharp-edgedbouldersthatringedthetinyclearing,jockeyingforasomewhatlevelplacetostandinanefforttodisguiseourmere-humancondition.

What’s in a name?marianl.Griffey

5222nW56thCourt,Gainesville,Florida32653

photographsbyC.KennethDodd,Jr.

1theUniversityofKansasrenameditsnaturalhistoryReservationin1986tohonorhenryFitch.

R e m e m b R a n c e

Apermanentarrayofdriftfencesandtrapsledtomanycapturesandrecapturesduringthe50-yearstudyoftheReservation’ssnakesandlizards.Checkingthetrapswasadailyritual.

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“this,”Fitchsaidwhenallhadarrived,“isagoodplaceforCopperheads.”severaleyesrolleddownwardtowardfeetnowfrozenandprecariouslyperchedatopcountlesspocketswhereslenderCopperheadsmighteasilycoil.ImentallycalculatedthedifficultyIhadingettingdownhilltothisscenicglen,andthelikelihoodofsurefailureingettingmynow-wearied,oxygen-deprived,flatlander’shideup-n-outintheeventthatsuchabeastielayundermyboulder. “here’sone,”Fitchannouncedcalmly.hiseyeswerefocusedonasmallishbouldernearhisownfeet.“I’llgetit,”heoffered,slowlyleaningtowardtheground.onethinarmstretchedoutinvitinglytowardthesnakeasheleanedhisbentframeeverclosertomotherearthandeden’sscourge,hishand,arm,andfacecomingincrementallynearertohisdeadlygoal. Anaudible,collectivegasprosefromthecircleofon-lookers.“Dr.Fitch,”someonesqueaked,“letoneofusgetit.” “no,”repliedtheunperturbedleaningman.“I’llgetit.” Andsohedid.Ascalmlyandgentlyasamotherwithanewborn,FitchmadehisacquaintancewiththeCopperheadbytherock,pickeditupwithouttheuseofhookorstaff,andrevealedhisprize,facebeamingwithpurejoy.hehelditfirmlybehindtheheadinawell-practiced,strong,andconfidenthand.Weallbreathedagain,butthefrownsofconcernforhissafetydidnotrelax. “oh,”hesaid,andmyheartskippedabeat(asI’msuremanyothersdid),fearingthathehadbeenbitten.but,no,hisexclamationhadsimplybeenoneofmildself-admonishment.“Iforgotmysnakebag,”hesaid.“I’llhavetotakethisoldgirlbacktothevantogetherstats.” Copperheadsecurelyinhandandwithoutfurtheradieu,Dr.Fitch set toagainwithhisGandhi-stride.skimmingoverthetalus,heledusanothermerrychase…uptheslope,beneaththetrees,throughthebriars,overthecrest,downacrossthemeadow.bythetimeweallhadreassembledontheroadbythetwovans,Dr.Fitchhadfoundhissnakebagandhadhalfthestatsrecorded.“Done,”heannounced,beamingatusallwithhissmile.“I’lltakeherbacknow.”

“letoneofus,”someoneoffered,perhapsconcernedforthethinlegsthatwouldhavetotraversetherouteback…andlikelyCopperheadcous-inscamouflagedinthedappledlightbeneaththetrees. “I’lldoit,”Fitchstatedgently,butwithafirmnessthatbroachednodiscussion.“sheandIareoldfriends,”headded,inthatsametoneofrever-enceforthesnakewithwhichI’dheardhisownnamespoken.Andoffhewent,acrossthemeadow,overthecrestintothebriars,andoutofsight.Afewheartbeatslater,hishoaryheadagaintoppedthehorizon.hismissionaccomplished,Isawhimpausebrieflyonthecrestamidstthebriars,hisfaceturnedskywardwithalookoflovethatencompassedthewholeearth.Dr.henryFitch.thenameechoesinmymemorywithatoneofhonorandakindoflovethatIcan’texplain.WordscannotdescribewhatI(anddoubtlesssomanyothers)continuetofeelfortheman,thescientist,and(Yes!)thelegend.everybutterflyremindsmeofthatgrandadventureintheKansaswoodsonasummerdayin1996.thoseluckyenoughtohavemethimunderstandmyinabilitytodescribehim.noonecanputhenryFitchintomerewords;hehadtobeexperienced. thosewhowillonlymeethimthroughreadinghisbiography,hislifetimeofscientificwritings,orothers’memoirswillnotbegintounder-standevenhalfthemanbehindthename—butthosewhotakethetimetohearthesubtextwithinthestoriesofthisremarkablegiantamongmenwilldoubtlessbeblessedfortheeffort.theyandfuturegenerationsmayonlymakeatip-of-the-iceberg,second-handacquaintancewiththemarveloushenryFitch,butohwhatadiscoverythatwillbe!thatsortofexperiencewillstillyourheart,turnyourthoughtsinward,andbroadenyourvisiontoencompassthewholeearth.Itwillmakeyousmileintothesky,callallthelittlelivingthings“oldfriend,”andcauseyoutostareatyourowntwofeetinsilentwonder,hopingthatyoutoocanlearntowalkwithaGandhi-strideandmakeaFitch-differenceinaworldmostlyunseenandun-experiencedbythe“civilized”masses. Couldtherebeanybetterwaytohonorthemanthantowalkinhislegendaryfootsteps?

An“oldfriend.”Dr.FitchcametoknowmanyoftheresidentsoftheFitchnaturalhistoryReservation.

Dr.FitchmeasuringaCommonGartersnake(Thamnophis sirtalis).

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new Perspectives on the ecology and natural History of the Yellow-bellied sea snake

(Pelamis platurus) in costa Rica: Does Precipitation Influence Distribution?

harveyb.lillywhite1,Alejandrosolórzano2,Colemanm.sheehyIII3,spencerIngley1,andmahmoodsasa4

1Departmentofbiology,UniversityofFlorida,Gainesville,Florida32611UsA2serpentarionacionalandmuseumofZoology,UniversityofCostaRica,sanJose,CostaRica

3Departmentofbiology,Universityoftexas,Arlington,texas76019UsA4InstitutoClodomiropicado,UniversityofCostaRica,sanJose,CostaRica

“...narrativeaccountsofrarelyobservedeventsinspecies’lifehistoriessometimeshavemuchmeritandvalue.evenasingleobservationmayconstituteavaluablecontributionandmaybeabreak-throughinunderstandingthespecies’ecology.”

Henry S. Fitch, 1987

morethan60 speciesof sea snakeshave evolvedand radiated tooccupytropicaloceanichabitats.themajorityofthese(50+spe-

cies)areentirelymarineanddonotintentionallyleaveseawater(heatwole1999).nonetheless,fewerthan2.5%ofallsnakespeciesliveinmarineenvironments,andtheevolutionarytransitionfromterrestrialtomarine

habitatsisconsideredtobeparticularlydifficult(lillywhiteetal.2008a).osmoregulatorychallengesareparamount,andallmarinespeciesofsnakes(indeedallmarinereptiles)haveevolvedsaltglandstoassistwithelimina-tionofexcesssaltthatisingestedwithdietorincidentaldrinking.Waterbalanceisarelatedbutdifferentissue,andthebodywaterthatislostpas-

Fig. 1. Pelamis platurus onaCostaRicanbeach.thesesnakesaresometimesstrandedonbeachesduetostrongoffshorewindsthatcausethesnakestodrifttowardtheshore.

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sivelytotheseaorinfecesandsecretionsmustbereplenishedeitherfromfreshwatersourcesorindirectlyfromfoodorseawater.theindirectsourcesinvolvefurtherloadingofsaltsand/ormetabolitesthatmustbeeliminatedwithfurtherlossesofsomewater. Recently,lillywhiteetal.(2008b)demonstratedthatthreespeciesofseaKraits(Laticauda spp.)requirefreshdrinkingwatertoreplacebodywaterdeficits,asthesesnakesdehydrateinseawaterinspiteofthefactthattheypossesssaltglands.evidently,inthesespecies,saltglandsareimportanttoionbalancebutareinsufficienttomaintainwaterbalanceinmarineenvi-ronments.basedoncurrentliterature,representativespeciesbelongingtofouroutoffivemajorlineagesofmarinesnakeshavebeenshowntorequirefreshwater(orverydilutebrackishwater)forthemaintenanceofnormalwaterbalance(lillywhiteetal.2008b).therefore,changesinpatternsofprecipitationrelatedtoglobalwarmingmayhaveimportantimplicationsforthesurvivalanddistributionofmarinesnakes. Whetherso-called“trueseasnakes,”whicharefullymarinespecies,requirefreshwaterforwaterbalanceremainstobedetermined.Wearecurrentlyinvestigatingthisquestion,andwehavesomeevidenceforafresh-waterrequirementintheYellow-belliedseasnake,Pelamis platurus (Figs.1&2).basedonlaboratoryinvestigationsofwaterandsodiumfluxesinthisspecies,DunsonandRobinson(1976)foundthatfastingPelamis dehydrateinseawater(−0.4%bodymassperday),andtheydocumenteddrinkingoffreshwaterwhenitwasofferedtodehydratedsnakes.snakeskeptinseawa-tersurviveforlongperiodsiffedonfreshwaterfish(DunsonandRobinson1976;As,pers.obs.),butwhetherthesesnakescansurviveforlongperiodsifkeptinfullseawaterandfedonmarinefishremainsunclear.

yellow-bellied sea snakes (Pelamis platurus)thisspeciesofseasnakehasthebroadestdistributionofanysnake.ItrangesfromthecoastalwatersofeasternAfricaalongthesouthernAsiancoaststoJapan,southwardandeastwardtoAustraliaandislandsofthewesternpacific,thenceeastwardtotheAmericas.Itistheonlyseasnakethathasreachedtheeasternpacificocean,andistheonlyspeciesthatispresentonthepacificCoastofCentralAmerica.thissnakeisentirelymarine,givesbirthtolivingyoungatsea,andispelagicinhabits.Itisaspeciesofsmalltomoderatesize,reachingamaximumlengthofabout1m(solórzano2004).Pelamis hasbeenregardedasa“passivesurfacedrifter”thatisoftenwaftedbycurrentstolatitudesatwhichitisnotabreedingresident(Dunsonandehlert1971).becauseofitscosmopolitandistribu-tionandpelagichabits,thisspeciesofseasnakeisofcentralimportancetotestingthehypothesisthatfreshwaterisrequiredforphysiologicalwaterbalanceinmarinehabitats. observationsofothersandourowncollectingexperiencesinCostaRicaindicatethatPelamis arefoundingreatestnumbersinassociationwithoceanic“slicks”(Kropach1971,1975;Dunsonandehlert1971).slicksaresmoothglassystreaksformingdriftlinesintheoceanresultingfromtheaccumulationoffoam,floatingplantparts,andotherdebris(flotsam;Fig.3).theyoccurinconvergencezoneswhereorganicmaterialaccumulates,flattensthewater,andcreateszonesofconcentratedbiologicalactivity.slicksmaycontaincontaminatefilmsoforganicoilsprobablyderivedfromdiatoms,andtheybecomediscerniblebecauseoftheirdampingeffectonsmallwavelets.theconvergencezonesarecreatedbyavarietyofphysicalprocessesandcurrentssuchasinternalwavesandeddies(ewing1950).slicksarebiologicallyimportantbecauseplanktonbecomeconcentratedintheconvergencezonesandcreategoodplacesforzooplanktonandfishestofindfood.Indeed,slickshavebeenregardedasbiologicalcommunities.theycanalsobeharmfultosealife,however,becausetheyalsoconcen-tratenoxiousdebrisandpollutants.slickscanberelativelysmallandshort-lived,buttheycanalsoformdriftlinesextendingforhundredsofmetersorkilometers.theyarelikelyagentsfavoringdispersalofseasnakes,buttheirdynamicsinthiscontextareverypoorlyunderstood.Weproposethreehypothesesforwhyseasnakesassociatewithoceanicslicks.First,averypopularnotionintheliteratureisthatslicksprovideimprovedfeeding

opportunitiesduetooccurrenceoffishesseekingshelterbeneaththefloat-ingdebris(Kropach1975,heatwole1999).Asecondhypothesisisthatconvergenceofwatersconcentratesfloatingseasnakesjustasitdoesfloat-ingdebris,whichwouldbeapurelypassivephenomenon.Insupportofthisidea,othershaveconjecturedthatPelamis probablydoesnotactivelyseekoutslicks(Dunsonandehlert1971).Athirdhypothesisiscamouflage,fortheseasnakesresemblesomeofthefloatingpartsofplantsbothincolorandinform,oftenmimickingfloatingstickstowhichfishareattracted(hunterandmitchell1968).thiscouldreducepredationonseasnakesaswellasfacilitatetheirfeedingopportunities.Kropach(1975)alsosuggestedthatgatheringatslicksincreasedmatingopportunitiesforseasnakes,butthiswouldseemdependentononeoftheotherreasonsfortheprimaryassociationofsnakeswithslicks.

the Coastal distribution of Pelamis in Costa RicaYellow-belliedseasnakesaregenerallydistributedalongtheentirepacificcoastofCostaRica,andareusuallyfoundwithin1–20kmofthecoast(Voris1983,solórzano2004).especiallydensepopulationshavebeennotedatseverallocationsinrelativelycloseproximitytothecoastline.theseincludeGolfodepapagayoandassociatedbays(bahíaCulebraandseveralplayasimmediatelytoitssouth),Caboblancoatthesouthendofthenicoyapeninsula,watersbetweenbahíadeCoronadoandthenorthernendoftheosapeninsula(puntasanJose),andwithintheGolfoDulcenearthesouthernendofthecountry.thesepopulationsareknowntousfromtheexperiencesofoneofus(As)orfromliterature.otherdensepopulationsalmostcertainlyoccurelsewhere,buthavenotbeenbroughttoourattention. Wearepresentlyexploringthehypothesisthatthecoastaldistribu-tionofPelamis ispartlydependentontheavailabilityoffreshwater.thiswas suggested for theeuryhalinemarine snakeAcrochordus granulatus (lillywhiteandellis1994)andwasdemonstratedtobethecasewherespeciesofLaticauda,or“seaKraits,”havebeenstudiedonalocalscaleatorchidIsland,taiwan(lillywhiteetal.2008b).threespeciesofLaticauda areabundantatsiteswithaccesstoafreshwaterspringorestuary,whereasfewornosnakesarefoundatstrictlyseawatersites.moreover,theabun-danceofthesesnakesalsovarieswithrainfall,andfewersnakesarefoundduringperiodsofdroughtorlowrainfall.Asdiscussedelsewhere(lillywhiteetal.2008b),Laticauda spp.exhibitapatchydistributionthatcorrelateswithlow-salinitytropicalwatersinthewesternpacificandIndianoceans.Also,apreliminaryassessmentofseasnakedistributionintropicalsouthAsiaandtheIndo-Australianregionrevealsthatthespeciesdiversityofallseasnakespeciescorrelateswithmeanannualrainfall(lillywhiteetal.2008b). severalsourcesoffreshwaterarepotentiallyavailabletoseasnakes.Inadditiontoprecipitation,theseincludesprings,estuaries,ortemporaryoutflowsfromthecoast.heavyrainfallformsatemporaryfreshwaterlensattheoceansurfaceduetothedensitydifferencefromseawater,andsuchlensesaretheonlysourceoffreshdrinkingwaterintheopenseaawayfromtheimmediatecoastline.thefactthatPelamisinCostaRicacanbefoundnearertothecoastduringthedryseason(As,unpubl.obs.)ispotentiallyrelevant;andinmexico,evenmorepronouncedseasonalchangesindis-tributionareevident(hlandCs,unpubl.obs.;seealsoDuellman1961).thesesnakesmightbemovingtocloserassociationswithcoastalestuariesduringperiodicdroughts,basedonknowledgeofsiteswithwhichwearefamiliarfromourownfieldwork.Pelamis areoccasionallyreportedfromestuaries(shuntov1966).Indeed,variousspeciesofseasnakesareobservedinestuaries,andtheysometimesmoveconsiderabledistancesintorivers(Dunson1975).Alternatively,Pelamis might appeartobemoreabundantnearerthecoastduetoprevailingstrongerwindsduringthedryseason,whichcouldwashthesesnakesontotheshoreattimes(solórzano2004).numerousreputableobservershavedescribedstrandingsofPelamis onbeaches,usuallyinassociationwithstorms(Dunsonandehlert1971).slicksalsomightformingreaternumbernearthecoastalbaysduringthe

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Fig. 2. Pelamis platurus illustratingsomeofthevariationofcolorpattern,whichvariesfrommostlyblacktoblackandwhiteoryellowtoallyellow.

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dryseason.thesehypothesesremaintobetestedwiththealternativeexpla-nationsinmind.

Conclusionsomemarinesnakes,includingcertainseasnakes,havebeenshowntobedependentonaccesstofreshwaterforwaterbalanceandoccuringreaterabundancenearfreshwatersources(lillywhiteetal.2008b).Wedonotknowwhetherthemajorityofseasnakespecies,whichareentirelyaquatic,

alsorequirefreshwater.physiologicalstudiessuggestthatthepelagicspe-ciesPelamis platurus dehydratesinseawaterandwilldrinkfreshwaterinthelaboratory(DunsonandRobinson1976).thus,onemightinferthatPelamis requiresfreshwaterforwaterbalance,butthisaspectofitsphysiol-ogyremainstobeproperlytestedanddocumented.themovementsofthispelagicspeciesarelargelyunknown,butsomefeaturesofitsdistributioninCostaRicasuggestthatitmightassociatemorecloselywithcoastalsourcesoffreshwater,suchasestuaries,duringtimesofperiodicdrought.thepotentialinfluenceoffreshwateronmovementsofthisspeciesmightaffectmetapopulationdynamicsinimportantwaysthatremaintobeexplored.Wearecurrentlyinvestigatingthephysiology,distribution,andgeneticstructureofPelamis populationsinthesecontexts.Knowledgeofthewaterrequirementsofseasnakes—andPelamis inparticular—willenhanceourunderstandingofthedistributionofthesemarinereptilesandhowtheymightrespondtofuturechangesinthedistribution,intensity,andperiod-icityofprecipitation.

AcknowledgementstheresearchonwhichthisarticleisbasedwassupportedbythenationalGeographicsociety,Committee forResearch andexploration (grant#8058-06tohbl), thenationalscienceFoundation(grant#Ios–0926802tohbl),andagrantfromtheexplorersClub(tosI).WearegratefultoAdánbarreraforassistancewithboattransportationandlocatingofseasnakesinCostaRica.

Literature CitedDuellman,W.e. 1961.the amphibians and reptiles ofmichoacán,mexico.

University of Kansas Publications of the Museum of Natural History 15:1–148.

Dunson,W.A.1975.Adaptationsofseasnakes,pp.3–19.In:W.A.Dunson(ed.),The Biology of Sea Snakes.Universityparkpress,baltimore.

Dunson,W.A.andG.W.ehlert.1971.effectsoftemperature,salinity,andsur-facewater flowondistributionof the sea snakePelamis.Limnology and Oceanography 16:845–853.

Dunson,W.A.andG.D.Robinson.1976.seasnakeskin:permeabletowaterbutnottosodium.Journal of Comparative Physiology b108:303–311.

ewing,G.1950.slicks,surfacefilmsandinternalwaves.Journal of Marine Research 9:161–187.

Fitch,h.s.1987.thesinofanecdotalwriting.Herpetological Review 18:68.

heatwole,h.1999.Sea Snakes. KriegerpublishingCo.,malabar,Florida.

hunter,J.R.andC.t.mitchell.1968.Fieldexperimentsontheattractionofpelagicfishtofloatingobjects.Journal du Conseil (Conseil international pour l’Exploration de la Mer)31:427–434.

Kropach,C.1971.seasnake(Pelamis platurus)aggregationsonslicksinpanama.Herpetologica 27:131–135.

Kropach,C.1975.theYellow-belliedseasnake,Pelamis,intheeasternpacific,pp.185–213.In:W.Dunson(ed.),The Biology of Sea Snakes.Universityparkpress,baltimore.

lillywhite,h.b.andt.m.ellis.1994.ecophysiologicalaspectsofthecoastal-estu-arinedistributionofacrochordidsnakes.Estuaries 17:53–61.

lillywhite,h.b.,C.m.sheehyIII,andF.ZaidanIII.2008a.pitviperscavengingattheintertidalzone:Anevolutionaryscenarioforinvasionofthesea.BioScience 58:947–955.

lillywhite,h.b.,l.s.babonis,C.m.sheehyIII,andm-Ctu.2008b.seasnakes(Laticauda spp.)requirefreshdrinkingwater:Implicationsforthedistribu-tionandpersistenceofpopulations.Physiological and Biochemical Zoology 81:785–796.

shuntov,V.p.1966.DistributionofseasnakesinthesouthChinaseaandeastIndianocean.Zoolicheskij Zhurnal 45:1882–1886.

solórzano,A.2004.Snakes of Costa Rica. Institutonacionaldebiodiversidad(Inbio),CostaRica.

Voris,h.K.1983.Pelamis platurus, pp.411–412. In:D.h.Janzen(ed.),Costa Rican Natural History.theUniversityofChicagopress,Chicagoandlondon.

Fig. 3.slicksseennearplayadelCoco,CostaRica.(A)Anarrowslickidentifiedbyfoamydriftline.(b)Flotsamanddebrisassociatedwithaslick.(C)Abroaderslickwithoutdebris;notethecalmwaterinthecenterofthephotograph.

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observations and comments on the Diet of the many-banded Krait

(Bungarus multicinctus multicinctus) in TaiwanJean-Jaymao1,Gerrutnorval2,Chih-lianghsu1,Wei-hanChen1,andRayGer1

1DepartmentofForestry&naturalResources,nationalIlanUniversity,no.1,sec.1,shen-lungRd.,Ilan,taiwan260(jjmao@niu.edu.tw)2Appliedbehaviouralecology&ecosystemResearchUnit,Departmentofenvironmentalsciences,

UnIsA,privatebagX6,Florida,1710,RepublicofsouthAfrica

themany-bandedKrait(Bungarus multicinctus multicinctus;Fig.1)occursnaturally inburma,theIndo-Chinesepeninsula,southern

China,hainan,hongKong,andtaiwan(U.s.Departmentofthenavy1991,ZhaoandAdler1993).Wefoundadead-on-road(DoR)maleon2november2007onthenorthernCrossRoad(Routeno.7)closetominchi(elevation1,021m;tWD97,n24.614164°,e121.482272°),YilanCounty,northerntaiwan.snout-ventlength(sVlinmm),tail-length(tlinmm),andbodymass(bming)were740mm,100mm,and98.9g(excludingstomachcontents),respectively.Weimmediatelynoticedthatanundigestedsnake’stailandthekrait’sownheartwereprotrudingfromthedamagedpartoftheupperthroatregion(Fig.2).Dissectionrevealedtwoophidianpreyitemsinthedigestivetract,ajuvenilemale

mountainWolfsnake(Lycodon ruhstrati ruhstrati;sVl260mm,tl82mm,bm6g)andanadultfemalemountainKeelback(Pseudoxenodon stej-negeri stejnegeri;sVl490mm,tl88mm,bm41.1g).theanteriormostportionsofbothpreyitemswerepartlydigested,whichindicatedtheyhadbeeningestedinheadfirstpositions(Fig.3). lueetal.(2002)suggestedthatthecolorationofL. r. ruhstrati isaformofbatesianmimicry,causingittoresembleB. m. multicinctus,althoughthesetwospeciesdifferintheirdefensivebehavior(norcal[sic]andmao2006).mao(1970)reportedcannibalisminB. m. multicinctusintaiwan.therefore,thekrait-resemblingcolorationofL. r. ruhstratimayprovideadegreeofprotectioninmostpredatorencounters,butnotagainstpredationbyB. m. multicinctus.

Fig. 1.themany-bandedKrait(Bungarus multicinctus multicinctus)intaiwanoccursinavarietyofhabitats,manyofwhich(e.g.,lowgrassyareas,ricepaddies,irrigationditches)areassociatedwithwater.

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Pseudoxenodon s. stejnegerioccursmainlyinmountainousregions atelevationsofabout1,000mabovesealevel,butB. m. multicinctususuallyoccursatlowerelevations.pope(1935)reportedB. m. multicinctusat700minKuatun,mainlandChina,andassumedthatthatwasthespecies’maxi-mumelevationaldistribution.Kuntz(1963)statedthattheyoccurprimarilyatlowelevations(below600m)intaiwan,andlueetal.(2002)surmisedthatthissnakespeciesoccursmainlyataltitudesbelow500m,andthattheyonlyrarelycanbefoundat500–1,000m.ourobservation,however,cannotconclusivelyprovethatB. m. multicinctusoccursnaturallyinminchi,becausesomesnakes,especiallytheChineseCobra(Naja atra),many-bandedKrait,andorientalRatsnake(Ptyas mucosa)arereleasedbycertainreligiousgroupsalongRouteno.7,especiallybetweensze-lanandminchi(mao,pers.obs.;YoandWong2007).however,ourobservationdoesshowthatsnakescanbeexploitedbyB. m. multicinctusinmountainousareas. Kuntz(1963)indicatedthatB. m. multicinctusintaiwanoccursinavarietyofhabitats(e.g.,lowlandwoodedareas,bamboothickets,grassyareasassociatedwithwater,ricepaddies,irrigationditches),whereasmao(1970)simplystatedthatB.m.multicinctusprefershabitatsnearwater.many-bandedKraitsreportedlyfeedprimarilyonloaches,frogs,skinks,snakes,andmice(mao1993,Zhaoetal.1998).slowinski(1994)notedthat,comparedtotheotherkraitsinthegenus Bungarus(e.g.,B. ceylonicus,B. fasciatus,andB. caeruleus),B. m. multicinctusappearstobemorepiscivorous(63.16%)than ophiophagous(34.21%).thisconclusionwasbasedontheresultsofmao

(1970),whoalsoreportedthatlargerindividualstendtobemoreophiopha-gousthansmallerones,andthatpreycorrelatedcloselywithhabitat,althoughhefailedtoprovidehabitatdescriptions,andlistedonlytaoyuanCountyasacollectionsite.linetal.(1995),ontheotherhand,reportedB. m. multicinctustobemoreophiophagous,andattributedpiscivorytoontogeneticchangesindietortolow-elevationwetlandpollutionand/orhabitatdestructionforindustrialandcommunitydevelopment.basedonthepreyitemsreportedandthecollectionsitedescriptioninmao(1970),wedeterminedthatallpreytypesdescribedwerespeciesrestrictedtolow-elevationricepaddies,ponds,andadja-centareas.ourconclusionisfurthersupportedbydescriptionsofthedomi-nantlandutilizationofthetaoyuanareaatthattime(Yang1994),whichwereidealforwetlandfaunaandflora.thereportoflin(1995),incontrast,wasbasedonspecimenscollectedbyroadsampling,especiallyinmountainousareas.Wethusconcludedthattheapparentlycontradictorydietarypreferencesreportedbymao(1970)andlin(1995)maymerelyreflectsamplingofdif-ferenthabitats,leadingtothequestionwhetherhabitatheterogeneitycouldaccountforunusualpreyexploitationbyB. m. multicinctus. In1997–2009,weconductedecologicalstudiesonsnakesinwetland(e.g.,ricepaddies,drainageditches,ponds,streams,etc.)andnon-wetland

Fig. 2.the road-killedmany-bandedKrait,with its heart and the tail of aPseudoxenodon stejnegeri stejnegeriprotrudingfromthedamagedsectionofitsbody.

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Fig. 3.Fromlefttoright:theroad-killedBungarus multicinctus multicinctusanditsprey:Lycodon ruhstrati ruhstratiand Pseudoxenodon stejnegeri stejnegeri.notethepartlydigestedheadregionsofthesnakesthatwereconsumed.

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table 1.preyofmany-bandedKraits(Bungarus multicinctus multicinctus)collectedfromwetland(e.g.,pondsandricepaddies)andnon-wetland(e.g.,secondaryforestsanddrycultivatedareas)habitatsintaiwan.Anasterisk(*)indicatesthataspeciesoccursinbothhabitattypes,but,inthiscase,itwasrecordedfromabetelnutpalmplantation,whichisclassifiedasdrycultivatedland.References:1thisstudy;2mao1970;3Dayetal.1994;4linetal.1995;5maoetal.2006;6lin2008;7mao,unpubl.data;8norval,unpubl.data;9Ji-shouYang,perscomm.

Wetlands Non-wetlandssnakes Enhydris plumbea(n=9)2 Ramphotyphlops braminus (n=1)4

Bungarus m. multicinctus(n=1)2 Amphiesma stolatum*(n=1)7

Xenochrophis piscator(n=3)7 Cyclophiops major (n=2)3,7

Orthriophis taeniura friesei(n=1)8

Lycodon r. ruhstrati (n=1)1

Pseudoxenodon s. stejnegeri(n=1)1

Protobothrops mucrosquamatus(n=1)5

Viridovipera s. stejnegeri(n=2)7,9

lizards Plestiodon chinensis formosensis(n=1)2 Plestiodon elegans(n=2)4

Eutropis multifasciata (n=1)6 Sphenomorphus indicus (n=1)4

Fish Misgurnus anguillicaudatus (n=10)2

Fulta alba (n=14)2

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areas(e.g.,drycultivatedlands,betelnutpalmplantations,mountainousandfoothillsecondaryforests,bamboothickets,etc.)oftaiwan.Apartfromchanceencounters,wealsomadeuseoftrapstocollectsnakes.Inwetlandhabitats,weemployedfloatingfunneltraps,baitedwithloachestocollectsemi-aquaticsnakes,and,exceptforthesewatersnakes(i.e.,ChineseWatersnake,Enhydris chinensis;plumbeousWatersnake, E. plumbea;RingedWatersnake,Sinonatrix annularis;oliveKeelback,S. percarinata suriki;andChequeredKeelback,Xenochrophis piscator),B. m. multicinctus wasthemostfrequentlycollectedspeciesandalwaysateallofthebaitinthetrap.Ashortdrift-fence/funnel-trapunitwasutilizedinnon-wetlandareas.Again,someB. m. multicinctus werecollected,althoughthenearestpermanentwatersources(e.g.,stream,wetland,pond,etc.)wereasfaras1kmaway.WecomparedthepreyitemsofallB. m. multicinctusencounteredinthesehabitatsandthoserecordedintheliterature(table1). exceptforspeciessuchastheFive-linedblue-tailedskink(Plestiodon elegans), IndianForestskink(Sphenomorphus indicus),ChineseGreensnake(Cyclophiops major),L.r. ruhstrati, P. s. stejnegeri,andChineseGreentreeviper(Viridovipera stejnegeri stejnegeri),whichoccurtomoderateeleva-tions(~1,000m)intaiwan,mostofthereptilianpreyofB. m. multicinctusarespeciesthatoccurinthelowlands(lueetal.2002).WealsofoundthatB. m. multicinctus fromwetlandhabitatspreymorefrequentlyonloaches,eels,andsemi-aquaticsnakesinthatorderoffrequency,whereasB. m. mul-ticinctusfromnon-wetlandormountainousforesthabitatspreyprimarilyonterrestrialorsemi-arborealsnakes(e.g., C. major;V. s. stejnegeri;taiwanhabu,Protobothrops mucrosquamatus,Fig.4)andterrestrialskinks(e.g., P. elegans,S. indicus). pope(1935)reportedpredationonmiceandfrogsbyB. m. multicinc-tus,butneitherofthesepreytypeswererecordedfromB. m. multicinctusinnature.Consumptionofmiceandfrogsappearstobebasedonobserva-tionsundercaptiveconditions;atbest,theyareraredietaryitemsofB. m.

multicinctus,despitereferencestothecontrary(e.g.,maki,1935).Undercaptiveconditions,wehaveobservedthatafewB. m. multicinctusacceptnewbornmiceasprey,whichsuggestsindividualvariationinacceptingvari-ousprey.Asforfrogs,eventhoughmaki(1935),Kuntz(1963),andmao(1970,1993)listedfrogsaspreyitemsofB. m. multicinctus,wewereunabletoconfirmthatpossibility.WhileworkingattheWildlifeAnimalshelter,nationalpingtungUniversityofscienceandtechnologyin2001,thefirstauthorneverobservedB. m. multicinctusattemptingtopreyontwosym-patricwetlandfrogs(CricketFrog,Fejervarya limnocharis andtaiwaneseFrog,Hoplobatrachus rugulosus)thatwereofferedasfood.FrogsrecordedinstomachcontentsofB. m. multicinctusmighthavebeenpreyofothersnakes,suchasthebuff-stripedKeelback(Amphiesmastolatum),E. plumbea,orX. piscator,whichhadbeeneatenbykraits.however,regardlessofpreytype(snakes,loaches,eels,orskinks),allshareonefeature—acylindricalbodyshape.thiswouldindicatethatB. m. multicinctusexhibitsapreferenceforcylindricalpreyitems,asdosomeseasnakes(e.g.,VorisandVoris1983). Inconclusion,preyavailabilityindifferenthabitattypesappearstobereflectedinthedietofB. m. multicinctus,withonlycylindricalbodyshapeasacommondenominator.Consequently,studiesdescribingpreytakenbykraitsshouldconsiderthetypeofhabitatfromwhichtheanimalsinques-tionweretaken.

Literature CitedDay,Y.t.,h.I.Wu,andC.C.Chen.1994.bandedKrait(Bungarus multicinctus)

fedonGreenVinesnake(Eurypholis major).Notes and Newsletter of Wildlifers2(3):4(inChinese).

Kuntz,R.e.1963.snakesoftaiwan.Quarterly Journal of Taiwan Museum16:17–19.

lee,W.J.andK.Y.lue.1996.Apreliminarystudyonthefoodhabitsofsnakesintaiwan.Biological Bulletin of the National Taiwan Normal University31:119–121(inChinese).

Fig. 4.Amany-bandedKraitcatchingandconsumingataiwanhabu(Protobothrops mucrosquamatus).

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lin,D.n.2008.noskink—IntroductionandpestcontrolofinvasiveMabuya multifasciata.Quarterly Journal of Nature Conservation61:30–36(inChinese).

lin,h.C.,K.F.hwung,andt.h.lu.1995.Foodofsnakesfromtaiwan.Notes and Newsletter of Wildlifers 3(2):19–21(inChinese).

lue,K.Y.,m.C.tu,andG.s.shang.2002.The Transition Kingdom — Guidebook of Amphibians and Reptiles of Taiwan.sWAn,taipei(inChinese).

maki,m.1931.A Monograph of the Snakes of Japan.Dai-Ichishobo,tokyo.

mao,J.J.,G.norval,W.b.Kung,andh.n.Chang.2006.Bungarus multicinctus multicinctus(many-bandedKrait).Diet.Herpetological Review37:350.

mao,s.h.1970.Foodofthecommonvenomoussnakesoftaiwan.Herpetologica26:45–48.

mao, s.h. 1993. Common Terrestrial Venomous Snakes of Taiwan. nationalmuseumofnaturalscience,specialpublicationno.5,taipei.

norcal[sic],G.andJ.J.mao.2006.mountainWolfsnake(Lycodon ruhstrati ruh-strati):Adescriptionofthedefensivebehavior.Sauria28(4):47–50(notethatthesurnameofthefirstauthorisatypographicalerrorfornorval).

pope,C.h.1935.The Reptiles of China.theAmericanmuseumofnaturalhistory,newYork.

slowinski,J.b.1994.thedietofkraits(elapidae:Bungarus).Herpetological Review25:51–53.

U.s.Departmentofthenavy(bureauofmedicineandsurgery).1991.Poisonous Snakes of the World.Doverpublications,newYork.

Vorish.K.andh.h.Voris.1983.Feedingstrategiesinmarinesnakes:Ananaly-sisofevolutionary,morphological,behavioralandecologicalrelationships.American Zoologist23:411–425.

Yang,s.l.1994.theirrigationsystemevolvedonthesocialandspatialorganiza-tionofthetaoyuanplateau.Unpublishedm.s.thesis,nationaltaiwannormalUniversity,taipei(inChinese).

Yo,C.W.andZ.J.Wong.2007.preliminaryinventoryofherpetologicalresourcesofnorthCrossRoad.Quarterly Journal of Nature Conservation59:38-47(inChinese).

Zhao,e.m.andK.Adler(eds.).1993.Herpetology of China.societyforthestudyofAmphibiansandReptiles.oxford,ohio.

Zhao,e.m.,m.h.huang,andY.Zong.1998.Fauna Sinica, Reptilia. Volume 3: Squamata, Serpentes.sciencepress,beijing,China(inChinese).

blocked-flight aggressive behavior in snakes

D.brucemeans

CoastalplainsInstituteandlandConservancy,tallahassee,Florida32303

everynaturalistaccumulatesfieldobservationsandexperiencesworthyofexpandingintofull-blownresearchprojects.moreoftenthannot,how-

ever,theinformationlanguishesundevelopedinmemory,paperpiles,andthesedays,computerfiles.someinterestingobservationsonsnakebehaviorIhavemadeovertheyearsareamongmymanyunfinishedprojects. experiencednaturalistsknowthatmostsnakesfleewhenencounteredinnature.moresedentary,slow-moving,orcrypticsnakeswillremainstillorcoiledinordertogounnoticed.Rattlesnakes,whendisturbed,coilandrattle.Afterawhile,theyalsomaymoveawayfromtheintruder,perhapscontinuingtorattleastheybreakoutoftheircoilandflee.Afewsnakes,however,dosomethingdifferent.

Fig. 2.ACottonmouthmouth-gapingasitcrawlstowardtheauthor.

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Fig. 1.ACottonmouth(Agkistrodon piscivorus)“chasing”theauthorwhileengagedinaggressivebehaviorduringblockedflightwithheadraised,striking,flatteningitsbody,vibratingitstail,andmovingdirectlytowardtheauthor.thissnakedidnotfollowtheauthorwhenhesteppedaside,butracedtosafetyintothewaterbeyond.

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thatdifferentbehaviorwas firstdisplayed tomewhenanadulteasternCottonmouth(Agkistrodon piscivorus conanti)roseuplikeacobraandmenacinglycrawledtowardmeinfrontofadozenstudents(Fig.1).IhadjustguaranteedmyfieldclassthatCottonmouthsdonotchasepeople.Inthosefewmoments,asIrapidlybackedupfromtheapproachingsnake,apotentialbehavioralstudywasborn(seeChapter8inmeans2008). Wehadencounteredthesnakeonewarmlateafternooninmay,whensnakes—andCottonmouthsespecially—aremostlikelytomove.Itwascrossingawide,sandyroadintheApalachicolanationalForestoftheFloridapanhandletheadjacentswampfromwhichtheCottonmouthhademergedwasdryingupfromaseveredrought,andthesnakeprobablywaslookingfornewaquatichabitatoradryingwaterholefromwhichtogorgeontheconcentratedaquaticanimals. Assooftenhappenswithsnakes,itstoppedcrawlingasmyvehicleapproachedandwasfullystretchedoutina“frozen”posture.IlinedupthestudentsontheroadshoulderfromwhichthesnakehadcomeandItookapositionontheotherside,blockingitsforwardmotion.ourmove-mentsstimulatedittocoilupanddoits“Cottonmouth”display,withitsattentionfocusedonme,whowasclosesttothesnakeandgesticulatingtothegroup.Afterafewminutes,thesnakesuddenlyroseupcobra-likeandbegancrawlingrapidlyatmewiththeforwardone-fourthofitsbodyraised. Withitsneckspreadslightly,thesnakecontinuedtoadvanceasIbackpedaled,makingafewstrikingparriesasitnearedmyfeet.WhenIsteppedsidewaysfromthesnake’strajectory(whichwasdirectlytowardaswampontheothersideoftheroad),Iwasgratifiedthatthesnakemain-taineditsoriginaldirectionanddidnotturntofollow(or“chase”)me.Its“aggressive”behaviorobviouslywasablufftoassistthesnakeinmakingitsgetawayintothesafetyoftheswamp. Realizing that this poorly trafficked, wide sandy road betweentwoswampsofferedanidealsitetostudythisbehavior,mywife,Kathysteinheimer, and I road-cruised this 0.5-km stretch ten times in thenext two years. she videotaped 16 episodes of me interacting withCottonmouths.Cottonmouths“chased”me10of17timesthatIblockedtheirmovement.Itsimplyappearstobebluffingbehavior,butaggressivenevertheless.Icallitshammed(bluffed)aggressionduringblockedflight.

Cottonmouthsengagedinsuchaggressiveflightexhibitedseveralcharacteristicbehaviors,includingcrawlingrapidlytowardme,raisingtheheadandneckofftheground,expandingthejaws,flatteningthebody,vibratingthetail,tongue-flicking,lungingandstrikingforward,andrarely,musk-squirting.Agreatdealofvariationexistsintheintensityofthesebehaviors.Afewdisplayedthebehaviorastheydashedtowardthecoveroftheroadsidevegetation.someassumedtheclassiccoil-upandmouth-gapedefensefirstandthenbrokeintoaggressiveflightwithinafewminutes. IwasabletoinitiateflightbehaviorinacoiledCottonmouthsimplybyflippingitoveronitsbackbymeansofmysnakehook.Uponrightingitself,thesnakeendeduplyinginastraightposition.Ittheneithercoiledagainandmouth-gaped,orfled,atwhichtimeIcouldstepinfrontofit.otherCottonmouthswererelativelyplacid,coilingrepeatedlyandrefusingtoflee.ofthe17,onehiditsheadunderitsbodywhencoiledandrefusedtobreakoutofthatposture.Anothermouth-gapedasitcrawledrapidlytowardme(Fig.2).AnotherinterestingbehaviorwenotedisthatwhenfleeingCottonmouthscrawledoutofthesandyroadbedandenteredtheroadsidevegetation,theyspeduptheirforwardmotionandbecameevenmoreanimatedintheiraggressivebluffing. Allenandswindell(1948)werethefirsttodescribeaggressiveflightintheCottonmouth:“onethreefooter,whenapproached,struckupwardsohardthatheliftedhisbodyofftheground.headvanced,coilingandstrikingrepeatedly,followingtheretreatoftheobserver.othershavebeenseentochargedeliberatelywithopenmouth.mostattackersseemtodosoinabluffingmannerratherthaninviciouspursuit.” morerecently,studieshaveanalyzedCottonmouthdefensivebehav-iorsinresponsetobeingsteppedonorapproachedclosely,andtobeingpickedupwithanartificialarm(GibbonsandDorcas2002,GlaudasandWinne2007).Inthesestudies,cagedsnakesorthosewithunrestrictedopportunitiestofleewerelesslikelytodisplayaggressiveflightbehavior.Apparently,thebehaviorismostlikelytobetriggeredwhenfree-rangingsnakesareblockedalongachosentrajectoryaftertheyarealreadyinflight. myintentioninvideotapingthese“chase”episodeswastodoastudyofCottonmouthbehaviorduringblockedflightbyanalyzingthefilmedsequences.Alas,whenIreturnedfromanoverseastrip,Idiscoveredthatthe

Fig. 3.AneasternCornsnake(Pantherophis guttatus)crawlingtowardtheauthor,strikingforward,asheblocksitsescapefromadirtroad.

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tapecontainingtheepisodeshadbeenaccidentallyoverwrittenwithsoapoperas,sotheprojecthasnotbeenresurrected.becausethetwoofuswerebusyvideotapingtheshort-lastingbehavior,wetookfewstillphotos,whichonlyshow“snapshots”ofthefullbehavior. overtheyears,Ihaveencounteredsimilarshammedaggressionduringblockedflightbyseveralspeciesofsnakes.thebehaviorisinducedonlywhenoneplacesone’sbodyinfrontofamovingsnake,blockingitsescape.manysnakes,especiallycolubrids,willturnandfleeinanotherdirection,butsomedonot.InsnakesassmallaspigmyRattlesnakes(Sistrurus miliarius barbouri),Istimulatedsimilarbluffedaggressionepisodesintwoofsevenindividuals.IevenencounteredaneasternCornsnake(Pantherophis guttatus)crossingaroadthatroseupandtriedtobluffmeoutofitsway(Fig.3).

InVictoriastate,Australia,Ideliberatelysteppedinfrontofacrawl-ingeasternbrownsnake(Pseudonaja textilis)andgotanothersurprise(Fig.4).notonlydiditrisecobra-likeandmovethreateninglytowardsme,butitalsoseemedpurposefullytoexaggeratetheundulationsofitsbodyintheloosegraveloftheroad.thisfast-movingsnakeisextremelyvenomousandresponsibleformoresnakebitedeathsinAustraliathananyotherspe-cies(Whitakeretal.2000).Fortunately,asIsteppedoutofitstrajectory,itdidnotturntowardmebutattemptedtomakegooditsescapeinthedirectionitwasfleeing.Whittakerandshine(1999)described455defen-

siveresponsesoffree-rangingeasternbrownsnakestoexperimentalcloseencounterswithhumans,butscoredonly12oftheseinwhichthesnakeadvancedtowardstheobserver.ofthese,onlythreeadvanceswerejudgedtobeoffensive.Intheirstudy,however,theydidnotreportattemptstoblockthepathofthesnakeonceitbegantoflee.ResearchershaveobservedthatmanyotherlargeAustralianelapidsreacttoanintruderbyrisingup,flatteningtheneck,openingthemouth,andcontinuinginthedirectionoftheintruder(Johnson1975,Greer1997). mymostinterestingencounterwithabluffingsnakehastobeanexam-pleofbehavioralmimicry.onemorninginJune2002,IwaswalkingalongthesparselyvegetatedfloodplainoftheescambiaRiverinwesternFloridawhen,suddenly,asnakeroseupcobra-likeandcrawledrapidlyuprightwhileIwalkedatasafedistancealongsideittakingphotosofwhatIassumedwasanotherCottonmouthengagedinaggressivebluffing.Itsbodyproportions,pattern,andbehaviorweresolikeaCottonmouththatIdidn’tdiscoveritstrueidentityuntilIlookedatmydigitalreadout.ItwasaperfectCottonmouthmimic,asouthernbandedWatersnake,Nerodia fasciata(Fig.5). Divertingmanyspeciesofsnakesfromthebeelinetrajectoryoftheirintendeddirectionisoftendifficult.thishasbeenparticularlyevidenttomeduringattemptstophotographmovingsnakes.manyseemfixatedontheoriginaldirectionoftheirmovementsothatonecannoteasilymakeasnakereverseitsheading.Dragthesnakebackwardsanditwillrepeatedlycrawlforwardalongitsprevioustrack. Ibelievethatthe“aggressivebluffing”behaviorIhavewitnessedintheCottonmouth,pigmyRattlesnake,Cornsnake,easternbrownsnake,andbandedWatersnakewereelicitedunderthespecialcircumstancesofencoun-teringthesnakesinhabitatsfamiliartothemandbyblockingtheirescapealongadirectiontheyhadchosen.Futurestudiesofaggressivebehaviorsexhib-itedduringblockedflightshouldtakethesea priori conditionsintoaccount. Iprobablydonothavetimeinmylifetotraveltheworldlookingforopportunitiestoblocktheescapepathofwildsnakes,butmaybebyreport-ingtheseobservations—aboutwhichIfeelsurehenryFitchwouldhaveenjoyedreading—willstimulateothernaturaliststowatchfor“bluffedaggressionduringblockedflight”insnakeseverywhere.Isuspectthatmostexperiencedfieldherpetologistshavehadsimilarencounterswithsnakes,andthelistofsnakespeciesthatdothismaybelong.Icannothelpbutwonderifshammedaggressionbehaviorsaremoreprevalentinvenomoussnakes,sincetheyaremorelikelytosurviveanattackbyenvenomatingtheintruderifthebluffdoesn’twork.maybesomeonewillconductin-depthstudiesandamorecompletereviewofthisinterestingbehavioralphenom-enoninsnakes,althoughIurgesnake-loversnottotrythesepotentiallydangerousexperiments,andprofessionalherpetologistswhomightdososhouldwearpropersnakebootsandothersnakebite-preventionapparel,unlikemeinFigures1and4.

Literature CitedAllen,e.R. andD.swindell. 1948.theCottonmouthmoccasinofFlorida.

Herpetologica4(supplement1):1–16.

Gibbons,J.W.andm.e.Dorcas.2002.DefensivebehaviorofCottonmouths(Agkistrodon piscivorus)towardhumans.Copeia2002:195–198.

Glaudas,X.andC.t.Winne.2007.Dowarningdisplayspredictstrikingbehav-iorinaviperidsnake,theCottonmouth(Agkistrodon piscivorus)?Canadian Journal of Zoology85:574–578.

Greer,A.e.1997.The Biology and Evolution of Australian Snakes.surreybeatty&sons,Chippingnorton,newsouthWales,Australia.

Johnson,C.R.1975.DefensivedisplaybehaviorinsomeAustralianandpapuan-newGuineanpygopodidlizards,boid,colubridandelapidsnakes.Zoological Journal of the Linnean Society56:265–282.

means,D.b.2008.Stalking the Plumed Serpent and Other Adventures in Herpetology.pineapplepress,sarasota,Florida.

Whitaker,p.b.andR.shine.1999.Responsesoffree-rangingbrownsnakes(Pseudonaja textilis,elapidae)toencounterswithhumans.Wildlife Research26:689–704.

Whitaker,p.b.,K.ellis,andR.shine.2000.thedefensivestrikeoftheeasternbrownsnake,Pseudonaja textilis(elapidae).Functional Ecology14:25–31.

Fig. 5.AsouthernbandedWatersnake(Nerodia fasciata)mimickingtheaggressiveflightbehavioroftheCottonmouth,sham-strikingforwardwithheadraised,bodyinflated,andjawswidened.

d. B

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Fig. 4.Aneasternbrownsnake(Pseudonaja textilis)comingstraightfortheauthorashebacksaway.

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mostnestingfreshwaterturtlesminimizetheirtimespentterrestrially,leavingthewatertonestandreturningimmediatelyfollowingnest-

ing(burkeetal.1994).exceptionstothisincludesometurtlesthatmigratelongdistancestonest(e.g.,Emydoidea blandingi;Roweandmoll1991)andspeciesthatburythemselvesafternesting(e.g.,Kinosternon flavescensandK. subrubrum;Iverson1990,burkeetal.1994).putativereasonsforburrowinginKinosternonincludenestattendance/defenseandawaitingconditionsthatwouldreducestressorincreasesurvivalduringtheturtles’returntowater(Iverson1990,burkeetal.1994).

northAmericansoftshells(Apalonespp.;Fig.1)arewellknownfortheircharacteristicbehaviorofburrowingintosandandothersoftsub-stratesunderwater(Webb1962,GrahamandGraham1991);however,littleisknownregardingtheirpropensitytoburrowterrestriallyafternest-ing.Wehaveobservedshallowterrestrial“troughs”inassociationwithnest-ingcrawlsofA. muticaandA. spinifera onsandbarsthatresultfromfemalescrawlingjustunderneaththesurfaceofthesandandresurfacingsomedis-

Terrestrial burrowing in nesting softshell Turtles (Apalone mutica and A. spinifera)

michaelV.plummer1andJ.seanDoody2

1Departmentofbiology,hardingUniversity,searcy,Arkansas72143,UsA(plummer@harding.edu)2schoolofbotanyandZoology,AustraliannationalUniversity,AustralianCapitalterritory,0200,Australia(sean.doody@anu.edu.au)

photographsbytheseniorauthor.

Afternesting,femalesoftshellshavebeenobservedtoburrowforvariabledistancesunderthesandsurfaceincloseproximitytothejust-completednest.WedescribeterrestrialburrowingbehaviorinApalone muticaandA. spiniferafromKansas,Arkansas,andlouisianaanddiscusspossiblereasonsforthebehavior.

Fig. 1.maleApalone spinifera(top)andmaleA. mutica(bottom).

Fig. 2.softshellhabitatonsandbarsoftheKawRiverinKansas(top)andnestinghabitatalongtheWhiteRiverinArkansas(bottom).

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tanceaway.WealsohaveobservedtheprocessoftroughexcavationinoneA. mutica. Collectively,wehaveobservedatotalof8troughsofA. muticaonsandbarsoftheKansasRiverinKansas(plummer1976,unpubl.data,7troughsofA. muticaontheWhiteRiverinArkansas(plummer,unpubl.data),and7troughsofA. spiniferaand4unidentifiedApalonetroughsontheComiteRiverinlouisiana(Doody1995).troughsvariedinlengthfrom0.5–4.0m(mean2.2±1.24msDinlouisiana)andabout20–30cminwidth.troughs(Fig.3)variedinshapefromstraighttocurvedtosinuoustofigure-eightshaped.onetroughthatappearedtobemadebyonefemaleA. muticaconsistedofthreeseparatesegmentsincloseproxim-ity,eachapproximately2–3minlength. mostnestingcrawlsofA. muticaandA. spiniferadidnotexhibittroughs(72of77;93.5%nestsofA. muticaand198of209;94.7%nestsofA. spinifera).however,5of8(63%)troughscontainednestsinKansasand11of15(73%)troughscontainednestsinlouisiana.troughswerefoundonlyduringthenestingseason(latemay–midJulyforbothspecies).exceptforbasking,thisperiodisgenerallytheonlytimethehighlyaquaticsoftshellsarefoundinterrestrialsituations(Webb1962).Wehavefoundbothpre-andpost-nestingadultfemales(Fig.4)burrowedtypicallyattheexitendofthetrough,butwehavenotfoundhatchlings,juveniles,adultmales,orsubadultfemalesintroughs.nestsmayoccuranywhereinthetrough,butusuallyweretowardoneend,mostoftenattheentrance(Fig.3b). Fewliteraturereferencesaddressnestingtroughsortheirfunction.harper(1926)notedthatnestingsoftshells(probablyA. ferox)sometimes

burrowedbeforereturningtowater.Apalonespp.generallynestonopensandbarsonsunnydays(Webb1962,plummer1976,Doody1995),whenextendedexposuretohightemperaturescouldpresumablybelethal.onehypothesisforthefunctionoftroughsisthatmothersburrowtotemporar-ilyescapehighsurfacetemperaturesandawaitoptimalconditionsforreturntowater.thisbehaviorissimilartothatofmudturtles(Kinosternon),whichnestlongdistancesfromtheiraquatichabitats(burkeetal.1994). Alternatively, burrowing turtlesmay exhibit cryptic behavior inresponsetodisturbanceorpredationrisk.Insupport,oneA. muticabur-rowedterrestriallyafterbeingdisturbedbymVpwhileshewasnesting.burrowingbyApalone is common inaquatic situations (GrahamandGraham1991).Forexample,whenapproachedfromsomedistance,indi-vidualturtleswillreadilyburythemselvesunderthesandinshallowwater(GrahamandGraham1991;JsD,pers.obs.;mVp,pers.obs.). Anotherpossiblefunctionisthatnestingturtlesburythemselvestomorecloselyassessthesuitabilityofsubstratecharacteristicssuchastem-perature.however,only5%of68A. spiniferanestsinonestudywereassociatedwiththebehavior(Doody1995).Finally,mothersmaysim-plyrespondtoaphysicalconstraint.extendedhotweatherconditionsorcoarse-grainedsandmayrendersurfacesandnotcohesive,causingthesandtocaveinonitselfwhenaturtleattemptstoexcavateachamber(Doodyetal.2004;Fig.5)orwhenhatchlingstrytoemergefromthenestchamber(mortimer1990;Fig.6).Asaconsequence,thecloselyrelatedpig-nosedturtle(Carettochelys insculpta)maynestatlowerelevationsonsandbarsduetothefriablesandathigherelevations(Doodyetal.2004);however,lowerelevationsincreasetheriskoffloodmortality.byburrowing,softshell

plUmmeRAnDDooDY

Fig. 3.(A)Acurvedtrough(dashedoutline)ofApalone spiniferaontheComiteRiverinlouisiana.(b)Astraighttrough(dashedoutline)ofA. muticaontheWhiteRiverinArkansas.theresearcherisprobingtofindtheexactlocationofthenest.

Fig. 4.Femalesoftshells,suchasthisApalone spiniferafromGinCreek,Arkansas,canbequitelarge.

A

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motherscouldexcavateanestincohesivesandbyremovingtheuppermostloosesand.thisputativebehaviorisperhapsasimplerversionofbody-pittingdisplayedbyseaturtlesandtheAmazonianGiantRiverturtle,Podocnemis expansa(schaubleetal.2006,Vogt2008).body-pittingmayaidturtlesattemptingtonestonsandbarsthathavealayerofloose,drysand.Insupportofthishypothesis,Apalonenestsassociatedwithtroughsweredeeperthanthosewithouttroughs(Doody1995). Althoughwehavegeneratedhypothesesfortheunusual“troughing”behaviorexhibitedbysoftshells,currentevidencedoesnotallowustofavoranyparticularone.Focalobservationsofnestingturtleswouldbeparticu-larlyusefulbutdifficultgiventherelativelylowfrequencyofthebehavior.Accordingly,thethermoregulationhypothesiswouldbedifficulttotest.however,theothertwohypothesesmaybemoreeasilytestable.First,mockpredatorsorsomeotherdisturbancecouldbeintroducedtonestingturtlestotestthepredationriskhypothesis.second,thecohesivesandhypothesiscouldbeaddressedbyattemptingtoexcavateanartificialnestchambernexttothetrough.Failuretocreateachamberwouldlendsupporttothisidea(Doodyetal.2004).

AcknowledgmentsAfter informinghenryFitchabout theunusualbehaviorofanestingfemaleApalone muticathatmVphadobservedontheKansasRiverduringthe1973nestingseason,henrysaid,“Yououghttolookintothat.”Wellhenry,hereitis—thanksforthesuggestion.

Literature Citedburke, V.J., J.W. Gibbons, and J.l. Green. 1994. prolonged nesting forays

byCommonmudturtles (Kinosternon subrubrum).American Midland Naturalist131:190–195.

Doody,J.s.1995.Acomparativenestingstudyoftwosyntopicspeciesofsoftshellturtles(Apalone muticaandApalone spinifera)insouthcentrallouisiana.Unpublishedm.s.thesis,southeasternlouisianaUniversity,hammond.

Doody,J.s.,A.Georges,andJ.e.Young.2004.Determinantsofreproductivesuccessandoffspringsexinaturtlewithenvironmentalsexdetermination.Biological Journal of the Linnean Society80:1–16.

Graham,t.e.andA.A.Graham.1991.Trionyx spiniferus spiniferus(easternspinysoftshell).buryingbehavior.Herpetological Review22:56–57.

harper,F.1926.talesoftheokefinokee.American Speech1:407–420.

Iverson,J.b.1990.nestingandparentalcareintheturtle,Kinosternon flavescens.Canadian Journal of Zoology68:230–233.

mortimer,J.A.1990.theinfluenceofbeachsandcharacteristicsonthenest-ingbehaviorandclutchsurvivalofGreenturtles(Chelonia mydas).Copeia1990:802–817.

plummer,m.V.1976.someaspectsofnestingsuccessintheturtle,Trionyx muti-cus.Herpetologica32:353–359.

Rowe,J.W.ande.o.moll.1991.Aradiotelemetricstudyofactivityandmove-mentsoftheblanding’sturtle(Emydoidea blandingi)innortheasternIllinois.Journal of Herpetology25:178–185.

schauble,C.,R.Kennett,ands.Winderlich.2006.Flatbackturtle (Natator depressus)nestingatFieldIsland,Kakadunationalpark,northernterritory,Australia,1990–2001.Chelonian Conservation and Biology5:188–194.

Vogt,R.2008.Amazon Turtles.Graficabiblos,lima,peru.

Webb,R.G.1962.northAmericanrecentsoft-shelledturtles(Familytrionychidae).University of Kansas Publications, Museum of Natural History13:429–611.

Fig 5.AfemaleApalone muticaexcavatinganestalongtheWhiteRiver,Arkansas.

Fig. 6.hatchlingApalone mutica readytoemergefromanest(thetop5cmofsandhasbeenbrushedback)alongtheComiteRiver,louisiana.

82 IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010

apparent competition by Grass frogs (Rana temporaria) and common Toads

(Bufo bufo) for breeding sitesIvanVergner

Zbysov33,CZ-28565ZbysovvCechach,CzechRepublic(ivan.vergner@atlas.cz)

photographsbytheauthor.

theGrassFrog(Rana temporaria;Fig.1)iswidespreadthroughoutmostofeurope,rangingfromnorthernspaintotheUrals(absentfrom

southernandcentralIberia,muchofsouthernItaly,andtheCaucasus)andeasttowesternsiberiaandnorthernKazakhstanthroughnorthernGreeceandbulgaria.thespecieshasapatchydistributioninmountainouspartsofthebalkans.Ithasbeenrecordedfromsealeveltoelevationsapproaching2,700mabovesealevel(asl)inthepyrenees. theCommontoad(Bufo bufo;Fig.2) iswidespread ineurope(excluding Irelandandmostmediterranean islands, althoughpresentonsicily)andnortherneurasia,withpopulations inpartsofwestern

Asia(turkey,syrianArabRepublic,andlebanon)andnorthernAfrica(morocco,Algeria,andtunisia).Ineurope,thespeciesispresentinmostareas(includingtheUnitedKingdomandscandinavia),rangingasfareastasnorthernKazakhstanandeasternsiberia.Inthemiddleeast,itisfoundthroughmuchofturkey,northwesternsyrianArabRepublic,andhasrecentlybeenrecordedfromtwomountainouslocationsinlebanon.InAfrica,ithasaveryfragmentedrangeinthemoremountainousregions.thespecieshasanaltitudinalrangeofsealevelto3,000masl;however,innorthernportionsoftherangethespeciesoccursonlyatlowerelevations. IobservedreproductiveactivityofGrassFrogsataforestpondonthesoutheasternedgeofthemiddlebohemianRegionatanaltitudeof~410masl(49°48’n,15°21’e;Fig.3).oncequitecommonintheregion,populationsofthesefrogshaverecentlybeendeclining,withextendeddryperiodsthroughthespringandsummerofrecentyearsplayingamajorrole.Droughtconditionsleavefrogbroodswithalowprobabilityofsurvivinguntilmetamorphosis. Reproductiveactivityforlocalamphibiansbeginswithspringwarm-ingtrends.on3April2009,largenumbersofmaleCommontoads(Bufo bufo)appearedatthepond,andIobservedegg-layingandfertilizationdur-ingthefollowingtwodays.thefirstvocalizingmaleGrassFrogsbegantocongregateon6April.ovipositingbeganthateveningandcontinuedthroughthefollowingday(Fig.4). onthemorningof7April,agroupofabout15maleGrassFrogswereinterspersedwitharowdycollectionofabout10Commontoads.malesofbothspecieswereactivelyvocalizing.Askirmishensuedamongthemalesofbothspecies,leavingindividualtoadsandfrogslimpingandinjured.onebat-tlebetweentwoGrassFrogsandoneCommontoad(Fig.5)wasparticularlyheated,asthesemalesapparentlycompetedforprimecallingandmatingsites.

VeRGneR

Fig. 2.AnadultfemaleCommontoad(Bufo bufo).

Fig. 3.thebohemianforestpondwherefrogsandtoadsapparentlycompetedforcallingandmatingsites.

Fig. 1.AnadultmaleGrassFrog(Rana temporaria)approachingthebreedingpond.

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Fig. 4.CongregatingGrassFrogsandeggmasses.

Fig. 5.AheatedconflictbetweentwomaleGrassFrogsandonemaleCommontoad.Clutchesoffrogeggsarevisibleinthewater.

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submerged calling by oregon spotted frogs (Rana pretiosa) Remote from breeding aggregations

Jaybowerman

sunrivernatureCenter,p.o.box3533,sunriver,oregon97707(frogs1@sunrivernaturecenter.org)

Frogcalls,largelyassociatedwithreproduction(i.e.,advertisementandreleasecalls),havebeenthesubjectofmuchinterestandstudy(schiötz

1973,Rosenandlemon1974,Wells1977,Fellers1979,Duellmanandtrueb1986,Kelley2004).Althoughwemostlyhearfrogcallsandcho-rusescomingfromthesurfacesofponds,anumberofspeciescallwhilesubmerged.Xenopusandotherpipids,allhighlyaquaticfrogs,callwhilesubmerged(elepfandt1996,Christensen-Dalsgaardandelepfandt2004).platz(1993)describedanewspeciesofleopardfrog(Rana subaquavocalis;butseeGoldbergetal.2004)thatexclusivelyemitsmatingcallswhilesub-mergedthatareinaudibleatthesurface.RanidsfromthewesternUnitedstates,taxonomicallydistinctfromtheirkinintheeasternU.s.(CnAh,2009),haverelativelyweakcallsthatdonotcarryfarfrombreedingloca-tions.severalofthesewesternranidscallbothatthesurfaceandwhilesub-mergedwheninornearbreedingaggregations(storm1960,licht1969,briggs1986,mactagueandnorthern1993). theoregonspottedFrog(Rana pretiosa)isahighlyaquaticexplosivebreederthatoverwintersundericeinmuchofitsextantrangeandspawnssoonaftericeleavestheponds(hayes1994).eggsaredepositedincom-munalaggregationsofafewtosometimesmorethanahundredeggmasses.Inthesunriverareaincentraloregon(elevation1,320m),breedingbeginssometimebetweenearlymarchandearlyApril,dependingonconditions.Althoughthebreedingseasonmaylastuptoamonth,breedingactivityatindividualovipositionsitestypicallyspansaperiodoftwotofivedays(pers.obs.).Duringbreedingevents,malesgatheratovipositionsites,call-ingactivelyatthesurfacewithbreedingtakingplacemostlyduringtheday. Recordedadvertisement callsofR. pretiosa consistof a stringoflow“clucks”orknocks,emittedatarateof3–7/sec,withpeakintensitybetween600–900hz.Releasecalls,emittedbybothmalesandfemales,weremoreofasingleordouble“squawk”givenatarateof1–2squawks/secandwithafrequencyspectrumsimilartoadvertisementcalls.Releasecalls

canbeelicitedalmostanytimebypickingupafrogwiththethumbandindexfingersplaceddirectlybehindtheforelimbs.RecordingsofR. pretiosa advertisementandreleasecallsatthesunrivernatureCenterareavailableatwww.sunrivernaturecenter.org. here,IreportforthefirsttimethatR. pretiosaalsocallsactivelyfromsubmergedlocationstenstohundredsofmetersawayfromandseveraldayspriortotheformationofbreedingaggregations.

Methods sub-surface listening was conducted using a DolphineAR™hydrophone(modelDe100-6-112),witha6-mcableandpreamporDolphineARpRohydrophoneconnecteddirectly toaZoomh-4digitalrecorder.listeningwasconductedbycastingthehydrophonefromshoretothelengthofthe6-mcable,allowingitsosink,listeningfor1min,retrievingapproximately1m,andlisteningagain.Recordingsweretakenwhengoodqualitycallingsignalsweredetected. twositeswereutilizedforsubmergedcallingsurveys.bullfrogpond(bFp,43°51’01”nx121°26’50”W)isa0.3-haisolated(nohydrologicAdultoregonspottedFrog(Rana pretiosa).

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sonogramofthemaleadvertisementcallofRana pretiosa.

sonogramofthereleasecallofRana pretiosa.

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connectionstonearbywaterbodies),excavatedpond,somewhatirregularinshape,approximately100mlong,and10–25mwide.maximumdepthis2.5–3m.springrunoffraisesthewaterlevelasmuchas1m,creatingashallowarmthatextendsupto150mfromoneendofthepermanentportionofthepond.breedingtakesplaceinthedistalportionofthistem-poraryarm.themaximumdistancefromanypointwithinthepermanentpondtotheovipositionsiteisapproximately200m.egg-masscountsoverthepast10yearsrangedfrom21–200+(median24).markandrecapturestudiessuggestonlylimitedmigrationintooroutofbFp(unpubl.pers.obs.).ovipositiongenerallytakesplaceover1–3days. lakeAspen(lA,43°53’10”nx121°26’47”W)isa2.5-haexcavatedpondwithyear-roundconnectionstootherwaterbodies,includingtheDeschutesRiver.Waterlevelsarecontrolledbyweirsandfluctuatebynomorethan30cm.Along-termmarkandrecapturestudy(unpubl.data)hasdocumentedthatseveralhundredadultR. pretiosamovetolAinthefalltooverwinter,enteringviaashallowoutletchannel.InmarchandApril,frogsmigrateoutoflAtospawnatacommunalovipositionsiteknownasDuckpondmarsh(Dp,43°53’28”nx121°26’52”W)approximately1kmaway.AtDp,eggmassesaredepositedinseveralcollectiveclustersoveranareaofapproximately10x80m.spawningatDpspans2–4weeks,usuallyinburstsof2–5daysatanyonecollectiveaggregation(unpubl.pers.obs.).DuetothelargersizeoflA,noattemptwasmadetosystematicallysurveytheentireshoreline.Instead,thehydrophonewasdeployedatlocations

alongabout200mofshorelineoneithersideoftheoutletchannelusedbyfrogstomovetotheovipositionsite.

ResultsbFpwasice-coveredduringsurveyson14march2002,exceptforopenwaterataseepatthesouthend.by21march,thespringthawhadpro-vidednumerousopeningsaroundtheedgesofthepond,although5–8cmoficeremainedovermuchofthepond.theonlysoundsdetectedbyhydrophoneon14and21marchwerebubblesescapingfromtheice.Atapproximately1100hon28march,advertisementcallsofmaleR. pretiosaweredetectedatadepthofapproximately0.5mneartheedgeofapatchofCattails(Typhussp.)withinthepermanentpond.threemalesappearedtobecallingatthatlocation.Additionalsmallgroupsofcallingmalesweresubsequentlydetectedatfouradditionallocationsscatteredaroundtheperimeterofthepond.Ifounddetectablecallsonlyinanarrowzone2–3mfromshoreatdepthsof0.5–1m.movingthehydrophonemorethan2–3mawayfromacallingindividualwassufficienttolosethecallbecauseofsoundattenuation.nofrogcallswereaudibleatthesurface,andallfrogsappearedtobeconcealedinsubmergedgrassorsedgeonthebottom.theweatheron28marchwascool(~12°C)withaheavyovercast.Isawonefrogatthesurfaceofthepondduring2hofsurveying,amaturefemalethatremainedatthesurfaceforapproximately5minafterdetection,thensubmergedandwasnotseenagain. on30march,ahighpressureweathersystemarrived,bringingwarmsunnyweather.on31march,abreedingaggregationofmalesformedatthehistoricbreedinglocationandbreedingcommenced.Atotalof21eggmassesweredepositedoverthecourseofapproximately24–36h,withbreedingcompletedsometimeon1April.eighteeneggmasseswereclus-teredtogether,withthreesatellitemassesdeposited<3mfromthemainaggregation.thedistancefromthebreedingsitetothevarioussubmergedcallinglocationswas80–150m.Duringoviposition,10–20maleswereobservedcallingfromthesurfaceinthevicinityoftheeggmasses.Callingwasaudibleupto30maway.somemalesalsowereobservedcallingfromthebottomin10–15cmofwater,andtheircallswereaudiblebutmuffled.nocallingwasdetectedawayfromthebreedingsiteon30march. hydrophonelisteningatbFpin2003and2005failedtodetectanysubmergedcallingpriortoorremotefromtheformationofasurfaceaggre-gationandsubsequentoviposition.At1000hon15march2004,asinglemalewasdetectedcallingfromasubmergedposition~10mfromtheeven-tualovipositionsite.by1330h,severalmaleshademergedattheoviposi-tionsiteandwerecallingintermittently.thatevening,from2030–2130h,allsurfaceactivityattheovipositionsitehadceased,butanumberofmaleswerecallingactivelyattheafternooncallinglocationandattwoadditionallocationsinthepermanentportionofthepondapproximately80mfrom

Authorholdingpre-ampandcableofaDolphineARhydrophoneontheshoreoflakeAspen.

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Communalovipositionsiteshowing~18newlylaideggmassesadjacenttoandon12eggmasseslaidseveraldaysearlier.Greenalgaearedevelopingwithinoldereggmasses.

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schematicofbullfrogpondshowing2002and2006locationsofsubmergedcallingandsubsequentbreeding.

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theovipositionsite.thefollowingday,alldetectableactivityhadshiftedtotheovipositionsiteandbreedingcommenced. In2006,iceremainedonthepondsuntillatemarchandwaterstoodatrecordhighlevels.on3April2006,Idetectedseveralmalescallingfroma20-mzoneinthemiddlesectionofthetemporaryarmofthepond.thiscallingzonewas20–40mfromtheovipositionsiteusedinprioryears,and40–60mfromthesitethatwouldbeusedin2006.Afterdark,at2100h,severaldozenmalesweredetectedcallingatdepthsof30–60cmwithinthissamenarrowband,about20mlong.multiplecallingmalescreatedacontinuouschorus.Atthistime,nosurfaceactivitywasdetectable.From1000–1130hon4April,Idetectedsubmergedcallingatthissamelocation,butnomorethantwoorthreefrogswereheard.nocallingwasdetectedfromwithinthepermanentpond.by1400h,afewmaleshadsur-facedandwerecallingnearthehistoricovipositionsite.on5April,surfaceactivityhadshiftedabout20mfartherupthetemporaryarmandspawningcommenced.overthenextthreedays,46eggmassesweredepositedinonemajorclusterandfivesatellitemasses2–5maway. lakeAspen,averydifferentsite,yieldedsomewhatdifferentresults.At1130hon4April2002,onefrogwasdetectedcallingabout4moff-shorenearthepondoutletatadepthofabout1.5–2.0m.Callingwassporadic,withpausesoftenlastingseveralminutes.Asecondindividualwasdetectedatasimilardepthabout30mtothewest.nofurthercallingwasdetectedthatyearandnosubmergedcallingwasdetectedduringseveraldaytimesurveysin2006.At1300hon28march2007,individualmalesweredetectedattwoofsixlocationswherethehydrophonewasdeployed.At2100hthatevening,however,1–3malescouldbeheardatfourofsixlisteninglocations.At1100hthefollowingday,asinglefrogwasdetectedcallingatoneofthelocations.thesecallinglocationsareapproximately1kmfromthemainovipositionsite.

DiscussionAdvertisementcallsofmaleRana pretiosa, ahighlyaquatic frog,weredetectedcomingfromsubmergedlocationsseparatedfromovipositionsitesbytenstohundredsofmetersandsometimesprecedingtheformationofbreedingaggregationsbyseveraldays.submergedcallingactivitywascon-siderablygreaterafterdark,eventhoughmostbreedingactivityoccursdur-ingtheday.oncebreedingaggregationsformedatovipositionlocations,malesweredetectedonlyinthevicinityofthebreedingaggregation,withcallingrestrictedtothesurfaceoratdepthsshallowenoughforthecallstobeaudibleabovewater. platz(1993)suggestedfourpossibleselectiveadvantages forsub-merged calling: Avoiding predators, avoiding temperature extremes,extendingthebreedingand/ordevelopmentalseason,andincreasingsound

transmissionrange.Incentraloregon,theprincipalpredatorsonR. pre-tiosaincludeGartersnakes(Thamnophis sirtalis),Greatblueherons(Ardea herodias),mink(Neovison [=Mustella]vison),andRaccoons(Procyon lotor).mostofthesepredatorsarevisual,daytimehunters.Callingwhilesub-merged,especiallyatnightandatdepthsgreaterthan30cm,wouldhelpR. pretiosaavoidthesepredators,exceptforraccoons. IncentraloregonandindeedthroughoutmuchoftherangeofR. pretiosa, temperaturesonclearnightsduringthebreedingseasonfallrap-idlyaftersunset,andovernighttemperaturesfrom-10–-5°Carecommon,althoughwatertemperaturesafewcentimetersbelowthesurfacetypicallyremainat8–14°C.submergedcallingwouldthusavoidthecoldtempera-turesatthesurface.IncreasingthelarvaldevelopmentalperiodisprobablynotimportantforR. pretiosa atsunriver,wheremetamorphosisisnormallycompletedinlateJulyorAugust,butmightbeimportantatsomehighele-vationssiteswheresnowandicecandramaticallyshortentheactiveseason.Despitethedensityofwater,submergedcallingdoesnotappeartoincreasesoundtransmissionrangeatthedepthsandintheenvironmentusedbyR. pretiosa.ladichandbass(2003)statedthat“…inshallowwaters,soundpropagationisverymuchlimited….”Indeed,Ifoundsubmergedcallingwasoftenundetectableatdistancesgreaterthan1–3mfromthesourcewhenthefrogswerehiddeninsubmergedvegetation. manyifnotallofthespeciesofRanafoundinthewesternUnitedstatesapparentlycallbothatthesurfaceandwhilesubmergednearbreed-ingaggregations(storm1960,licht1969,briggs1986).thisappearstobethefirstreportofsubmergedcallingbyanyofthesespeciesthatissepa-ratedfrombreedingaggregationsbothspatiallyandtemporally.AnecdotalreportsplaceamplecticpairsofR. pretiosa atvaryingdistancesfrombreed-ingsites.however,whetherremotepairingissimplyopportunisticasfrogsmovetowardbreedingsitesorifactivemateselectionmaybetakingplaceremotelyfrombreedingsitesandrelatedtosubmergedcallingisunclear.severalimportantquestionsremaintobeanswered.Isactivemateselec-tiontakingplacepriortoarrivalatbreedingaggregations?Ifso,thenwhatisthefunctionoflargeaggregationsofunpairedmalesfoundcallingatthesurfaceatovipositionsites?Conversely,ifmateselectionisprincipallyaccomplishedinthebreedingaggregationsatovipositionsites,thenwhatfunctiondoessubmergedcallingserve?Doesthepresenceofbothsurfaceandsubmergedcallinginseveralspeciesofwesternranidssuggestthatthiscladeisevolutionarilyintransitionfromonestrategytowardtheotherorthatcompetingselectivepressureshelpmaintaindualstrategies?Dootherwesternspeciesthatemitadvertisementcallsbothatthesurfaceandsub-merged(i.e.,R. cascadae,R. boylii,andR. luteiventris)engageinsubmergedcallingpriortoandatsomedistancefromtheformationofbreedingaggre-gations?Answeringthesequestionswillprovideabetterunderstandingoftheecologyofthisentiregroupofanurans.

Literature Citedbriggs,J.l.1987.breedingbiologyoftheCascadeFrog,Rana cascadae,withcom-

parisonstoR. aurora andR. pretiosa.Copeia1987:241–245.

Christensen-Dalsgaard,J.andA.elepfandt.2004.biophysicsofunderwaterhear-ingintheClawedFrog,Xenopus laevis. Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology176:317–324.

CnAh(Center fornorthAmericanherpetology).2009.standardCommonandCurrentscientificnames(Frogsandtoads).www.cnah.org/nameslist.asp?id=3.

Duellman,W.e.andl.trueb.1986.Biology of Amphibians.mcGraw-hill,newYork.

elepfandt,A.1996.UnderwateracousticsandhearingintheClawedFrog,Xenopus,pp.177–191.In:R.C.tinsleyandh.R.Kobel(eds.),The Biology of Xenopus.oxfordUniversitypress,newYork.

Fellers,G.m.1979.Aggression territoriality, andmatingbehaviour innorthAmericantreefrogs.Animal Behaviour27:107–119.

Goldberg,C.s.,K.J.Field,andm.J.sredl.2004.mitochondrialDnAsequencesdonotsupportspeciesstatusoftheRamseyCanyonleopardFrog(Rana subaquavocalis).Journal of Herpetology38:313–319.

CommonGartersnake(Thamnophis sirtalis)swallowingaspottedFrog(Rana pretiosa).

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hayes,m.p.1994.thespottedFrog(Rana pretiosa)inWesternoregon.partI.technicalReport94-1-01.oregonDepartmentofFishandWildlife,salem.

Kelley,D.b.2004.Vocalcommunicationinfrogs.Current Opinion in Neurobiology14:751–757.

ladich,F.andA.h.bass.2003.Underwatersoundgenerationandacousticrecep-tioninfisheswithsomenotesonfrogs.In:s.p.Collinandn.J.marshall(eds.),Sensory Processing in Aquatic Environments.springer,newYork.

leonard,W.p.,l.hallock,andK.R.mcAllister.1997.Rana pretiosa (oregonspottedFrog).behaviorandreproduction.Herpetological Review28:28.

licht,l.e.1969.ComparativebreedingbehavioroftheRed-leggedFrog(Rana aurora aurora)andtheWesternspottedFrog(Rana pretiosa pretiosa)insouthwesternbritishColumbia.Canadian Journal of Zoology47:1287–1299.

mactague,l.andp.t.northern.1993.UnderwatervocalizationbytheFoothillYellow-leggedFrog(Rana boylii).Transactions of the Western Section of the Wildlife Society29:1–7.

platz,J.e.1993.Rana subaquavocalis,anewspeciesofleopardFrog(Rana pipienscomplex)fromsoutheasternArizona.Journal of Herpetology27:154–162.

Rosen,m.andR.e.lemon.1974.thevocalbehaviorofspringpeepers,Hyla crucifer.Copeia1974:940–950.

schiötz,A.1973.evolutionofanuranmatingcalls:ecologicalaspects,pp.311–319.In:J.l.Vial(ed.),Evolutionary Biology of the Anurans.Universityofmissouripress,Columbia.

storm,R.m.1960.notesonthebreedingbiologyoftheRed-leggedFrog(Rana aurora).Herpetologica16:251–259.

Wells,K.D.1977.thesocialbehaviorofanuranamphibians.Animal Behaviour25:666–693.

suburbandevelopmentinWoodway(asuburbofWaco),mclennanCounty,texashadbegunin1964,whenIstarteda46-yearprojectto

recordthearea’snaturalandunnaturalherpetofaunalhistory(Gehlbach2002).the80-hasavannawaspartofaranchtotallyerasedbysuburbiaduringthestudy,andabout20%ofadjacentwoodlandwaslost.oneearth-dammedstockpondwaskeptforfishing;theotherwasdemolishedbyfloods.habitatprotectionwasinadvertentlyaffectedbycitybuildingcodesthatsavedforestedravineswithsteepslopes,creeksandtheirterraces,

andbyaplanned-unitdevelopment’s6-haprivatenaturepreserveofcreek,forest,andwoodlandin1984. thisstudypresentsfeaturesofextirpation,survival,andgeneralabun-danceofeachspeciesofamphibianandreptileanddescribesherpetofaunalchangeduringthestudysite’ssuburbandevelopment.Ihopeitwillstimu-lateotherstudiesthatsupportconservationandeducationaluseofnaturepreservesinourever-growingU.s.cities.

suburbanization of a central Texas Herpetofauna

FrederickR.Gehlbach

Departmentofbiology,baylorUniversity,Waco,tX76798

photographsbytheauthor.Allweretakenatthestudysite.

Reptilesandamphibiansweresurveyedonthebalconesescarpmentincentraltexasastheareawastransformedintosuburbia.offourhabitats,oak-junipersavannawastotallydestroyedandaquarteroftheadjacentoak-juniperwoodlandwaseliminated,buttwocreeksandmostdeciduousriparianforestremained.extirpationof12of30species(40%)included55%ofnineamphibianspecies,50%oftwoturtles,33%ofsixlizards,and30%of13snakes.herpetofaunalrichnessdeclinedexponentiallyfrom1965to1990withincreasedhousebuilding,thenstabilizedthrough2005asbuildingslowedandstopped.speciessurvivalwasrelatedtouseofculturalandremainingnaturalhabitats,secretivebehavior,andhumaninterest.

AmatingpairofCope’sGraytreefrogs(Hyla chrysoscelis). strecker’sChorusFrog(Pseudacris streckeri).

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MethodsFieldworkcoveredthe120-hastudysite(elevation165–196m)onthebalconesescarpment in allmonths in1964–2009.originalhabitatswerematureevergreenoak-junipersavanna,alivestockpasturewithtree“islands”(locallycalledmotts),twopermanentcreeks(eachwithastockpondina12-mdeepravinewithdeciduousriparianforest),anddeciduousoak-juniperwoodlandonlimestoneslopeswithrockoutcropsabovetheforestedravines.observationalandcatchandreleasesurveysinmarch–octoberincludeda3.3-kmtransect(graduallyreducedbyhousebuildingto1.2km)inallhabitats2–4times/week,20coveredpitfalltrapsarranged

randomlyin200m2ofprotectedriparianforestandcheckeddailyinApril–June1984–1989,anddailydead-on-road(DoR)recordson3–5kmofcitystreetsborderingnaturalhabitatsandamonghousesinthestudysiteand1kmbeyonditonallsides. speciesaregroupedasfollows:(1)extirpated(noevidenceforfiveconsecutiveyears),(2)uncommonsurviving(notedlessfrequentlythanweekly),or(3)commonsurviving(noteddailytoweekly).proportionalherpetofaunalchangeswereanalyzed,aswasthecumulativenumberofextirpatedspeciesperfive-yearperiodin1965–2005relativetotheconcur-rentrateofhousebuilding.

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Greatplainsnarrowmouthtoad(Gastrophryne olivacea). smallmouthsalamander(Ambystoma texanum).

texasspottedWhiptail(Aspidoscelis [Cnemidophorus] gularis gularis).

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scientificnamesfollowCollinsandtaggart(2009)orsubsequentupdatesonthewebsiteoftheCenterfornorthAmericanherpetology(www.cnah.org);ifrecentlychanged,previousnamesareinbracketstoassistrecognition.

ResuLtsextirpated species:

Woodhouse’stoad(Anaxyrus[Bufo]woodhousii woodhousii).Uncommon;disappearedby1970withoutevidenceoflocalreproduction. Cope’sGraytreefrog(Hyla chrysoscelis).Uncommonmarch–Aprilbreederinastockpondandadjacentcreekpoolsinriparianforestuntilgamefishwerestockedinthepondandsomeescapedintothecreekin1980;thisfrogdisappearedin1982. strecker’sChorusFrog(Pseudacris streckeri).Commonnovember–Februarybreederinthesavanna’srainpoolsfurtherformedaslivestockwallowsinnaturaldepressionsuntileliminatedby1975. Greatplainsnarrowmouthtoad(Gastrophryne olivacea).UncommonbreederinsavannarainpoolsduringApril–Juneuntil1975,butdidnotdisappearuntil1979. smallmouthsalamander (Ambystoma texanum).Common in theriparianforest;bredduringoctober–Decemberrainsincreekpoolsandthestockpondthatbecameafishingpond.thesecond-mostfrequentDoRamphibian;goneby1982. ornateboxturtle (Terrapene ornata).only two records in thesavanna;noneafter1975. texashornedlizard (Phrynosoma cornutum).Common in thesavannauntil1975;subsequentlyuncommoninsuburbanvacantlots

until1978.Apronounceddeclinethroughoutcentralandeasterntexascoincidedwithinsecticidesprayingfromairplanesinthe1970stoeradi-catepopulationsofimportedFireAntsthatexpandedduringsuburban-ization. texasspottedWhiptail(Aspidoscelis [Cnemidophorus] gularis gularis).Commoninthesavannauntil1975andlateruncommoninvacantlotsamonghouses;goneby1986.

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broad-bandedCopperhead(Agkistrodon contortrix laticinctus).

texaspatchnosesnake(Salvadora grahamiae lineata).

AnadultYellowbellyRacer(Coluber constrictor flaviventris).

Groundsnake(Sonora semiannulata).

AjuvenileYellowbellyRacer(Coluber constrictor flaviventris).

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texaspatchnosesnake(Salvadora grahamiae lineata).thisrarestofthe13originalsnakespecieswasfoundonlytwiceinoak-juniperwood-land;notseenafter1988. broad-banded Copperhead (Agkistrodon contortrix laticinctus).Uncommon in the two natural wooded habitats and occasionally inwoodedsuburbanyardsuntil1970.Awinterden5maboveacreekinalimestonecrevicehad19sunningadultsinoctober1967;thedenwasapparentlydestroyedbydynamiteandlaterbulldozedduringdevelopmentofplanned-unithousing;extirpatedby1971. linedsnake(Tropidoclonion lineatum).onlythreeindividualswerefoundundergroundsurfacecoverinthesavanna,allbefore1975,makingthisthesecondrarestsnakeintheoriginalherpetofauna. Groundsnake(Sonora semiannulata).Uncommon;1–2foundannu-allyundersurfacecoverinevergreenwoodlanduntil1983.bothplainbrown-coloredindividualsandsnakeswithblackheadsandneckspotsonadullorange-browngroundcolorwerepresent.

uncommon surviving speciessouthernleopardFrog(Lithobatessphenocephalus utricularius [Rana sphe-nocephala]).onceacommonbreederincreekpoolsinmarch–may,andstillcommonincreeksandpondsoutsidethestudysite.Graduallylessabundantinthestudyplot;notrecordedin2007–2009,henceextirpationispossible. northernCricketFrog (Acris crepitans). Inhabitscreek sidesandbreedsincreekpoolsinApril–may,but,liketheotheranurans,eggsandtadpolesarewasheddownstreambyfloodsenhancedbystreetdrainage.effectivereproductionseemsincreasinglylessinfrequentforthisandallofthesuburbancreekbreedersthatremaincommontoabundantelsewhereinlocalruralareas. three-toedboxturtle (Terrapene carolina triunguis).originallyuncommoninriparianforestandrareinevergreenwoodland.since1991,onlyonemarkedadultmalehasbeenfoundannually(butnotin2009),sothepresentstatusisunknown.Afewliberatedpetsfromoutsidetheregionwereknownbefore2000,butextirpationisnowquitepossible. YellowbellyRacer(Coluber constrictor flaviventris).moderatelyabun-dantintheoriginalsavannaandwoodland,butgraduallymoreuncommoninremainingwoodlandandwoodland-yardedges,whereithuntslizardssuchasnorthernGreenAnolesandtexasspinylizards.originally1–3DoRrecordsperyear,butonlyonesince2000,reflectingreducednum-bers. Flatheadsnake(Tantilla gracilis).thisfossorialspeciesissecretivebutpersistsinremainingforestandwoodland.equaltoRoughearthsnakes,Flatheadsnakesweretheleastfrequentcatchinpitfalltrapsandaretheonlysurvivingsnakenotyetfoundondevelopedproperty.thisisthethird

mostfrequentDoRsnake,mostoftenonroadsborderingorlessthan75mfromremainingnaturalhabitat. texasGartersnake(Thamnophis sirtalis annectens).Alwaysuncommon,althoughitpersistsinriparianforestnearcreeksandoccasionallyappearsinsuburbanyardsandgardensthatadjoinremainingnaturalhabitat. easternhognosesnake(Heterodon platirhinos).Anothersnakeorigi-nallyuncommonandraretoday.becauseofitsinitialaggressive,neck-flat-tening,defensivebehavior,itiscommonlythoughttobedangerousandissometimesconfusedwiththeCopperhead,hencesummarilykilled.Coastalplaintoadsareknownfood.A48-cm(totallength)adultwasactiveat16°CinearlyDecember.oneDoRwasrecordedin2007.

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Anadulteasternhognosesnake(Heterodon platirhinos)exhibitingadefensivedis-playagainstapredator.

Coastalplaintoad(Ollotisnebulifer [Bufo valliceps]).

mediterraneanGecko(Hemidactylus turcicus)onthebrickwallofahouse.

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Common surviving species:Coastalplaintoad(Ollotisnebulifer [Bufo valliceps]).themostabundantanuranandmostfrequentDoRinallyears,apparentlybecauseindividu-alscongregateonstreetstofeedoninsectsbelowstreetlights.layseggsincreekpoolsandbackyardpondsduringApril–Augustrains,thelongestbreedingseasonamonganurans.Itistheonlyamphibianthatutilizesarti-ficialpondsatsuburbanresidences.Whetheranindividualbreedsmorethanonceannuallyisunknown.suburbanitesdonotmindtoadsontheirproperty.

RioGrandleopardFrog(Lithobates [Rana]berlandieri).breedsincreekpoolsduringAugust–october.Increasinglylesscommon,although,asnotedforotheranurans,abundantinpondsoutsidethestudyarea.streetandyardwaterdrainsintocreeks,sofloodedeggsandtadpolesand/ordis-solvedtoxinsmaybedetrimentstocontinuedexistence. mediterraneanGecko(Hemidactylus turcicus).IntroducedtoWaco,texasintheearly1950s(b.brown,pers.comm.)andtothestudysiteby1982.livesonbrickandstonewallsandinsidehousesandotherbuildings;hatchlingsappearanytimeexceptDecember–February.Adultsareactive

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northernGreenAnole(Anolis carolinensis carolinensis)onapottedporchplant.

texasspinylizard(Sceloporus olivaceus).

WesternRatsnakes(Scotophis obsoletus[Elaphe obsoleta])readilyclimbtrees.

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throughouttheyearduringnightswithtemperaturesabove~15°C,butinwinteronlyifbrickshavebeensun-heated.oneDoR. texasspinylizard(Sceloporus olivaceus).Inopenforest,woodland,andwoodedsuburbanyardsonthegroundandrocksandintrees(seealsoblair1960).Rarelyclimbsherbaceousvegetation.Gardensandbackyardrefusepilesarenestsites,alsoforGreenAnoles,Groundskinks,WesternRatsnakes,andprobablyotherspecies.themostfrequentDoRlizard. northernGreenAnole(Anolis carolinensis carolinensis).Commonaroundhouses,lesssoinforestandwoodlandedges.livesinplantedherba-ceousvegetation,shrubs,andtreesingardens,andclimbspottedpatioandporchplantsandhousewalls.thisspeciesincreaseddramaticallyduringsuburbanization,benefitingfromhumaninterestandplantingsthatattractitsinsectfood.two–threebroodsperyearinJune–september.Adultsareactivethroughouttheyearinsunnyweatherattemperaturesabove~15°C.thethirdmostfrequentDoRlizard. Groundskink(Scincella lateralis [Lygosoma laterale]).onlyslightlylessabundantthantheGreenAnole,butrarelynoticedbysuburbanitesbecauseof itssecretiveterrestrialhabits inandundergroundcover inwoodedyardsandgardens,especiallyatforestandwoodlandedges;2–3broodsperyearinJuly–september(seealsoFitchandGreene1965).thisisthemostfrequentreptileinpitfalltraps,andtheonlyabundantspecieswithoutaDoRrecord. RoughGreensnake(Opheodrys aestivus).moderatelycommoninopenriparianforestandbrushyedgesalongcreeksandinyardsandgar-dens,whereitisunnoticedlyinglengthwiseonlowfoliagewhilesit-and-waitforagingforinsects.Itsgreencolorationandbehaviorhideitsoeffec-tivelythatIneverheardofonekilledbypeople.oneDoRrecord.

WesternRatsnake(Scotophis obsoletus[Elaphe obsoleta]).thislargestoflocalsnakes(to2mtotallength)iscommoninforestandwoodland,lesssoonwoodedsuburbanproperty.ItisthesnakeIammostfrequentlyaskedtoremove;forexample,a1.5-madultcoiledinsideacookingpotinakitchencabinet.Activeinmarch–november;eggshatchinAugust–september.hatchlingsandadultseatlizards,includingmediterraneanGeckosonbrick-walledhousestosecond-floorheights.Anestpredator

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RoughGreensnake(Opheodrys aestivus).

Rough earth snake (Virginia striatula) in pre-shedding condition (note the“cloudy”eye).

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ofopenandcavity-nestingbirds,includingsmallowls(Gehlbach1994).FourthmostfrequentDoRsnake. texasblindsnake(Rena[Leptotyphlops]dulcis).Abundantinsub-urbia,mostlyduringApril–Juneunderflatgroundcoverobjectssuchaspathwaysteppingstones;alsodugupinflowerbeds,foundaliveinswim-mingpools,ongaragefloors,ineasternscreech-owlnests(Gehlbachandbaldridge1987)andFireAntmounds(baldridgeandWivagg1992).thesecondmostfrequentlyrecordedDoRsnake,andtiedwiththetexasbrownsnakeforsecondmostfrequentpitfallcatch. texasbrownsnake(Storeria dekayi texana).likeothersmallinver-tebrate-eatingsnakes,encounteredmostlyduringthespringundergroundcoverafterrains.liveslargelyinriparianforestbutalsoinsuburbanyards,althoughlesscommonlythanblindandRoughearthsnakes.Itisthesec-ondmostfrequentpitfallcatchandthirdmostabundantDoRsnake. Roughearthsnake(Virginia striatula).themostabundantsuburbansnakeandDoRreptileofanyspecies.lawnandgardenwateringmightpromoteitsearthwormfood,whilesmallsize,burrowingbehavior,browncoloration,anduseofground-surfacecovercontributetoitsexistenceinsuburbia.Activeinmarch–november.

Discussionsurvivalwasnilforspeciesthatdependedmostlyontheerasedsavanna.strecker’sChorusFrogs,Greatplainsnarrowmouthtoads,andornateboxturtlesdidnotsurvive,whiletexashornedlizardsandtexasspottedWhiptailsremainedonlytemporarilyonlargevacantgrassysuburbanlotsintheformersavannaarea.otheramphibiansbredindecliningnumbersincreekpoolsorwereextirpatedbystockedgamefish.however,onetoad,threelizards,andfoursnakesbegantolivepartlyinsuburbanyardsandgardens.mostpeopledidnotmindtoads,buttwoofthelizardsandallofthesnakessurvivedinsuburbanhabitatsbybeinginconspicuous. nativespeciesmostcommonlyencounteredinsuburbiawere, inorderoffrequency,northernGreenAnole,Coastalplaintoad,Roughearthsnake,texasspinylizard,Groundskink,texasblindsnake,andRoughGreensnake.theyandothersurvivorsbenefitedfromthestudysite’sheat-island(landsberg1981),whichaveraged1.5°Chigherthanacorrespondingruralvalueinmarch–october(Gehlbach1994),andsub-urb-forestedgehabitatthatsupplied40%morepotentialinsectpreythan12mawayintheforestinteriorinseptember–october(D.Quigley,pers.comm.). humanattitudeswerenegativetowardthelargestsnakesandcontrib-utedtotheextirpationofCopperheadsanddeclinesinYellowbellyRacersandeasternhognosesnakes.homeownerssometimesaskedmetoremove

largesnakes,butnotthoselessthan~15cmintotallength,whichmanybelievedwere“worms.”smallsize,browncolor,soilorground-litterhabi-tat,andgreencolorcombinedwithsit-and-waitforaginghidsomelizardsandsnakesinsuburbia.Conversely,thenorthernGreenAnole’sreddewlapdisplays,bodycolorchangescoordinatedwithperchcolor,andprominentuseofcultivatedplantsdrewhumaninterestandprotectionthatmighthaveenhancedtheobviousincreaseinthenumbersofthisspeciesinsuburbia. Creek-breedinganuranswerenegativelyaffectedbywaterrunofffromcitystreetsandprivateproperties.theseunnaturalsourcesprobablycon-tainpetroleumandpesticidetoxins,andstreetdrainsconcentraterainintoflashfloodsthatwasheggsandtadpolesoutofthecreeks.thus,southernleopard,RioGrandeleopard,andnorthernCricketfrogsdeclined—butnotCoastalplaintoads,perhapsbecausetheirbreedingseasonis1.7timeslongerthanthoseoftheotheranurans,andhencearemorelikelytoavoidfloods.Also,onlythistoadbredinandmighthavebenefitedfromartificialpondsinsuburbanyards. twelve(40%)ofthestudysite’s30originalspeciesofamphibiansandreptilesdisappearedduringsuburbanization.thisfractionissignificantlylessthanthe67%extirpationof21speciessurveyedbyminton(1968)inanoldersuburbofIndianapolis,Indiana(c2=7.2,p=0.007).Amphibianlosseswerecoincidentwithdegradedaquatichabitatsinbothsuburbsandproportionatelyhigherthanthoseofreptiles:75%versus61%ofspeciesinIndianaand55%versus38%intexaswithoutintra-orinter-sitediffer-ences(c2<0.23,p>0.63). therateofherpetofaunaldeclineinthistexasstudywas1–5spe-ciesperfive-yearperiodofexponentiallyincreasingsuburbangrowthdur-ing1965–1995.subsequently,noadditionalspecieswerelostassuburbangrowthbecameprogressivelyslowerthrough2005.Duringninefive-yearperiodsthrough2005,thechangingandthenstableherpetofaunawasstronglycorrelatedwiththecumulativenumberofcitybuildingpermitsforthestudysiteandsurrounding1-kmwidesuburbanarea(3°polynomialregression,r2=0.95,p=0.001).

AcknowledgementsColleagues,family,friends,andstudentsjoinedhunts;nancyGehlbachwaspresentmostfrequentlyandmadehelpfulcommentsonthemanu-script.DenaQuigleyprovidedinsectabundancedatafrommalaisetraps.henrys.Fitchparticipatedinatransectsurveyinthe1970s,andIamhon-oredtodedicatethisstudytohismemoryinrecognitionofhisfriendshipandexemplaryprofessionalcareerinherpetologyandecology.

Literature Citedbaldridge,R.s.andD.W.Wivagg.1992.predationonimportedfireantsbyblind

snakes.Texas Journal of Science44:250–252.

blair,W.F.1960.The Rusty Lizard: A Population Study.Universityoftexaspress,Austin,texas.

Collins,J.t.andt.W.taggart.2009.Standard Common and Current Scientific Names for North American Amphibians, Turtles, Reptiles, and Crocodilians.6thed.CenterFornorthAmericanherpetology,lawrence,Kansas.

Fitch,h.s.andh.W.Greene.1965.breedingcycleoftheGroundskink, Lygosoma laterale.University of Kansas Publications, Museum of Natural History15:565–575.

Gehlbach,F.R.1994.The Eastern Screech Owl: Life History, Ecology, and Behavior in the Suburbs and Countryside(2nded.2008).texasA&mUniversitypress,Collegestation.

Gehlbach,F.R.2002.Messages from the Wild: An Almanac of Suburban Natural and Unnatural History.Universityoftexaspress,Austin.

Gehlbach,F.R.andR.s.baldridge.1987.liveblindsnakes(Leptotyphlops dulcis)ineasternscreechowl(Otus asio)nests:Anovelcommensalism.Oecologia71:560–563.

landsberg,h.e.1981.The Urban Heat Island.Academicpress,newYork.

minton,s.A.,Jr.1968.thefateofamphibiansandreptilesinasuburbanarea.Journal of Herpetology2:113–116.

teXAsheRptoFAUnA

texasblindsnake(Rena[Leptotyphlops]dulcis)emergingfromthesoil.

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Red milk snake Taking large Prey late in the season

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photographsbytheauthor.

Redmilksnakes(Lampropeltis triangulum syspila)aresecretiveandlittleisknownabouttheirbehaviorwhencomparedtomanyother

northAmericansnakes(ernstandernst2003).theyaredietarygeneral-istsandareknowntopreyonmammals,birds,lizards,snakes,reptilianeggs,amphibians,fish,earthworms,slugs,andinsects(seereviewinernstandernst2003). milksnakeshave relatively smallheadsandmouthsandnarrowbodies,andtheygenerallyappeartoprefersmall,narrowprey(FitchandFleet1970,ernstandernst2003).Lampropeltis t. syspila(totallength[tl]144–812mm)issmallerthanL. t. triangulum(tl175–1140mm;Williams1988)andpresumablyeatssmallerprey.AmongthemammalianpreyrecordedforL. triangulumarevoles(Clethrionomys gapperi, Microtus ochrogaster, M. pennsylvanicus, M. pinetorum;seereviewinernstandernst2003).Fitch(1999)reported36preyitemsfrom19L. t.syspila and8scats.nineofthepreyitemswereMicrotus ochrogaster;8ofthosewerespecificallyidentifiedasnestlingsandsizewasnotspecifiedfortheother. on28september2007at1433h,IinvestigatedarodentdistresscallinUnionCounty,Illinois.thecallcamefrombeneathleaflitterinawoodedareanexttoagravelroadbetweenlimestonebluffsandswamp.Isawaleafmoveandliftedit,revealinganadultL. t. syspilawiththreecoilsaroundanadultWoodlandVole(Microtuspinetorum).theheadofthevoleprotrudedfromthecoils,butthesnake’sheadwasnotvisible.At1436h,thesnakestartedsearchingfortheheadofthevoleand,at1439h,itgraspedtherostrumofthevoleandbeganswallowing.Fourpeoplequietlyobservedtheeventandtookflashphotographsasunobtrusivelyaspossible.thesnakeremainedintentonconsumingitsmealdespitethedisturbance. thesnakefinishedswallowingthevoleat1519h(40minswallow-ingtime)andbeforethepreyhadreachedthesnake’smidbody,itstartedcrawlingoutofsightintotheleaflitter.Duringtheevent,wenotedthatthesnakehadashriveled,blindlefteye.theweatherwassunnywithanambientairtemperatureofapproximately30°C. thesnakewasnotrestrainedinordertoallowittofinishitsmeal,andsothelengthandweightofthesnakeandthevolewerenotrecorded.thevolewasidentifiedusingfeaturesvisibleinthephotographs.Microtus pinetorumhasauniformreddish-blackdorsumandashorttail(18–24mm)relativetoitshindfeet(approximately16mm).thehabitat(leaflitterinoak,hickory,andmapleforests)alsowasconsistentwiththatusedbyM. pinetorum(hoffmeister2002). thisfeedingeventwasnotablylateintheyearforaL. t. syspilatobeconsumingprey.FitchandFleet(1970)recorded20naturalfooditemsforL. t.syspilainnortheasternKansas.AllwereconsumedinlateApril,may,orearlyJune,withtheexceptionofonePlestiodon fasciatusinasnakecapturedon25september1964.tomyknowledgethepres-entaccountisthefirstunambiguousreportofL. t. syspilaconsuminganadultM.pinetorum.

AcknowledgementsIthankmichaelDloogatchforreviewingthisnoteandAmysullivanforidentifyingthevole.

Literature Citedernst, C.h. and e.m. ernst 2003. Snakes of the United States and Canada.

smithsonianInstitutionpress,Washington,D.C.

Fitch,h.s.1999.A Kansas Snake Community: Composition and Changes over 50 Years.KriegerpublishingCompany,malabar,Florida.

Fitch.h.s.andR.R.Fleet.1970.naturalhistoryofthemilksnake(Lampropeltis triangulum)innortheasternKansas.Herpetologica24:387–396.

hoffmeister, D.F. 2002. Mammals of Illinois. University of Illinois press,Champaign.

Williams,K.l.1988.Systematics and Natural History of the American Milk Snake,lampropeltistriangulum.2ndrevisededition.milwaukeepublicmuseum,milwaukee,Wisconsin.

AdultRedmilksnake(Lampropeltis triangulum syspila)constrictingandswallowingalargeWoodlandVole(Microtuspinetorum).

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observations on a Wild King cobra (Ophiophagus hannah), with emphasis

on foraging and DietDhirajbhaisare1,VipulRamanuj1,p.Gowrishankar1,m.Vittala1,mattGoode2,andRomWhitaker1

1AgumbeRainforestResearchstation,suralihalla,Agumbe,shimogaDistrict,Karnataka,India(arrs.india@gmail.com)2schoolofnaturalResources&environment,UniversityofArizona,tucson,Arizona85721(mgoode@ag.arizona.edu)

since2005,wehaveconductedresearchonKingCobras(Ophiophagus hannah)intheforestsandplantationssurroundingAgumbeRainforestResearchstation(ARRs)locatedintheWesternGhatsofsouthernIndia.Inmarch2008,westartedaradiotelemetryproj-ectandtodatewehavetrackedfouradultKingCobras(threemales,onefemale)forvaryingperiods.AlthoughwehavedocumentedanumberofinterestingfacetsofKingCobrabiology,thisnoteisconfinedtoobservationsmadeonasinglemale(m2)fromJune–november2009.Duringthistime,weobservedm2capturingandconsuming26pitvipers(twohump-nosedpitVipers,Hypnale hynale,and24 malabarpitVipers,Trimeresurus malabaricus).Weprovidedetailsoftheseremarkableobservations,andbrieflydiscusstheminthecontextofourongoingeffortstounderstandandconservethesecharismaticserpents.

KingCobrasinsouthernIndiaaredistributedalongthelengthoftheWesternGhats(thechainofhillsjustinlandfromthewesterncoast-

lineofpeninsularIndia),fromthesouthernmosttip,calledKanyakumari(blackRockestate),northtoGoa(bondlaWildlifesanctuary).therange

ofKingCobrasinthispartofIndiacoincideswithareasofextremelyhighrainfallandcorrespondingwetdeciduous,riparian,andevergreenrainforest(smith1943).theKingCobraisuncommonthroughoutitsrange,andmostofitsoriginalhabitathasbeendestroyed.Inaddition,KingCobrasare

Radiotrackingintherainforestischallenging,withvegetation,hillocks,andevenheavyraininterferingwithsignals.

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typicallykilledonsightthroughoutmostoftheirrange.however,hinduslivinginthemalnadregionofKarnatakastate,includingourstudyarea,thevillagesandReserveForestsaroundAgumbe,actuallyworshipKingCobrasandthesnakesarerarelyharmed,eveniftheyenteraresidence.thistolerantattitudeforthelargestvenomoussnakeintheworldhasnotonlyfacilitatedourresearchandeducationalefforts,butalsohasprovideduswithover100opportunitiesto“rescue”KingCobrasfromhumanhabi-tations(shankar,inprep.). Asinferredbytheirscientificname,KingCobrasfeedprimarilyonsnakes,includingtheirownspecies(wehaveobservedintraspecificpreda-tioninbothcaptiveandwildindividuals).theonlyotherpreyreliablyreportedfromthewildweretwospeciesofmonitorlizards(Varanusspp.),theCommonIndianmonitor(V. bengalensis;smith1943)andtheWatermonitor(Varanus salvator;C.traeholt,pers.comm.).twoofus(RWandpGs)haveobservedKingCobrasoverthepastseveraldecades,andwehavefoundthatadultsnakes(meantotallength[tl]ofmales~300–400cm;females~225–275cmtl)intheWesternGhatspreymainlyontheubiq-uitousIndianRatsnake(Ptyas mucosa;meantl~175cm,maximumtl~350cm,meanmass~1,250g).AlthoughwecommonlyobserveIndianRatsnakesinopenandswampyareasofbothcultivatedandwildland,werarelyobservethemindeepforest.UnusuallylargemaleKingCobras(>400cmtl)areknowntopreyonIndianpythons(Python molurus);oneofus(RW)wasshownphotosofamaleKingCobrakillingandswallowingamedium-sizedpython(~210–240cmtl),anda366-cmindividualkilledandswal-loweda280-cmReticulatedpython(Python reticulatus,shebbeare1947). somespeculation(b.G.Fry,pers.comm.)suggeststhatKingCobrasmaybeimmunetothevenomofviperidsandelapids,andthattheymay

takespecial“precautions”whenfeedingonpreyspecieswithlongfangs.ZoosaroundtheworldthatexhibitKingCobraseitherconditionthemtoeatdeadrodentsthathavebeenpreparedwithodorsfromsnakesand/orfeedthemsnakesthathavediedonexhibit.thishasresultedinalonglistofvenomousandnon-venomoussnakes,someasexoticasAfricanGaboonVipers(Bitis gabonica),thathavebeenfedtocaptiveKingCobras(s.pfaff,pers.comm.).however,anunderstandingoftrophicecologybasedoncaptivefeedingrecordsislessthansatisfactory.KingCobrashavebeenobservedfeedingonbandedKraits(Bungarus fasciatus)inthewild(saravanakumar,pers.comm.),andhereinwedescribewhatappearstobeselectivehuntingofvenomouspitvipers.

MethodsWeobtainedallfeedingobservationsreportedherewhileradiotracking“m2,”amale(tl=321cm,mass=4,900g),intothecoelomiccavityofwhichweimplanteda25-gtemperature-sensitiveradiotransmitter(modelAI-2t,holohilltd.,ontario,Canada).Wereleasedthesnakeonedayafterthesurgeryon28march2009atitsexactpointofcapture(~1kmfromtheAgumbeRainforestResearchstation[ARRs]).overthenexttwodays,thesnakemoved2.3km,climbingdownasteepescarpmentthatrenderedradiotrackingimpossible.thesignalwasloston31march,butrecoveredtenweekslateron7Juneafterincredibleperseverancebythetrackingteam.thesnakehadmoved~6kmstraight-linedistancefromwhereitwaslost,andwehavetrackedhimeverydaysince10June. Weradiotrackedm2eachmorning,stayingwithhimthroughoutthedayuntilnightfall.ondayswhenthesnakedidnotmove(i.e.,20con-secutivedayswhenshedding),westillobtainedafixeveryhour.Ateachfix,werecordedasuiteofenvironmentalparameters(e.g.,ambientandbodytemperature,relativehumidity,cloudcover),Gpscoordinates,andhabitatvariables(e.g.,canopycover,dominantplantspecies,sheltertype).Ifthesnakewasmovingorvisiblewhenlocated,wemadeeveryefforttoobserveitsbehavior.Althoughourinitialprotocolwasnottogettooclosetothesnakeforfearofinfluencingnaturalbehaviororcausingittomoveorhide,wequicklylearnedthatm2didnotseemtobedisturbedbyobserverswithinafewmeters,aslongastrackersremainedfairlystill.theabilityto

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closelyapproachthesnakehasenabledustomakethedetailedobservationsreportedoninthispaper. ourdataindicatethatKingCobrasareprimarilydiurnal,although,onafewoccasions,wetrackedanimalsafterdarkandrecordedsomemove-ments,onceuntil2200h,butthatwasexceptional.m2almostalwaysfoundasheltersitewellbeforesunsetandsometimesasearlyas1400h.Dependingonweather,suchasrainfall,ambienttemperature,andavail-abilityofsunlight,m2typicallyemergedfromhisrestingplacearound0900h(range=0830–1200h).Wealwaysstayedwithintherangeoftheradiotransmittersignal,becauseevenshortmovementscouldresultinlos-ingthesignalinthedenserainforestenvironment.Inaddition,thisallowedustoinferchangesinbehaviorofthesnakebymonitoringvariationsinsignalintensityandrate.Whenfeasible,wekeptthesnakeundersporadicline-of-sightobservationusingbinoculars.Wetookdetailednotesdescrib-ingallmovementandbehaviorsobserved,includingwhatappearedtobeobviousforagingbehaviorasrevealedbyincreasedtongue-flickingandapparentscent-tracking,presumablyinresponsetopreyodors.

Results and DiscussionDuring2008,wetrackedourfirstadultmaleKingCobra(m1)formorethan100kmoveranine-monthperiodthroughawiderangeofland-scapes,muchofithuman-dominated(e.g.,arecanut[betelnut]planta-tions,ricefields,andcommunityforests).Unfortunately,wesubsequentlylosthim.however,thosedataandothersfromover100“rescued”snakessuggeststhatatleastsomeKingCobras“prefer”human-dominatedland-scapes.perhapsKingCobrasaredrawntoman-madefeaturesbecausetheyattractrodents,whichinturnprovideforanincreasedrelativeabundanceofIndianRatsnakes. Duringthefirsttwomonths,wetrackedm2throughamixofdenseintactforest,degradedcommunityforests,andrarelyplantations.however,attheonsetofthemonsooninJune,m2movedintodeepforestnotfarfromARRs,wherehehasremained,exceptforonebriefperiodof22dayswhenheretreatedintoatermitemoundnearfarmlandson20september,emergingnewlyshedon12october.between7Juneand22november,m2moved35.6kmthroughanarea(whichwearetentativelyreferringtoashis“homerange”)totaling6.3km2. Wenaivelyassumedthatm2wouldnotbeveryactiveduringthemonsoon,becauseannualrainfall inAgumbeaveragesover7,000mmandhaspeakedatover11,000mmduringthefour-monthmonsoonsea-soninJune–september.Weassumedthatheavyandpersistentrainfallwouldcausefeedingtotaperoffuntilthereturnofwarmer,dryerweather.however,m2wasfirstobservedfinding,killing,andeatingamalabarpit

Viper(Trimeresurus malabaricus)on6Julyattheheightoftherainyseason.overthefollowingfourandahalfmonths,weobservedm2eatingatotalof26pitvipers,twoofwhichwerediminutivehump-nosedpitVipers(Hypnale hynale). Whetherornotm2movedonanygivendayusuallydependedonwhetherornotitwasraining,rainfallintensity,andpresenceorabsenceofdirectsunlight.ondrier,sunnierdays,m2typicallybaskedfor15–45minbeforebeginningtohunt.Foragingactivityconsistedofapparentlypurposefulsearching,characterizedbyconstanttongue-flickingandactiveprobingofcrevices,burrows,treebuttresses,andvegetativedebris.Datafromdaysonwhichweobservedm2feedingononeormorepitvipers(n=23),hespentfrom37minto>7hperdayforaging(mean=3.5h/day). Whenm2apparentlydetectedanodortrailofapitviper,hisforagingactivitybecamevisiblymoreintense,sometimesinducing“headjerking,”followedbysustainedtongue-flickingalongthepresumedodortrailuptoaheightof20minthecanopy.onseveraloccasions,pitvipersseemedalarmedbythecloseapproachofm2,causingthemtofallorlaunchfromtheirperch(malabarpitVipersareprimarilyarboreal,pers.obs.).Acon-siderableamountoftimespentforagingbym2consistedofclimbinginlowerandmid-canopy.ofthe24malabarpitVipersconsumedbym2,17(70.8%)werefoundandcapturedintrees,shrubs,vines,ordeadbranches,andseven(29.2%)werefoundandcapturedontheground. ourobservationssuggestthatKingCobrasareabletodetectmove-mentfromrelativelylongrange,perhapsasfaras50–100m.WehaveobservedKingCobrasintentlywatchingusaswemovedthroughdenseforestthickets.WealsohaveobservedaKingCobrathatappearedtobewatchingaflockofheronsflyingoverhead.however,theirvisualacuitymaybepoor,astheydonotseemtoeasilyrecognizestationaryobjects,betheypreyanimalsorpotentialthreats(suchasourselves).India’sfor-estpitvipersaretypicallynocturnalambushpredators,remainingseden-tarybyday(pers.obs.).Indeed,wehaveobservedpitvipersintheexactsamelocationandevenpositionforasmanyassevendays.KingCobrasareunlikelytoseeamotionlesspitviper.Infact,weobservedm2directlypursueandsubduepitvipersthathehaddisturbed,causingthemtomove.Whenunsuccessfulatcapturingthepitviper,m2reliedonchemosensorysearchingtofollowtheodortrail.Weobservedm2losetheodortrail,suchaswhenthepitviperfellintoorcrossedoverwater,buthepersistedinhissearchuntilhepickedupthetrailagain.Remarkably,heevenseemedabletosuccessfullyfindandfollowodortrailsinheavyrain.Althoughpitviperswouldoftenremainmotionless,theywouldinvariablymovewhenm2gottooclose,therebycausinghimtostrikeimmediately. Inonestartlingtwo-hourepisoderecordedonvideo,weobservedtheKingCobralocatingalargegravidfemalemalabarpitViper,causingittofallfromitsperchabout15mabovethegroundandtrytoevadecapture.

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m2promptlydescendedandfollowedthepitviperodortrailthreetimesaroundandacrossalarge,swiftstreambeforefinallycatchingandkillingit.Althoughthepitviperwaseasilyvisibletous,m2passedclosetohispreyseveraltimes,apparentlysointentonpickingupolfactorycuesthathefailedtoseethesnake.Atonepoint,theodortrailwasstrongenoughtoinducem2tobiteastonetwice,providingfurtherevidenceofthepromi-nenceofolfactionovervisionundersomecircumstances. Wealsodocumentedinterestingpreyhandlingbehavior.In16oftheobservedpreycaptures,oncem2sawthepitviper,heapproachedtheheadend(malabarpitVipershaveanarrowneckandwide,triangularheadcharacteristicofviperids)andgraspedthesmallersnakeeitherjustbehindordirectlyonthehead.ourobservationsofKingCobrasfeedingonIndianRatsnakesindicatethattheyarenotparticularaboutwhichpartofthebodytheygrab,eventhoughretaliatorybitesfromIndianRatsnakes,includingontheheadoftheKingCobra,commonlyoccur.perhapsm2wastryingtoavoidbeingbittenbythepitvipers.Innineofthepreda-tioneventsweobserved,theKingCobrareceivedwhatappearedtobeapenetratingretaliatorybite.WhilethescalesofaKingCobraprobablypre-ventpenetration,apitviperpresumablycouldpiercetheinterstitialskinbetweenscales.Whenbittenbypitvipers,m2invariablydroppedhisprey,waitinganaverageof10minuntilthepitviperwasimmobilizedfrombeingenvenomated.AlthoughmalabarpitVipervenomhasrelativelylowtoxicity(Dhananjaya,pers.comm.),thebiteofthehump-nosedpitViperhascausedkidneyfailureandoccasionaldeathsinhumansinsrilanka(desilva1990). Weobservedm2capturingalarge,gravidfemalepitviper,graspingitatmidbody,onlytoreceivewhatappearedtobeapenetratingbiteonthesideofthehead.m2immediatelyreleasedthepitviperand,withheadand

neckerectuptoapproximately50cm,appearedtowatchtheenvenom-atedpreyclimbanearbybranchtoaheightofabout6m.m2remainedmotionlessinthispositionforthenexthalfhour,mouthpartlyopen,with-outtongueflickingorreactinginanyway.eventhepitviper,fallingdeadfromabranchafter20min,approximately1maway,andtheapproachofthetrackerwithinapproximately3mtocheckforswellingatthesiteofthebite(nonewasevident),didnotelicitaresponse.After30mininthisuprightposition,m2cameoutofhisapparent“trance”andimmediatelystartedsearchingforthepitviper,ignoringthedeadsnakeinfrontofhimandinsteadclimbing(allthewhilevigorouslytongueflicking)tothepointwherethesnakefell.hethenreturnedtothegroundtoeventuallyfindandswallowthedeadsnake. themaximumtimeforapitvipertobeconsumed,startingwiththepointatwhichitwasbittenandendingwhenitwasswallowed,was67minandtheminimumtimewas4min(mean=14min).In16predationevents,m2graspedthepreybytheheadorneckand,inafewcases,immediatelyswalloweditalive,completingthetaskwithinafewminutes.Inthecaseofasmallhump-nosedpitViper(meanadulttl~25cm,meanmass<40g),m2simplygraspedthesnakeatmidbodyandswalloweditwhileitwasdoubledover,withoutbotheringto“jaw-walk”towardsthehead.Fourpitvipersweregraspedatmidbody,threebythetail,andwefailedtoobservethreeothercaptures.elevenofthesnakeswerestillmovingorclearlyalivewhenswallowed,12wereimmobileandthreewefailedtoseeclearly. Weestimated12ofthepitviperstobe~60cmtl(thesewereprob-ablyallfemales,severalappearedgravid),fivewereapproximately45cmtl(averageadultmalesize),andsevenwereroughly30cmtl.Weobservedm2eatingtwopitvipersperdayonthreeseparatedays.thecobrafre-quentlydefecatedafteraswallowingepisode.

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Duringthisstudy,m2spentapproximately50%ofhisactivehoursforaging/huntingforprey,inthiscasepitvipers.ononeoccasion,hespentoverfivehourscontinuouslyforagingbeforehelocatedapitvipertoeat.hewasnotobservedsearchingfor,pursuing,orcapturinganyotherspeciesofsnakeduringthistime,despitethepresenceofotherspeciessimilarinbodymasstosmallerpitvipers(seebelow).sincewefollowedm2everydaythroughouthisentirediurnalactivityperiod,wealmostcertainlyrecordedthevastmajorityofpredationeventsoverthefull140-dayperiod.Duringthattime,ourfieldteamwaswithm2forabout8hperdayforatotalof1,120h.Duringthistime,weobservedm2atotalof74.89hoursengagedinobviousforagingbehaviorthatledtoanactualpredationevent.onaver-age,m2consumedapitviperevery5.38days,andspentanaverageof2.88hforagingforeachofthe26pitvipershecaughtandconsumed. InJuly–november,m2primarilyrestrictedhisactivitytodeepfor-est.thisperiodoftimecoincidedwithheavyrainfallintheAgumbearea.Duringrainyperiods,werarelyobservesnakesspeciesthatareknowntobetypicalpreyofKingCobras(e.g.,IndianRatsnakes).Inaddition,wehaveobservedIndianRatsnakesfarmorefrequentlyinopenmead-owsandagriculturallands,presumablywithhighpreydensities,thanindenseforest.Incontrast,pitvipersareapparentlymoreactiveduringthemonsoonseason,asevidencedbyourincreasedobservations.othersnakespecies,whicharetypicallyvisibleduringtherainyseasoninthevicinityofAgumbe,arebeddome’sKeelback(Amphiesma beddomei),large-eyedbronzeback (Dendrelaphis grandoculis), and theCommonVinesnake(Ahaetulla nasuta);however,wedidnotobservem2eatinganyofthesespeciesduringthetimeinwhichheconsumedthe26pitvipers.Althoughpurespeculation,perhapsKingCobrasasaspecies(orm2asanindividual)

preferpitvipers,ortheodortrailofpitvipersissomehoweasiertofollowthanthoseofothersympatricspecies. Alsointerestingtoconsideriswhetherornotthepitvipersthatm2consumedcouldprovidehimwithenoughenergytomaintainhishealth.Whenwecapturedm2inearlymarch,heweighed4,900g,whichisanaveragemassforahealthy321-cmKingCobra.Ifm2consumedanaver-age-sizedadultIndianRatsnake(~175cmtl,~1,250g)hisbodymasstopreysizewouldbeapproximately4:1.AsmallsampleoftenmalabarpitViperscapturedinthevicinityofARRsyieldedanaveragetlof45cmandanaveragemassof35g.Althoughwewereunabletoobtainmassesonthepitvipersconsumedbym2,weestimatedtherangeoftheirmassestobe30–60g,whichcorrelatestoanaveragemassofapproximately50g.Usingthis50-gestimate,themassratioofpredatortopreyisroughly98:1.Ifweaddtogethertheaverageapproximatedweightsofall26pitvipersconsumed,thepredator-preymassratiois3.77:1,whichisapproximatelyequaltothatofasingleIndianRatsnake.oneobviousquestionarisingfromthisexerciseiswhetheratotalpitvipermassof1,300gissufficientnutritionforaveryactive5-kgKingCobraoveraperiodof18weeks,nottomentionapresumedincreaseinpredationriskwhilemoving. basedonourobservations,m2apparentlyexpendedagreatdealofenergysearching,climbing(observedalmosteveryday),andpersistentlyandrepeatedlyfollowingpitviperodortrailsforwhatseemedtobeamea-gerreward.however,m2didnotappeartohavelostasignificantamountofmass,nordidhisbodyconditionappeartosuffer.nevertheless,smallsnakessuchaspitvipers,withtheattendantneedforlargeenergyexpendi-turetofindthem,areunlikelytofulfillthenutritionalrequirementsofsuchalarge,activesnake.

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Whetherornotm2willalterhisforagingtacticsduringtheupcomingmatingseason,whenKingCobrasofbothsexesbecomemuchmoreactive(judgingfromthedramaticincreasein“rescue”callsreceivedduringtheperiodofJanuary–may),remainstobeseen.perhapsm2willmoveintohuman-dominatedlandscapes,suchasplantationsandcropfields,whererelativeabundanceofIndianRatsnakesisapparentlymuchhigher.

AcknowledgementsGerrymartin,nikWhitaker,Charliepainter,loriKing,Aniruddhabelsare,Gowrimallapur,brijeshRaj,sandeshKadur,samirWhitaker,p.prashanth,R.sharmila,srinidhiKashyap,citizensofAgumbeVillage,andanumberofvolunteersputindaysofhardworkbrainstormingandprovidedlogisticalsupport,helpinthefield,andveterinarywork.We

aregratefultotheKarnatakaForestDepartmentfortheircollaborationandenthusiasticinterestinthisproject.WearegratefultothenationalGeographicsociety,DisneyWorldwideConservationFund,mohamedbinZayedspeciesConservationFund,RiverbanksZooandGarden,andGladysporterZooforprovidingfinancialsupport.

Literature Citeddesilva,A.1990.Colour Guide to the Snakes of Sri Lanka.R&Apubl.,UnitedKingdom.

Dunn,R.D.andK.W.brady.2001.snakeinhibitorsofphospholipaseA2enzymes.Biochimica et Biophysica(55803):1–9.

shebbeare,e.o.1947.themalayanCobras.Malayan Nature Journal2(2):25–32.

smith,m.A.1943.Fauna of British India, Vol. III. Serpentes.taylorandFrancis,london.

bhAIsARe

Googleearth®imageshowingsitesatwhichm2wasrelocated.ARRs=AgumbeRainforestResearchstation.

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CasaVerde, theweathered,palm-sheltered former fieldheadquar-tersoftheCaribbeanConservationCorporation’svenerableGreen

turtleresearchprogram,andnowitsmodernizedsuccessor,theJohnh.phippsbiologicalFieldstation,standsquarelybetweentwogreatbiologi-calworlds.totheforestretchestheCaribbeanseawithallitsteemingmarinelife.Itsedgehereistheblacksandbeachoftortuguero,namedforthesmallcoastalvillageadjacenttothestation,andthemostimportantseaturtlenestingsiteintheentirewesternCaribbean.totherearlietherichCostaRicanlowlandrainforestandacomplexnetworkoffreshwatercanals,rivers,andswampsthreadedthroughitsverdure. For thebiologist, few ifanyotherplacesonearthoffer somanyopportunities,thenatureofwhichdependssimplyuponthedoorfromwhichoneemerges.ArchieCarrandthelegionofbiologistsuponwhomthereputationoftortuguerohasbeenbuiltwentoutthefrontdoortothedarkbeachesandtheirdrovesofnestingGreensandleatherbacks.Instead,Igooutthebacktostudyanequallyfascinatingbutpoorlyknownassemblageofsmallerturtlesresidentintortuguero’sfreshwaterandforest

habitats.longignoredandovershadowedbythehuge,spectacularlyabun-dantseaturtles,thesenon-marinespecieshaveunobtrusivelygoneaboutthebusinessoflivingwithoutmuchattentionfrombiologists.however,recentinvestigationsthatIhaveconductedontheirecologyhaverevealedmuchofinterestinthesesmallpackages.theirnaturalhistory,uniqueandworthyofrecordforitsownsake,alsoprovidesinsightintotheoriginoftheneotropicalnon-marineturtlefaunaandtheevolutionarystrategiesthatallowitssurvival.oneofthefreshwaterspecieshasledmethroughaproverbialrevolvingdoor.Inspiteofmyoriginalintentionstostayinthejungle,ittookmebacktothebeachtoobserveacriticalphaseofitslifehistoryinthetortugueroecosystem. studyingtheturtlesoftortugueroisasingularexperienceinitself.Cruisingthegreen-walledstreamswhilefour-footiguanaslaunchthem-selvesfromoverhangingbranches,andparrots,toucans,andhowlermon-keysscold,gurgle,androaroverheadaddsadimensiontoturtletrappingthatcan’tbematchedinthetemperateZone.thepresenceofcaimans,bullsharks,andstingraysintherivers,andJaguars,fer-de-lances,androv-

The backdoor Turtles of TortugueroDonmoll

Departmentofbiology,missouristateUniversity,springfield,missouri65897(donmoll@missouristate.edu)

photographsbytheauthor.

T R a v e l o G u e

Adultmalemeso-Americanslider(Trachemys venusta),withacarapacelengthofabout40cm,onthebeachattortuguero.

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ingherdsofWhite-lippedpeccariesintheforestalsocontributespicetotheambience. oureffortstodatehaveyieldeddataconcerningfiveofthesixspe-ciesinhabitingthetortugueroarea.therarenarrow-bridgedmudturtle(Kinosternon angustipons)occasionallyturnsupattortuguero,butwehavenotyetlocateditsapparentlyratherspecializedhabitat.theobservedspe-ciesrangeacrossthenichespectrumfromthealmosttotallyaquaticsouthAmericansnappingturtles(Chelydra acutirostris)andmeso-Americansliders(Trachemys venusta),whichemergefromwateronlyforegg-laying,occasionalmovementintootheraquatichabitatsand,inthelatter,forsun-bathingonemergentlogs,tothesemiaquaticWhite-lippedmudturtle(Kinosternon leucostomum)andblackRiverturtle(Rhinoclemmys funerea),whichspendsignificantamountsoftimebothinwaterandwanderingontheforestfloor,tothefullyterrestrialbrownWoodturtle(Rhinoclemmys annulata),whichenterswateronlytodrinkorsoakinforestpuddles. besidesthesemoretraditionaltaxonomicandhabitatapproachestoclassifyingtortuguero’sturtles,ecologicalinformationmayhelptocat-egorizethemaccordingtotheirrespectivebiogeographichistories.Whilealloftortuguero’sturtlesarecertainlytropicalintermsoftheircurrentgeography,theirsurvivalheredependsuponstrikinglydifferentadaptiveapproachestocopingwiththetropicalenvironment.thesevaryingstrate-giesprovidecluestotheirpast.Fromthisperspective,twooftortuguero’sturtlesseemoutofplaceindeed. Reproductivepatternsareespeciallydiagnosticinthisregard.thetwomudturtlesandtheblackRiverandbrownWoodturtleslayoneoronlyasmallclutchofsmall(mudturtles)torelativelyverylargeeggs(blackRiverandbrownWoodturtles)ontheforestfloor,eitherasynchronouslythroughouttheyearoroverextendedperiodsduringtherainyseason.theycharacteristicallymakelittleornoefforttoburytheeggsotherthanscrap-ingalittleleaflitteroverthembeforetheypermanentlyleavethescene.thisreproductivepatternhasbeenobservedinsmalltomedium-sizedrainforest-dwellingturtlesinboththenewandoldWorldtropicsandisconsideredtobeindicativeoflong-termtropicalresidency—perhapsanadaptiveresponsetothemyriadeggandhatchlingpredatorsthatroamtherainforestfloor.Unpredictabilityinlayingtimeandlocationdiscouragespredatorsfromgangingupontheturtles’reproductiveeffortastheycan,forexample,onthenearbyGreenseaturtlenestingbeach. Alltheseaturtles,andtheworld’stemperateZonefreshwaterandterrestrialspecies,practiceaverydifferentnestingstrategy.theseturtlestypicallylayoneormorelargeclutchesofrelativelysmalleggsincarefullyconstructedsubterraneanneststhataresealedaftereggdeposition.nestingtakesplacewithinadistinctseasonandnestsoftenareconcentratedinsuit-ableareasthatprovidebotheasyaccessfornestingfemalesandthepropermicro-environmentforthedevelopingembryos.Aspredatorsoftenhome

inonthesenestingareaswithgreatefficiency,thesespeciesaremorelikelytosurviveasaresultoftheirabilityto“swamp”predatorsthroughtheirhighannualfecundityandthe longreproductive lifeofadult females.thesnappingturtlesandslidersoftortugueroreproduceinthistypicaltemperateZonepattern. tortuguero’scrazy-quiltamalgamofspecieswithsuchdivergentnest-ingstrategiesisbestexplainedbytherelativelyrecentinvasionofthecloselyrelatedancestorsofthelattertwospeciesintotheneotropicsfromtemper-atenorthAmerica.WithclosureoftheCentralAmericanisthmusduringtheplioceneandthesubsequentclimaticturmoilofthepleistocene,thetimewasripeforfaunalexchangesbetweenthetwocontinents.Duringdrierperiodswhentheforestshrankandrainforestspecialistswereatadisadvantage,theopportunisticsnappingturtlesandslidersprobablycolo-nizedCentralandsouthAmerica.today,thesnappersrangeasfarsouthasecuadorandthesliders’rangeextendsintotemperateArgentina.Duringwetterperiods,asinthecurrentinterglacial,however,theseJohnny-come-lateliesareapparentlyatadistinctdisadvantageinthere-coalescentrain-forest,astheirdistributionsareextremelydisjunctthroughthisvastareaandtheirpresentpopulationsizesoftenarelowcomparedtothetemperatehabitatsinwhichtheyoftendominatetheturtlefauna.Characteristically,bothspeciesarerelativelyuncommonattortuguero.onefocusofmyworkwastodeterminehowthesespeciesliveattortuguero,which,withtheexceptionoftheareaaroundthevillage,isstillcharacterizedbyprimaryrainforestasdenseasthatanywhereintheneotropics. Currently,twotropicalspeciesinthegenusChelydraandasmanyasninemainlandlatinAmericanspeciesinthegenusTrachemysarerec-ognized—allverycloselyrelatedtotheirtemperateZoneancestors,theCommonsnappingturtle(Chelydra serpentina) andtheslider(Trachemys scripta),respectively. Althoughlittlecomparativeinformationwasavailableconcerningtheecologyofsnappingturtlesinothertropicallocations(i.e.,limitednaturalhistoryobservationsinChiapasbymiguelAlvarezdeltoroandinColombiabythelateFedericomedemareavailable),thecollectivedatafromseveralstudiesofslidersintropicalmexico,belize,Colombia,Venezuela,andespe-ciallypanamaindicatedaremarkablysimilarecologytothatofpopulationsintemperatenorthAmerica.observedecologicaldifferencesinthetrop-ics(e.g.,greatersize,annualeggproduction,longernestingseasons,etc.)weremoreamatterofdegree,largelyrelatedtogreateropportunitiesforforaging,growth,andreproductionintheyear-longwarmthratherthansubstantivechangein lifestyle.WhileI foundessentiallythesamepat-ternsattortuguero,theslidersherewereuniqueintwocharacteristicsthatwarrantedfurtherstudy.First,theyarehuge,evenbytropicalsliderstan-dards,withsomefemalesattainingcarapacelengthsof44cmandweightsupto10kg.second,circumstantialevidencehadaccumulatedthatslidersusedCaribbeanseabeachesfornesting—averyunslider-liketrait,and

Ahatchlingslideratanestingsiteincocoplumscrubalongthebeachattortuguero.

Aslidernestingtrackontortuguerobeach.

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knowninonlyahandfulofothernon-marineturtles.turtlebiologistpeterpritchardhadpreviouslyobservedthatnon-marineturtletracksobservedontortuguerobeacheswerealwaysthoseofadultsliderfemales(whicharemuchlargerthanmales)andthatthediscardedcarcassesofslidersthathadbeencollectedonbeachesforfoodbytortugueroresidentswerealsoadultfemales.theseobservationssuggestedthatbeachedsliderswerenotjustwaifswashedoutofthetortugueroestuaryduringfloodperiods.Inaddition,inJune1989,Jimspotila,tomYocky,andIweresearchingforleatherbacknestsalongtortuguerobeachwhenweencounteredasmallstrangetrackthatwefollowedtoasinglesliderhatchlingonthelowerbeachnearthesurfline.Webacktrackedittoitsprobablenestingsiteinthecocoplumscruboftheupperbeachberm.theseobservationsintriguedme,andIresolvedtostudytheecologyofsea-beachnestinganditsadaptivesignificanceforslidersinthetortugueroecosystem.Iwasparticularlyinterestedinwhy,ofallthesliderpopulationsacrossthevastrangeofthesespecies,sea-beachnestinghadevolvedonlyhere,andwhetherthehugesizecharacteristicoftortuguerofemaleswassomehowlinkedtothisphenomenon. peterpritchard,whohasthoughtaboutmostthings“turtle”atsometimeoranother,hadpreviouslysuggestedthatbeachnestingwasprobablyaresponsetotheneedforopennestingsites,alwaysinshortsupplyinvirginrainforest,andthatlargesizeprovidedphysiologicaladvantagesduringtheperiodofseatravelrequiredtoreachappropriatenestingsites.AstudyoftheecologyofpanamanianslidersbymybrotheredwardmollandJohnleglerinthe1960shadestablishedthatgolfcourses,plantations,andothermoreopensiteswereusedtotheexclusionofrainforestfornestingintheir

CanalZonestudyarea,sothisfirstideamadesense.Asallseaturtlesandthoseotherfreshwaterspeciesknowntoventureintothesea,suchastheAsianriverterrapinsandAsiaticGiantandnilesoftshells,areverylargeanimalsbyturtlestandards,theassociationoflargesizeandseaworthinessalsoheldmerit.mysabbaticalleavefromteachingresponsibilitiesduringaspringsemesterseveralyearsagoallowedmethefreedomtowalkthetortuguerobeachesfromJanuarythroughmarch,thepeakoftheslidernestingseason,toacquiremoreinformation. FringedbyCoconutpalmandseagrape,theblacksandbeachoftortugueroextendsunbroken for35kmfromthetortugueroCanalestuaryinthenorthtotheparisminaRiverestuaryinthesouth.Alwaysbeautiful,itneverthelesscanbeadifficultplacetowork.byday,itquicklybecamelethallyhot,withsandsurfacetemperaturesapproaching70°C.Uponreturningfromearlymorningsurveys,Iusuallyenteredthecoolgloomoftheadjacentforestforarespite—oftenentertainedbywing-snappingmatingdisplaysofWhite-collaredmanakinsandslowlyforagingtroopsofblackhowlermonkeysinthetreetops.onmoonlessnights,thebeachcouldbesoblackastoloseitsfeaturesaltogether,andIwasregularlyupendedbybeachlitterandbysteppingoffnewlywave-cutbenchesthatchangedpositionnightly—but,iftherewerehardships,theyweremorethanoffsetbythelureofdiscoveryandthebeautyofthewildtropicalbeach.onebrightmoonlitnight,forexample,Isatwatchinganesting

sliderinthecocoplumscrubwhileanearly-seasonleatherback,spangledwithphosphorescenceandmoonglow,hauledoutofthesurftonestjustthirtymetersdownthebeach.onanothernight,asIreturnedtoCasaVerdeafterreachingthesouthernterminusofmysurveyroute,InoticedJaguartracksparallelingmyownfootprintsformorethanthreekilome-ters.Itwasabiganimal,apparentlywatchingmecuriously(Iprefernottothinkhungrily)fromthedarknessasItrekkeddownthebeach.Withoutdoubt,experiencessuchasthesewerethe“adventuresofanaturalistonremoteCaribbeanshores”thatArchieCarrhadinmindwhenhewroteThe Windward Roadnearly50yearsearlier. Gradually,throughobservationandradiotelemetricmonitoringofslidersintherivers,inthesea,andonthebeach,acharacteristicallyclock-wise,albeitvariable,nestingpatternemerged.slidersgenerallymovednorthwithrivercurrentsintotheseaviathetortugueroestuaryorbycross-inganarrowstripoflandthatseparatedtheriverfromtheseanorthoftortugueroVillage.theythenmovedsouthinthelongshorecurrentjustbeyondthesurflineuntiltheyhauleduptonestinthecocoplumscruboftheupperbeachafewkilometerssouthofthevillageintortugueronationalpark.spentfemaleswouldusuallyreturntothesea,movingdown-coastforabriefperiodbeforeemergingagaintotrekoverlandacross

tURtlesoFtoRtUGUeRo

AnadultsouthAmericansnappingturtle(Chelydra acutirostris)attortuguero

eggsexcavatedfromasnappernestneartortuguero.

snapperhabitatneartortuguero.

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thepeninsulatothetortugueroCanal.onceinthecanal,theymovednorthagainwiththecurrenttosuitablefeedingandbaskingareas.thepat-ternconservedenergybyutilizingtheprevailingcanalandseacurrents,andgottheturtlestotheappropriatenestingsiteswithlittletimespentonland,animportantpointthatI’lladdressbelow.theupperbeachcocoplumscrubnesting sitesprovidedamoderate thermal environment for eggdevelopmentwellawayfromthehighesttides,aswellasprotectivecoverforthelayingfemaleandemerginghatchlings.sliders,likemostturtles,havetemperature-dependentsexdetermination,inwhichwarmernestsornestdepthsproducefemalesandcoolersitesmales.substrate-temperaturemonitoringindicatedthatopenbeachtemperaturesweretoohotandprob-ablylethaltodevelopingeggswhileforestsoilswererootstrewn,soggy,andprobablytoocoolforadequatefemaleproduction.thecocoplum,however,providedfewobstructions,properdrainage,andtherightther-malregimefortheproductionofbothsexes.thelatterprobablyresultsfromacombinationofsingle-sexandmixed-sexnestsscatteredthroughthecocoplumscrub.Duetotherelativeproximityofthesesuitablebeachnestingsitestofreshwaterhabitats,astraight-linedistanceofakilometerorlessofforestedpeninsulainbetween,thesliderscansurviveattortuguero.Withoutit,asisoftenthecaseinmoreinlandrainforests,theyprobablycannot,andarethereforeusuallyabsentfromtherainforestturtlefauna. thequestionremains,however,whythefemalesliderssimplydon’twalkacrossthepeninsulatodeposittheireggsonthebeach?slidersarefullycapableofwalkingakilometerormoretolayeggsormovingfrompond

topondintemperatehabitats.Whyisitbettertorisktraveloverasubstan-tiallylongerdistancebyseainaphysiologicallydifficultenvironmentwithaformidablesetofpredatorsinordertoreachthebeachnestsite?theanswermaybelinkedtoboththesizeofthetortugueroslidersandtothevulner-abilityoftheirrelativelysmalleggstoadiversearrayofterrestrialpredators.Ithinkslidersherehaveresortedtoseatraveltoavoidleadingterrestrialpredatorstotheirnestsites.thiscouldoccurifpredatorsfollowedafemalewanderingoverlandtothenestingsite(orherscenttrail)orback-trailedhertothesiteasshemoveddirectlytowardthecanalthroughtheforestafternesting.Fromthisperspective,boththelongseatravelphaseinreachingthenestsiteandthereturntotheseaafternestingforabriefperiodbeforedown-coastemergenceandoverlandmovementtothecanalmakesense.Ialsothinkthatgreatsizehasevolvedasaconsequenceofthesea-travelphase.Whileitmaywellconferosmoregulatoryadvantagesbyloweringthebody-surface-to-massratio,workbybillDunsonofpennsylvaniastateUniversityhasshownthatevensmallerturtleshavearelativelylowrateofsodiuminflux.Ithinkthelargesizeherehasevolvedprincipallytopro-videthesea-goingfemaleswithinsuranceagainstthepredationrisksthathavebeenincreasedbytheirrelativelylongnestingmigrationsanden-routeexposuretomarinepredators.theoccasionalmutilatedcarcassesofsliderfemalesI’veobservedwasheduponthebeachunderscoretherealityofthedangers.WhitGibbonshasadvancedthegeneralhypothesisthatlargesizewouldconferanadvantageinreducingpredatoryandenvironmentalriskstoturtlesthatmusttravelfromtherelativesafetyoftheiraquatichome

brownWoodturtlehabitatinsecondarygrowthneartortuguero.brownWoodturtles(Rhinoclemmys annulata)arefullyterrestrial.

White-lippedmudturtle(Kinosternon leucostomum)fromtortuguero. White-lippedmudturtlehabitatattortuguero.

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rangefornestingorotherreasons.Asidebenefitofdecreasedtravelriskswouldbethatmoretimecouldbeexpendedandmoredistancetraveledinseekingthemostoptimalnestingsitesavailable.Ifstrongselectionreducesterrestrialtraveltominimizeeggpredation,thentraveltimeintheseamustincreaseaccordingly.theevolutionofgigantismattortugueroisprobablyaresponsetothesefactors. Ifthebeachesprovidetherelativelyopensitesnecessaryforslidernest-ing,Iwonderedwhatsitesmightbeusedfornestingbyresidentsnappingturtles.mystudiesoftheirecologyattortugueroindicatednosubstantivechangesfromtheecologyoftemperatepopulations.theyweresurvivingattortuguerobylivingtheirusualgeneralistlifestylecharacterizedbyhabitatflexibilityandomnivory.ReproductivestudiesbyRonbrooksinontarioandJustinCongdoninmichiganindicatedthatnorthernsnappers,likesliders,requirerelativelyopennestingsites.Isuspectedthattheywouldneedsuchnestingsitesattortugueroaswell,butIneverencounteredany

ontheseabeachesduringmyslidersurveys.byaccident,however,Istum-bledacrossanestingsnapperearlyoneFebruarymorningonanabandonedmilpa,anareaclearedandplantedforoneormoreyears,thenabandonedandallowedtoreturntoforest.theseareasremainrelativelyunshadedforseveralseasonsandprovideakeyhabitatforthesurvivaloftortuguero’ssnappers.byfocusingmysearchintheseareasadjacenttocaños(smallstreams),Iwasabletodiscoverseveraladditionalsnappernestsoveranextendedperiodofthespringdryseason.thesewereessentiallyidenticalinformandeggcharacteristicstotemperatenests.Attortuguero,aloosesortofmutualismexistedbetweenhumansandsnapperswherethebigtortugaslagartos(“Alligatorturtles”)occasionallywerecaughtforfoodbylocalresidents,and,inturn,wereprovidedwithnestsitesviamilpaagri-culture.perhapssnappersurvivalinthemodernneotropicsisdependentuponsuchhumanactivitieselsewhereinrainforestedareasaswell.treefallsnearwaterwaysmayprovidesimilarnestingopportunitiesbylettingmoresunlightreachtheground,butIneverencounteredanynestingsnappersinnaturalforestopeningsattortuguero. Ithinkoftheslidersandsnappersoftheneotropicsasthe“beverlyhillbillies”oftheturtleworld—truemisfits,butgamelytryingtomakeagoof it inanalienculture.thecollectiveecologicalevidence fromtortugueroandelsewhereindicatesthattheyhavenotyetblendedintotherainforestcommunityduringtheirbriefresidencehere.theirgeneralizedecologyandneedfornon-forestednestingsitesclearlyevolvedtocopewithadifferentenvironmentfarfromtheircurrenthome.theotherturtlesherehavebecomehighlyadaptedforlifeintherainforest.Reproducingandregularlyforagingontheforestfloorandeven,asourresearchindicates,beinginvolvedinsymbioticseed-dispersingrelationshipswithmanylocalplants,thesespecieshavetrulybecomepartofthefabricoftherainforestecosystem.Incontrast,thenewcomershaveextendedtheircharacteristicflexibilitytotheextremetosurvivehereinspiteoftheall-encompassingforest.byusingsuchtricksasbeachandmilpanestingtheyhavemanagedtohangon,barely,bytheskinoftheirjaws,astheforesthasreclaimeditsformerdomainduringthesewettertimessincethelasticeage.Fromthisperspective,theslidersandthesnappersoftheneotropicsmaybeamongthefewspeciestobenefitfromthecontinueddestructionoftherainforest.

tURtlesoFtoRtUGUeRo

thetortugueroRiverprovidedsliderandblackRiverturtlehabitat.tortuguerobeach.

blackRiverturtles(Rhinoclemmys funerea)spendtimebothinwaterandwanderingontheforestfloor.

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AJJAepstudentandahatchlingornateboxturtlewithradiotransmitterattached.

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Turtles in the Dust: effects of Hands-on scientific Training on a Group of behaviorally

at-risk students’ Knowledge and empathyJ.Alansosa1,oscarReyes2,andGadperry1

1DepartmentofnaturalResourcesmanagement,box42125,texastechUniversity,lubbock,texas79409,UsA2lubbockIsD,1909n.Akron,lubbock,texas79415,UsA

photographsbytheseniorauthor.

Abstract.—Wetaughtagroupofbehaviorallyat-riskstudentslessonsonbox-turtlebiologyandnaturalhistorybeforetranslocatinghatchlingornateboxturtles(Terrapene ornata ornata)toanatureareaadjacenttotheirschool.thestudentsthenassistedwithdatacollectionandultimatelyusedtelemetrytore-locatetheturtlesandindependentlycollectdata.eachstudentintheexperimentalgroupwasgivenaquestionnairepriortobeginningandagainaftercompletionoftheprogram,withquestionscoveringbox-turtlebiology,naturalhistory,andempathytowardanimals.studentsinacontrolgroupweregiventhesamequestionnaireatsimilartimes.studentsintheexperimentalgroupfelttheyimprovedtheirknowledgeconcerningbox-turtlebiologyandnaturalhistoryaftertraining,whereascontrolgroupstudentsdidnot.neithergroup,however,showedachangeinempathybetweenthepre-andpost-questionnaires.Incorporatingbehaviorallyat-riskstudentsinresearchprojectsmayenableresearcherstocompleteprojectswhileatthesametimeinvolvinganoften-neglectedportionofthepopulation.

Introduction

Interactionswithanimalsatayoungageimprovesocialdevelopmentandattitudestowardnatureinchildren(myersandsaunder2002).In

addition,companionanimalsintheclassroomareknowntoincreasesen-sitivitytowardhumansandpossiblydecreaseaggression(hergovichetal.2002).Gender,animalownership,andattitudestowardanimalsalsoaffectempathytowardhumans(taylorandsignal2005).however,noconsen-suslinksinteractionswithanimalsandincreasedempathytowardhumans(paul2000,taylorandsignal2005).Wealsoareunsurewhetherinvolvingstudentsinintensive,hands-onscienceprogramswithanimalsmightalsoincreaseempathytowardnature. thegoalsofthisstudyweretoincreasescientificknowledgeandfosterempathytowardanimalsinagroupofbehaviorallyat-riskstudentsbyengagingstudentsinahands-onresearchproject.Wepredictedthatasaresultofpersonalinvolvementinbox-turtleresearch,theexperimentalgroupofstudentswouldshowincreasedknowledgeandempathywhencomparedtoacontrolgroupthatreceivedneitherclassroomlessonsnoranopportunitytoparticipateinfieldresearch.

study systemtheJuvenileJusticeAlternativeeducationprogram(JJAep)isanalterna-tiveeducationprogramforstudentswhohavebeenexpelledfromanyoftheeightschooldistrictsinlubbockCounty,texas.theprogramfocusesondisciplineandattemptstobringstudents’knowledgeuptotheircur-rentgradelevel.moreinformationcanbefoundatwww.lubbockisd.org/JJAep/.A0.6-hanatureareaisadjacenttotheschoolandisbisectedbyanaturewalk. theornateboxturtle(Terrapene ornata ornata)rangesfromsouth-erntexastosouthDakotaandfromColoradotoIndiana.ItisthestatereptileofKansas(Dodd2001)andisoftenwell-likedwhereveritisfoundnearpeople.likemostmembersofthegenus,theornateboxturtlehasdeclinedthroughoutmuchofitsformerrange.however,urbanlubbockholdslargepopulationsthathavereceivedsomeattentioninrecentyears(sosa2009).

hAnDs-onsCIentIFICtRAInInG

hatchlingornateboxturtle(Terrapene ornata ornata).

110 IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010

MethodsWeworkedwith students enrolled in the JJAep fromApril 2008 tooctober2009.oneofus,theactingprincipalofJJAep(oR),selectedmalestudentsbetweentheagesof13and17whometminimumrequirements(goodattendanceandbehavior)toparticipateinthehands-onexperiment.beforetraining,theselectedstudentsweregivenashortquestionnairewithquestionsonbox-turtlebiology,naturalhistory,andattitudestowardsani-mals.Uponcompletionofthequestionnaire,oneofus(JAs)metwiththestudentstwiceweeklyfor45minuteseachtime.Duringtheinitialthreeweeks,welecturedthestudentsonbox-turtlebiologyandnaturalhistorywiththeassistanceofbooksandarticles.Wealsodemonstratedthepropertechniquesemployedbyscientistsinthestudyofboxturtles,suchasradiotelemetryandthreadtrailing,totheexperimentalgroupofstu-dents.beginningonthefourthweek,wetranslocatedhatchlingornateboxturtles,whichhadbeengiventothesouthplainsWildlifeRehabilitationCenter,tothenatureareaadjacenttotheJJAepcampus.twohatchlingsin2008andthreehatchlingsin2009weretranslocatedtothenaturearea.theexperimentalgroupofstudentsaidedintheattachmentofradiotrans-mittersandmeasurementsofeachturtlepriortotranslocation. Werelocatedeachhatchlingtwotofivetimesaweekfortheremain-deroftheactiveseason.thestudentsassistedwithrelocationsbiweeklyduringtheschoolyear.Foreachrelocation,werecordeddate,time,Gpscoordinates,weatherdata,habitat,andactivityoftheturtles.beforestu-dentsweredismissedforsummerbreak(approximatelysixweeksfollowingadministrationoftheinitialquestionnaire),eachstudentcompletedthequestionnaireagain.Acontrolgroupofstudentswasgiventhequestion-nairetwiceduring2009,withasix-weekinterval,butreceivednotrain-ing.thequestionnaireconsistedoffactualquestions,aswellassomeabout

individualattitudes.Forquestions1–11,whichfocusedonstudents’per-ceptionsoftheirknowledge(e.g.,“howmuchdoyouknowaboutboxturtles?”)answersrangedfrom1(“nothing”)to5(“everything”).Questions13–15offeredarangeofanswers(e.g.,forthequestion“thebestplaceforaboxturtleis?”answerswere“innature”;“inazoo”;“inacageasapet”;“thereisnogoodplace”;and“noneoftheabove”). Weusedananalysisofvariancetotestforchangesinknowledgebetweentheanswersonpre-andpost-questionnairesforthetwogroups(experimentalandcontrol).significantdifferencesbetweenpre-andpost-surveyswerethenexploredfurtherwithanindependentsamplest-testforeachgroup.Wetestedfordifferencesbetweenqualitativepre-andpost-sur-veyquestions(13–15)withachi-squareanalysis.Resultswereconsideredsignificantatα=0.05.

ResultsFivestudentsin2008andfivein2009beganintheexperimentalgroup.AttherecommendationoftheJJAepprincipal,however,twostudentsdidnotcompletetheentiretrainingprogramin2009.Inaddition,12studentsin2009werepartofthecontrolgroupandreceivednotraining.Aftertrain-ing,allstudentsintheexperimentalgrouphadseenaturtle,whereasnodif-ferenceinbox-turtlesightingsexistedbetweenpre-andpost-questionnairesforthecontrolgroup.Wefoundasignificantdifferencebetweencontrolandexperimentalgroupsforquestions1–9andquestion11(p<0.05),allofthembox-turtlespecific,butnotquestion10,whichinvolvedfactorsresponsibleforenvironmentaldegradation.Withintheexperimentalgroup,wefoundasignificantimprovementbetweenpre-andpost-responsesonquestions1–11(p<0.05inallcases).Incontrast,wedetectednoimprove-mentforthecontrolgroupforquestions1–11(p>0.05inallcases).no

sosAetAl.

JJAepstudentsandahatchlingornateboxturtles.

JJAepnaturetrailmarkerandrepresentativehabitat.

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statisticallysignificantdifferencesexistedbetweenthepre-andpost-answersforquestions13–15(p>0.05)foreithergroup.

Discussionourresultsindicatethatstudentsintheexperimentalgroupimprovedtheirknowledgeonbox-turtlebiologyandnaturalhistory.Questions13–15,however,wereanattempttodetermineiftrainingimpactedstudents’empa-thytowardboxturtles.theanswerchoicesmighthavebeenat theextremes,eitherapositiveornegativeresponsewithnoneutralchoices.Ifstudentshadamorediversearrayofanswerchoicesandweworkedwithalargersampleofstudents,webelievetheresolutionofwhetherstudents’empathychangedwouldimprovebecauseempathyisexpectedtoincreasewhendirectcontactwithanimalsisprovided(thompsonandGullone2003). At-riskstudentsarecapableofindependentlycollectingscientificdatawhengiventraining,despitethefactthatsomeprocedureswerecomplex(radio-telemetry)andothersrequiredmeticulousattentiontodetail(datarecording).theexperimentalgroupofstudentsisnowtrainedinadvancedscientificproceduresthatmosthighschoolstudentsdonothavetheoppor-tunitytolearn. Inconclusion,ourtrainingprogramwaseffectiveinteachingat-riskstudentsbiologyandnaturalhistoryofboxturtlesaswellasscientificskills.

Futurestudiesshouldfocusonframingquestionsonempathyinordertoimproveresolution.studiesalsoshouldexaminewhetherstudentswhopar-ticipateintraininghaveincreasedconfidencecomparedtostudentswhodonotreceivespecializedtraining.becauseJJAepisaschoolforstudentsexpelledfromtraditionalschools,theextraattentioncouldprovidestudentswithmoreconfidenceandincreasetheirmarketabilitylaterinlife.Ataminimum,studentswereexposedtonovelideasandapotentialcareerpath.

AcknowledgementsWe thank C. Klein for assistance in planning this experiment. theequipmentused in thisexperimentwaspartially fundedbythetexasherpetologicalsocietyandthetexasparksandWildlifeDepartment.thisexperimentwasconductedundertexastechUniversityIACUCprotocol#108021-05andIRbprotocol#501915.thisismanuscriptt-9-1187oftheCollegeofAgriculturalsciencesandnaturalResourcemanagement,texastechUniversity.

literature CitedDodd,C.K.,Jr.2001.North American Box Turtles: A Natural History.Universityof

oklahomapress,norman.

hergovich,A.,b.monshi,G.semmler,andV.Zieglmayer.2002.theeffectsofthepresenceofadogintheclassroom.Anthrozoös15:37–50.

myers,o.e.,Jr.andC.D.saunders.2002.Animalsaslinkstowarddevelopingcaringrelationshipswiththenaturalworld,pp.153–178.Inp.h.Kahnands.R.Kellert (eds.),Children and Nature: Psychological, Sociocultural, and Evolutionary Investigations.mItpress,Cambridge,massachusetts.

paul,e.s.2000.empathywithanimalsandhumans:Aretheylinked?Anthrozoös13:194–202.

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

taylor,n.andt.D.signal.2005.empathyandattitudestoanimals.Anthrozoös18:18–27.

thompson,K.l.ande.Gullone.2003.promotionofempathyandprosocial-behaviourinchildrenthroughhumaneeducation.Australian Psychologist38:175–182.

hAnDs-onsCIentIFICtRAInInG

JJAepstudentsusingradio-telemetrytorelocatetranslocatedturtles.

Asked“Whatisyourknowledgeonhowbigaboxturtlecangrow?”experimental-groupstudentsshowedsignificantlygreaterimprovementafterworkingwithturtlesthandidcontrol-groupstudents.similarpatternswereseenonallnaturalhistoryquestions(questions1–11onthepre-andpost-surveys).

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managementofhabitatsfortargetspecies,suchastheYellowAnaconda(Eunectes notaeus)inArgentina,canbebeneficialtomanyotherspecies,suchastheseneotropicalCormorants(Phalacrocorax brasilianus)atbanyadolaestrella.

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c o m m e n T a R Y

Is Wildlife management business or conservation — a Question of Ideology

JesúsA.Rivas

Departmentofevolution,ecology,andorganismalbiology,theohiostateUniversity,Columbus,ohio

photographsbytheauthor(exceptwherenoted).

Macroeconomics and Conservation Approaches

Ipublishedanarticlein2007aboutanacondaconservationandhowitcanbeaffectedbymacroeconomics(Rivas2007a).Iidentifiedpovertyas

theultimatethreattoconservationinlatinAmericaandhowconservationeffortswereboundtomeetwithlittlesuccessaslongaspovertyremainstheruleinruralareas.Ialsoidentifiedneoliberalpolicies1asoneofthemaincausesofpovertyandhighlightedhowwellintendedconservationefforts,basedonneoliberalmeasures,failtosolvethepovertyproblemsofruralregions—andthusalsofailintheirconservationgoals.Instead,theyworkasa“painkiller,”creatingtheillusionofasolution,providingatbesttempo-raryrelief,butinfactdistractingfromseekingrealsolutions.

What is true Conservation?AtthecoreofanydisagreementregardingconservationprogramsinlatinAmericaisthenotion,widelyheldamongmanyconservationbiologists,thatanyplanforwildlifemanagement,includingecotourism,isbydefini-tionaconservationplan.thisnotionhasbeenpromotedforthelastfewdecadesinordertocapitalizeonpeople’sincreasingenvironmentalaware-ness(e.g.,mansfied2009).however,considerableevidencesuggeststhatwildlifemanagementisnotde facto conservation.Considerabird-watchingoperationlocatedinanareainhabitedbyaveryshyandrarespecies.birdwatchersflocktothesiteduringthenestingseasontoseethisrarespe-cies,whichcanproduceaneconomicsurgeinthelocaleconomy.Althoughsuchanexamplemightbeconsideredaneffectiveconservationplan,ifthisrarebirdissoshythatthesteadyparadeoftouristscompromisesnestingsuccess,thispopulationcouldliterallybe“watched into extinction.”Atrueconservationprogrammusthaveconservationasitsprimarygoalandnotjustasabyproduct.Ifonlyabyproduct,thesystemcaneasilystrayintoaregularbusinessregulatedsolelybythebottomline—andonethatmightnotevenbesustainable. Wildlifeharvestingprogramsfall intooneofthreecategories:(1)businessesthatexploitanenvironmentalcommodityuntilitisdepleted.(2)businessesthatuseanenvironmentalresourceinasustainablemannerbutwithoutprovidingenougheconomicincentivestothestewardsoftheland.(3)programsthatusearesourcesustainablybutalsoprovidesubstan-tiveeconomicincentivesforlocalcitizenswhothenhavegoodreasonstoprotecttheenvironmentfromotherusesthatmightnotbesustainable.Whenthebulkoftheeconomicincentivebenefitsthelocalcommunities,theywillhavebothreasonsandresourcestopreventexternalenterprisesfromthreateningtheenvironment.Iwouldarguethatthefirstexampleisnotconservationatallandthatonlythethirdistrueconservation.thesecondexamplecan—andshould—takecreditforbeingsustainable,butjustbecauseitdoesnotdestroytheenvironmentisnotenoughtoconstrue

itasaconservationprogram.As a matter of ideology, the goal of a conservation program must be conservation.economicgaincanbeabyproductorameanstodoconservationbutitmustnotbethegoal.Also,themainbeneficiariesofatrueconservationprogrammustbethelocalcommunities.theyaretightlylinkedtothelandandwillmorelikelytrytoprotectanecosystemthatsupportsthem—iftheyhavetheresources.externalbusinessescaneasilymovetheiroperationelsewhereandarenottrulycommittedtothemaintenanceofthesystem.

Management of Anacondas in Formosa, ArgentinaInmy2007article(Rivas2007a),Ineverintendedtoprovideacompre-hensivereviewoftheArgentineanYellowAnacondamanagementprogramandIdonotintendtodosonow.myconcernthenandnowisthatman-agementprogramsthatallocatemostoftheprofittoaneconomiceliteprovideonlysuperficialrelieftotheproblemsofthelocalpeople,donotprotectthesystemagainstexternalinfluences,anddonotconstitutetrueconservation.Infact,theyhavethepotentialfordistractingusfromseekingrealsolutions. micucciandWaller(2007),andWallerandmicucci(2008)high-lightedanumberofpositiveelementsintheFormosaprogram.Inaddi-tion,theprogramhasdoubtlesslyincreasedtheeconomicstatusofthelocalpopulation.Frominterviewswithlocalpeople,Ilearnedthattheanacondaharvestcouldincreasetheiryearlyincomebyasmuchas50%.Ialsolearnedfromlawenforcementofficialsthattherateofcattlerobberyandcommoncrimeshaddroppedtohistoriclevelssincetheprogrambegan,whichtheyattributedtothelocalpeoplehavinglegalmeansofearninganincome.Whileallthesearedesirabletraitsinamanagementprogram,theydonot

CommentARY

1Inessence,neoliberalpoliciesseekstotransfermuchofthecontroloftheecon-omyfrompublictotheprivatesectorunderthebeliefthatitwillproduceamoreefficientgovernmentandimprovetheeconomichealthofthenation.

Formorethantwodecades,theVenezuelanspectacledCaiman(Caiman crocodilus)programgeneratedacontinuousprofitandwasoftencitedasanexampleofsustain-ablemanagementinafree-marketeconomy.however,thesystemcollapsedasaconsequenceofover-hunting,andtannersmovedtheiroperationstoothersites.

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differfromanyotherbusinessmovingintoanareaandtheymightfailtoprotecttheecosystemagainstnon-sustainableuses—becausetheincentiveofferedbytheanacondaprogram,asdescribedinmicuccietal.(2006),isnotenoughtoempowerthelocalpeople,nordoesitprovidethemwiththemeanstoopposeacorporatetakeoverinsearchofgreaterprofits. Wildlifemanagementprogramsaroundtheworldarenotaskedtomeetthesehighstandardstoqualifyasconservation.Iwouldalsoarguethatthisisthereasonconservationprogramsmoreoftenthannotshowpoorresults.thisandmyearlier2007papersareintendedtoraiseaware-nessabouteconomicsandpoliticsamongconservationbiologists,todesignmanagementprogramsthatnotonlyuseresourcesinasustainablemannerwithconservationasabyproduct,buttodesignthemwithconservationastheprincipalgoalandtoincludeinthemmeansofprovidinglocalcommu-nitieswiththeresourcestowithstandpressuresfromexternalsourcesthatpromotenon-sustainableusesinfavorofshort-termprofits2. Weshouldnotusetheterm“conservation”forprogramswithgoalsthatarenotprimarilyconservation-oriented.Forexample,catchingfishtosupplyhigh-endrestaurantsiscalledfishing,notfishconservation.Fishingoperationsaroundtheworldarefirstandforemostcommercialbusinesses—andfishingoperationshaveonmanyoccasionsover-fishedtheirstocks(e.g.,hutchingsandmyers1994,larkin1977,myersetal.1997).Furthermore,evensustainablefishingoperationsdonottrytodisguisetheirbusinessasconservationprograms.theanacondamanagementprograminFormosamaywellbealegitimate,sustainablebusinessthathelpsthelocaleconomy(likeanybusiness)andrelievespressureonthenaturalenvironmentbyprovidingjobs(asbusinessesoftendo)—but,ifconservationismerelyabyproduct,suchaprogramshouldnotbepresentedasconservation.

Globalization or No Globalization? that is the QuestionConservationeffortsbasedonglobalizationandthefreemarketareriskybecausetheyarenottime-testedmodelsandfallwithinanarrowcontextofeconomicprinciples.Wecannottrustourpreciousdiversitytosuchuntestedeconomicmodels.Free-marketeconomieshave largely failedintheonetasktheypurportedlyaredesignedtodowell:productionofwealth.theUnitedstatesisoneofveryfewcountries(basicallytheG83)thathavebenefitedfromafreemarketsystem—butthatisnotthecaseforthemajorityofthecountriesthathavetriedit.Furthermore,thecountriesthathavesucceededunderfree-marketeconomiesarecountriesthathavedestroyedmostoftheirpristinenaturalhabitats,asafreemarketreliesonconstanteconomicgrowth.Usingglobalizationandfree-marketmeasuresforconservationpoliciesisaresponsetoideologicalagendas,anditisnotdata-drivenorsupportedbyfacts(e.g.,mansfield2009). Idonotintendtoturnthiscommentaryintoadebateoneconomicsorpolitics,butwhenweapplyaconservationstrategythatistightlylinkedtoaneconomicideologywearesupportingthatideology,whetherwereal-izeitornot.Insistingonfree-marketmeasuresforconservationdespitetheirrepeatedfailurestoprotectbiodiversityisnotonlyineffectivebutshowsadherence—consciousornot—toideologicalpositionsthatareintrinsicallyatoddswithconservationprinciples.

tylenol ConservationAsIarguedinmy2007articles,temporarymeasurescanandshouldbedevelopedtoaddressandrelieveshort-termproblems.todifferentiatethemfromrealsolutions,Ilabeledthem“tylenolConservation,”astheyworklikeapainkiller,amelioratingsymptomsofadiseasetheyarenotintendedtocure.Amanagementprogramthatrelieveslocalpovertywhilewesearchforrealsolutionsisawelcometoolaspartofaconservationprogram,butitisitnotconservationbyitself—anditshouldnotreplacethesearchforarealsolutionanymorethanapainkillershouldreplacethesearchforarealcure. manyoftheconservationsolutionsweseekintoday’sworldaredes-tinedtofailbecausetheyrelyonthesameneoliberalframeworkresponsibleforthepovertythatislargelyresponsibleforthefailureofconservationprograms,andtheyprovideonlytemporaryandsuperficialrelief.thisis

2Imaginethatacorporationwantedtodrainlargeportionsoftheswampfromwhichanacondasarebeingharvestedto,forexample,plantoilpalmsforthepro-ductionofagro-fuels.thisoperationwilldestroythehabitat,butwouldalsoofferpermanentemploymentwithcomparableorsuperiorincometowhatthelocalsmakefromwildlifeharvesting.Willthelocalsbewillingtoopposethisoperationtoprotectthehabitat?Willtheyhavetheresourcestoopposethecorporation?Icontendthatitisonlyconservationiftheanswertothesequestionsisyes.

IstartedtostudyGreenAnacondas(Eunectes murinus)inVenezuelain1992inordertoexplorethepossibilitiesforsustainableuse.Duetothecollapseofthecai-manprograminthemid-1990s,theVenezuelangovernmenthaltedotherharvest-ingprograms.Consequently,noattempttoharvestanacondasevermaterializedinVenezuela.Conservationbiologistsoftenbelievethattheirapproachtoconservationispure conservation,strictlyscientific,orsomehowdevoidofpoliticsorideology.however,managementprogramsbasedonafree-marketeconomyrelyonconstantgrowth,whichisintrinsicallyatoddswithconservationprinciples.scientistswhofailtorealizethisareatriskofbecomingunwittingtoolsofeconomicagendasthattheydonotunderstandorwithwhichtheymightnotevenagree.

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large, non-aquatic animals have been unable to flourish in most Capybara(Hydrochoerus hydrochaeris)habitats(hoogesteijnetal.1997).Infact,Capybaraare,forthemostpart,thelonelargeherbivoreinmostoftheirnaturalhabitats.Assuch,thenormallargepreypredatorhasnotevolved,andcapybaracanbefarmedinanalmostcompletelynaturalsetting.Consequently,manyconservationistshavestronglypushedforgovernmentallysubsidizedCapybarafarming.

3A forumfor theworld’smajor industrializeddemocracies (Canada,France,Germany,Italy,Japan,Russia,UnitedKingdom,Unitedstates)todiscussissuesofmutualorglobalconcern.

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whyIseektoredefinewhatwedoinconservationbypromotingagreaterawarenessofthepoliticalandeconomicframeworkinwhichwefunction.notdoingsocanrenderusunwittingtoolsofeconomicandpoliticalide-ologiesthatcompromisethesuccessofconservationefforts.

Referenceshoogesteijn,R.andC.A.Chapman.1997.largeranchesasconservationtoolsin

theVenezuelanllanos.Oryx31:274–284.

hutchings,J.A.andR.A.myers.1994.Whatcanbelearnedfromthecollapseofarenewableresource?AtlanticCod,Gadus morhua,ofnewfoundlandandlabrador.Canadian Journal of Fisheries and Aquatic Sciences51:2126–2146.

larkin,p.A.1977.An epitaph for the conceptofmaximum sustainedyield.Transactions of the American Fisheries Society106:1–11.

micucci, p.A. and t. Waller. 2007. the management of Yellow Anacondas(Eunectes notaeus)inArgentina:Fromhistoricalmisusetoresourceapprecia-tion.Iguana14:161–172.

micucci,p.A.,t.Waller,ande.Alvarenga.2006.programaCuriyú,pp.77–92.In:m.l.bolkovicandD.Ramadori(eds.),Manejo de Fauna Silvestre en la Argentina. Programas de Uso Sustentable.buenosAires,Argentina.

mansfield,b.2008.Globalenvironmentalpolitics,pp.235–346.In:K.Cox,m.lowandJ.Robinson(eds.),Handbook of Political Geography.sage,london.

mansfield,b.2009.sustainability,pp.37–49.In:n.Castree,D.Demeritt,D.liverman,andb.Rhoads(eds.),A Companion to Environmental Geography.blackwellpublishingltd.,oxford.

myers,R.A.,J.A.hutchings,andn.J.barrowman.1997.Whydofishstockscol-lapse?theexampleofcodinAtlanticCanada.Ecological Applications7:91–106.

Rivas,J.A.2007a.Conservationofanacondas:howtylenolconservationandmacro-economicsthreatenthesurvivaloftheworld’slargestsnake.Iguana14:74–85.

Rivas,J.A.2007b.Whatiswrongwithpainkillers,npR,theDemocraticparty,andconservationbiologists.The Axis of Logic(http://axisoflogic.com/artman/publish/article_25333.shtml).

Waller,t.andp.A.micucci.2008.Anacondaconservation:AreplytoRivas.Iguana15:51–53.

notonetoquestionitsluck,thisGreategret(Ardea alba)readilyexploitsthehuman-mediatedintroductionofGreenIguanas(Iguana iguana)ontoGrandCayman.

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I n v a s I v e s P e c I e s

Island Invaders: Introduced amphibians and Reptiles in the Turks and caicos Islands

R.GrahamReynoldsandmatthewl.niemiller

Departmentofecologyandevolutionarybiology,Universityoftennessee,Knoxville,tennessee37996,UsA(rgraham@utk.edu,mniemill@utk.edu)

theterm“invasive”usuallyreferstonon-nativespeciesthatarehav-ingatleastsomenegativeimpactonlocalfloralorfaunalcommuni-

ties.Invasivespeciesareamongthetopthreecausesofglobalbiodiversitydecline(lockwoodetal.2007),andunderstandingtheirdistributionsandavenuesofintroductionisimportantforregionalconservationandmanage-ment.moreover,anunderstandingofthebiologyandnaturalhistoryofinvadersmighthelptosuggestmethodsfortheircontrolandalsopredicteffectsonnativewildlife. Recently,much attentionhasbeenpaid to invasive reptiles andamphibians(Kraus2009).Reptilesandamphibiansmaybeespeciallygoodcolonizers,asevidencedbythenearlycircumtropicaldistributionofcertainspecies,suchasthebrahminyblindsnake(Ramphotyphlops braminus)andtheCanetoad(Rhinella marina). Islandsseemtobeparticularlyvulnerabletoinvasivespecies,asthesespeciesoftenarefreedfromthepressuresofnaturalenemies(predatorsandparasites)andcompetition(WhittakerandFernández-palacios2007).many tropical and subtropical islands contain unique herpetofaunalassemblagesthatarevulnerabletodisruptionbytheintroductionofnon-nativepredatorsandcompetitors.theWestIndiesisconsideredoneoftheworld’smostimportantbiodiversityhotspotsandtheregion’snativereptilesandamphibiansareofparticularconservationconcern(myersetal.2000,smithetal.2005).Centuriesofhabitatmodificationandtheintro-ductionofdamagingmammalianpredatorssuchasferalcats(Felis catus),Indianmongoose(Urva auropunctata),andblackRats(Rattus rattus)havenegativelyaffectedmanyreptilianandamphibianpopulations(e.g.,Iverson1978,Corke1992,smithetal.2005,tolsonandhenderson2006).Assuch,documentingandreportingthespreadandimpactofintroducedher-petofaunaremainsanimportanttask.

Fig. 1.theturksandCaicosIslandsarelocatedatthesouthernterminusofthebahamianArchipelago~130kmfromhispaniola.majorislandsandislandsmen-tionedinthetextarelabeled.

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Fig. 2.nativereptilesoftheturksandCaicos:(A)turksandCaicosRockIguana(Cyclura carinata),(b)turksandCaicosCurly-tailedlizard(Leiocephalus psam-modromus),(C)Caicosblindsnake(Typhlops platycephalus),(D)CaicosDwarfboa(Tropidophis greenwayi),(e)Antilleanskink(Mabuyasp.),(F)turksDwarfGecko(Sphaerodactylus underwoodi;female,left;male,right),(G)CaicosDwarfGecko(Sphaerodactylus caicosensis;male,bottom;female,top),(h)Caicos(hecht’s)barkingGecko(Aristelliger hechti),(I)turksIslandboa(Epicrates chrysogaster chryso-gaster),(J)southernbahamasAnole(Anolis scriptus scriptus).

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WehavespentthelastfouryearsworkingintheturksandCaicosIslands(Fig.1)andhavemadeanefforttodocumentthepresenceanddis-tributionofnon-nativespeciesofthisarchipelago.theturksandCaicosIslands,locatedatthesouthernterminusofthebahamianArchipelago,containtenspeciesofnativereptiles,eightofwhichareendemicatthespeciesorsubspecies level (Fig.2).theremainingtwospecies (Anolis scriptusandMabuyasp.)arewidelydistributedinthesouthernbahamasandthroughouttheWestIndies.noamphibianspeciesisnativetotheturksandCaicos.Asofoctober2009,sevennon-nativereptilianandtwoamphibianspecieshavebeendiscoveredintheturksandCaicos,nearlydoublingthenumberofreptilesandaddingamphibiansinaregionwherepreviouslytheyhadnotbeendocumented.tourismandlargeinternationalinvestmentshaveresultedinexplosivedevelopmentoftheseislandsandasurgeinthenumberofimmigrantsfillingjobsintheconstructionandserviceindustries.Combinedwiththerapidincreaseinimports,thelistofinvadersmightcontinuetogrow.belowwediscussthedistributionandabundanceoftheknowninvasivereptilesandamphibiansintheturksandCaicos,andencourageotherresearcherstodocumentsimilarinvasionsinotherregionsthatsupportauniquenativeherpetofauna.

Cuban treefrog, Osteopilus septentrionalis. Established.Cubantreefrogs(Fig.3)arewidely introducedintheCaribbeanandsoutheasternUnitedstates,withpopulations onpuertoRico, in theVirginIslands,upperlesserAntilles,Florida,andelsewhere(hendersonandpowell2009).thesefrogsappearabletotoleratexericconditionsthatwouldpreventmostotheramphibiancolonistsfrombecomingestablished.Aslongastheyhaveaccesstoephemeralorpermanentsourcesoffresh-water,thesefrogscanbreedprolificallyandbecomeabundant.Cuban

treefrogsareofparticularconcernbecauseoftheirvoraciousappetites,highdensities,andnoxiousskinsecretions.AlthoughthisspeciesisnativetothelittleandGreatbahamabanks,itisgenerallyconsideredarecentarrivaltotheturksandCaicos,whereitisfirmlyestablishedontheislandsofprovidenciales,Grandturk,northCaicos,andmiddleCaicos,andwilllikelybefoundonseveralotherislandswiththeexpansionofirrigationfordevelopment.onnorthandmiddleCaicos,thesefrogsreachexcep-tionaldensities,oftencoveringtheroadafteraheavyautumnrain.endemicCaicosDwarfboas(Tropidophis greenwayi)haveevenshiftedtheirbehaviortofeedonthesefrogs,occupyingtheinteriorwallsofoldstonewaterwellstocapturemetamorphsastheyclimbupthesidesfromthewaterbelow(RGR,C.Deal,andn.manco,unpubl.data).Colorphotovouchers,AustinpeaystateUniversity(ApsU)19024,19027.

greenhouse Frog, Eleutherodactylus planirostris. Established.GreenhouseFrogs(Fig.4)aresmallterrestrialfrogsthatstowawayinsoilandplants,andhavebeenintroducedtoJamaica,Florida,andtheturksandCaicosIslandsfromtheirnativerangeinCuba,theCaymans,andthebahamas(hendersonandpowell2009).thisspeciesisadirect-developer,meaningthatthelarvalstageiscompletedintheegg.eggsarelaidinmoistleaflitter,andhatchlingsemergeasminiatureadults.GreenhouseFrogsoccurintheturksandCaicosontheislandsofprovidenciales,Grandturk,northCaicos,andmiddleCaicos,andwilllikelybediscoveredonotherislands.Colorphotovoucher,ApsU19023.

Red-Eared slider, Trachemys scripta elegans. Not established.Red-earedsliders(Fig.5)arepopularpetswithlonglifespans,andoftenarereleasedwhenownersgrowtiredofthem.nativetotheeasternandmid-westernUnitedstates,thesefreshwateraquaticturtleshavebeenintroducedtomanyislandsintheWestIndies,frompuertoRicoandhispaniolatoGuadeloupeandmartinique(hendersonandpowell2009).theyrequireaconsistentsourceoffreshwaterand,althoughthistypeofhabitatisrareintheturksandCaicos,severalpondsbuiltfortheleewardgolfcourseonprovidencialeslikelyprovidethemainrefugeforthisspecies.onlyafewadultRed-earedslidershavebeencapturedandremoved,althoughafewmoreprobablyoccurinthesemanmadeponds.nohatchlingsorjuvenileshavebeenobserved;hencewedonotconsiderthisspeciestobereproduc-ing.Colorphotovoucher,ApsU19021.

Antillean slider, Trachemys stejnegeri malonei. Established?AsingleAntilleansliderwascollectedin1975frompineCayontheCaicosbank(W.Auffenberg,UF49423),althoughseveralotherindividualswerereportedin1997fromafreshwaterpondonthesameisland(seidel1986,Fig. 3.Cubantreefrog(Osteopilus septentrionalis)fromnorthCaicos.

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1988;leeandRoss2001).thisspeciesisbelievedtobeahumanintroduc-tiontotheCaicosCays(seidel1988,1996)fromGreatInagua.Weareunawareofanyspecimensreportedsince1997,althoughpersistenceofthispopulationfrom1973to1997wouldindicatethatreproductionmightbeoccurring,unlessthespeciesisbeingrecurrentlyintroduced,asituationthatseemsunlikelyonthisprivatelyownedresortisland.

Wood slave, Hemidactylus mabouia. Established.Woodslaves(Fig.6)arerapidlyexpandingtheirrangeintheWesternhemisphere,andarefirmlyestablishedintheturksandCaicosIslands.theselizardsareexceptionallygoodcolonizersand,ashumancommensals,areaffordedfrequentopportunitiestostowawayincargo,whichisprob-ablyhowtheyarrivedintheturksandCaicos.theyarewidelydistributedintheturksandCaicosandarecurrentlyknownfromsixlargeislands:providenciales,northandmiddleCaicos,southCaicos,Grandturk,andsaltCay(Reynoldsandniemiller2009),althoughtheylikelyoccuronmanymore.WehavediscoverednestsonsaltCay(Fig.7)andhaverecordedjuvenilesandhatchlingsonallsixislands.thesegeckosprobablydonotrepresentagreatthreattomostofthelocalherpetofauna,exceptontheislandsofnorthCaicos,FrenchCay,andbigAmbergris,wheretheendemichecht’s(orCaicos)barkingGecko(Aristelliger hechti)occurs.thesenativegeckosareecologicallysimilartotheintroducedWoodslave,

occupyingverticalsurfacesoftrees,rockwalls,andbuildingsandfeedingonsmallflyingorclimbinginsects.Colorphotovouchers,ApsU18047,18945,18946,and18949.

Mayaguana dwarf gecko, Sphaerodactylus mariguanae. Established?mayaguanaDwarfGeckosarenativetothebahamianislandsofmayaguanaandboobyCay,locatedabout65kmtothenorthwestoftheturksandCaicos.Althoughsmall(sVlto41mm;schwartzandhenderson1991),theyaremuchlargerthanthenativeS. caicosensis (sVlto32mm)and S. underwoodi(sVlto32mm).Dwarfgeckoslikelymovebetweenislandsquiteeasilybecauseoftheirsmallsize,highfecundity,andproclivityforinhabitingstacksofbuildingsupplies,suchascinderblocksandlumber,aswellaspottedplants,soil,andmulch.moleculardatasuggestthatnativeS. underwoodimovebetweentheislandsofGrandturkandsaltCayoncargo(ReynoldsandKoneczny,inreview). schwartz(1968)andschwartzandhenderson(1991)reportedS. mariguanaefromGrandturk.toourknowledge,thisspecieshasnotbeenseenthereinquitesometime,buttherecordstandsinhendersonandpowell(2009).thisrecordlikelydoesnotrepresentamisidentification,as40individualswerecollected(AlbertschwartzFieldseries[AsFs]10766)andcomparedtootherpopulationsandtoS. underwoodi(schwartz1968);however,whether theseS. mariguanae individuals represent ahumanintroductionoranextensionofthespecies’nativedistributionisunclear.Also,thatthisspecieshasapparentlynotbeenrecordedsinceschwartz(1968)andwasneverfoundonotherislandsontheturksbankisnotable.schwartz(1968)speculatedthattheseGrandturkS. mariguanaerepre-sentedanintroducedpopulation,andevidencecurrentlyavailablecertainlyweighsinfavorofthispostulate,henceweincludethisspecieshereasanintroducedspecies.Grandturkisaheavilydevelopedisland,yetendemicS. underwoodiappearstobefairlycommon,andthusS. mariguanaemayyetpersistthere.

green iguana, Iguana iguana. Established?Aspopularpets,GreenIguanas(Fig.8)havebecomeestablishedinmanyareasoutsideoftheirnativerangeinCentralandsouthAmerica,perhapsmostnotablyinsouthernFlorida,wheretheyoccurinveryhighdensi-tiesnearresidentialareas.IntheturksandCaicos,afewGreenIguanashavebeenfoundonGrandturkandprovidenciales.theylikelyrepresentreleasedorescapedpets.Reproductionhasnotbeendocumented,butthisseemsalikelypossibilityifseveralmatureadultsoccurinthesamearea.Colorphotovoucher,ApsU19019.Fig 6.Woodslave(Hemidactylus mabouia)fromsaltCay,turksandCaicosIslands.

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Cuban Knight Anole, Anolis equestris. Not established.likelyreleasedpets,fourCubanKnightAnoleshavebeencollectedandseveralmoresightedataGracebayresortonprovidencialesIsland.likeGreenIguanas,theselargearboreallizardsappeartothriveinappropriatehabitat,astheyalsoarequitecommonwheretheyhavebeenintroducedinsouthernFlorida(meshakaetal.2004).theyapparentlyarenotreproduc-ingintheturksandCaicos,althoughthatcannotyetberuledout,giventheabundanceoflushandirrigatedvegetationintheGracebayresortcom-plexes.Colorphotovouchers,ApsU19025,19026.

Brahminy Blind snake, Ramphotyphlops braminus. Established.essentiallyacircumtropicalspecies,thebrahminyblindsnake,whichisnativetotheIndiansubcontinent,hasproventobeanexceptionalcolo-nizerbecauseofitshabitofsequesteringitselfinpottedplantsanditspar-thenogeneticmodeofreproduction.thisspecieshasbeenfoundonbothprovidencialesandGrandturkandislikelytofinditswaytomanyoftheotherislandsinthearchipelago.Wecurrentlydonotknowwhetherbrahminyblindsnakescompetewiththeecologicallysimilarnativeblindsnakes(Typhlopssp.,sensus.b.hedges,in litt.),whicharemoreabun-dantonislandsotherthanGrandturkandprovidenciales.Colorphotovoucher,ApsU19022.

Eastern Corn snake, Pantherophis [Elaphe] guttatus. Not established.theCornsnake(Fig.9)isnativetothesoutheasternUnitedstatesandhasbeenintroducedelsewhereintheCaribbean,althoughitissuccessfullyestablishedonlyontheislandsofGrandCayman(schwartzandhenderson1991)andst.thomas(U.s.V.I.,hendersonandpowell2009).thesesnakesmighthavearrivedonislandsaseggslaidinthesoiloflargepot-tedtreesfromFlorida;however,escapedpetsalsoareapossiblemeansofconveyance.thusfar,threeadultshavebeenfoundonGrandturk,oneofwhichlaidaclutchofnon-viableeggsaftercapture(b.n.mancoandb.Riggs,pers.comm.).Colorphotovoucher,ApsU19020.

ConclusionstheturksandCaicoscontainauniqueassemblageofnativereptilespe-cies.threatsfromhabitatmodificationanddirectpersecution(i.e.,killingsnakes)aretakingatollonspeciessuchastheturksIslandboa(Epicrates chrysogaster)andtheturksandCaicosRockIguana(Cyclura carinata)(G.Gerber,unpubl.data;ReynoldsandGerber,inreview).threatsfrominva-sivemammalianspecies,suchascats,arewellestablished,andlocalreptilianpopulationshavesufferedtremendouslyfromtheirintroduction(Iverson1978,mitchelletal.2000).thedegreeofthreattolocalwildlifeposed

byintroducedreptilesandamphibiansislargelyunknownatthistime,however,thefirststepistodocumentsuccessfulinvasions.WeencouragefellowbiologistsandamateurherpetologistsintheWestIndiestotakenoteofintroducedspeciesandreportthemtolocalauthorities.

AcknowledgmentsWethankb.n.mancoandC.Dealforassistanceinthefield,andb.n.manco,b.Riggs,G.Gerber,m.hibbert,ande.salamancaforvaluableinformationregardingbothintroducedspeciesandreptilesintheturksandCaicos.thanksaswelltoA.F.scottforaccessioningphotovouch-erstotheAustinpeaystateUniversityCenterforFieldbiologymuseumin Clarksville, tennessee. We are grateful to W. Clerveaux and theDepartmentofenvironmentandCostalResources,turksandCaicosIslands,forscientificresearchpermits(#’s1-4RGReynolds),andtheturksandCaicosnationaltrustforlogisticalsupport.FinancialsupportfortheauthorswasprovidedbytheDepartmentofecologyandevolutionarybiology,Universityoftennessee,KnoxvillesummerResearchGrantprogram (RGR and mln), and the University of tennessee W.K.mcClurescholarshipforthestudyofWorldAffairs(RGR).thankstoR.W.henderson,R.powell,andG.perryforreviewsandhelpfulcom-mentsonthismanuscript.

ReferencesCorke,D.1992.thestatusandconservationneedsoftheterrestrialherpetofauna

oftheWindwardIslands(WestIndies).Biological Conservation62:47–58.

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

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

lee,D.s.andJ.p.Ross.2001.theCatIslandturtle,areptileofproblematicorigin,includingabibliographicreviewofthegenusTrachemysintheWestIndianregion,pp.36-47.In:C.Clark-simpsonandG.W.smith(eds.),Proceedings of the 8th Symposium on the Natural History of the Bahamas.GeraceResearchCenter,sansalvador,bahamas

lockwood, J.l., m.F. hoopes, and m.p. marchetti. 2007. Invasion Ecology.blackwellpublishing,malden,massachusetts.

meshaka,W.e.,Jr.,R.D.bartlett,andh.t.smith.2004.ColonizationsuccessbygreeniguanasinFlorida.Iguana11:155-161.

mitchell,n.,R.haeffner,V.Veer,m.Fulford-Gardner,W.Clerveaux,C.R.Veitch,andG.mitchell.2000.Cateradicationandtherestorationofendan-gered iguanas (Cyclura carinata)onlongCay,Caicosbank,turksandCaicosIslands,britishWestIndies,pp.206–212.In:C.R.Veitchandm.n.

Fig. 8.GreenIguana(Iguana iguana)capturednearlongbayonprovidencialesandbeingheldatthenationalenvironmentalCentre.

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Fig. 9.easternCornsnake(Pantherophis guttatus)capturedonGrandturk,turksandCaicosIslands,andbeingheldatthenationalenvironmentalCentre.

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Clout(eds.),Turning the Tide: The Eradication of Invasive Species.IUCnssC Invasivespecies specialistGroup, IUCn,Gland,switzerland andCambridge,UnitedKingdom.

myers,n.,R.A.mittermeier,C.G.mittermeier,G.A.deFonseca,andJ.Ke.2000.biodiversityhotspotsforconservationpriorities.Nature403:853–858.

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Reynolds,R.G.andm.l.niemiller.2009.Hemidactylus mabouia.(Woodslave).Distribution.Herpetological Review40:452.

schwartz,A.1968.thegeckoes(Sphaerodactylus)ofthesouthernbahamaislands.Annals of the Carnegie Museum39:227–271.

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

seidel,m.e.1988.Revisionof theWest Indianemydid turtles (testudines).American Museum Novitates(2918):1–41.

seidel,m.e.1996.CurrentstatusofbiogeographyoftheWestIndianturtlesinthegenusTrachemys (emydidae),pp.169–174.In:R.powellandR.W.henderson(eds.),Contributions to West Indian Herpetology: A Tribute to Albert Schwartz.societyforthestudyofAmphibiansandReptiles,Ithaca,newYork.

smith,m.l.,s.b.hedges,W.buck,A.hemphill,s.Incháustegui,m.A.Ivie,D.martina,m.maunder,andJ.F.ortega.2005.CaribbeanIslands,pp.112–118. In:R.A.mittermeier,p.R.Gill,m.hoffman, J.pilgrim,t.brooks,C.G.mittermeier,J.lamoreaux,andG.A.b.daFonseca(eds.),Hotspots Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial Ecoregions.CemeX,s.A.deC.V.,mexicoCity.

tolson,p.J.andR.W.henderson.2006.AnoverviewofsnakeconservationintheWestIndies.Applied Herpetology3:345–356.

Whittaker,R.J.andJ.m.Fernández-palacios.2007.Island Biogeography.2nded.oxfordUniversitypress,newYork.

cuban brown anoles (Anolis sagrei) in st. maarten

AxelFläschendrägerhalle,Germany(Axel.Flaeschendraeger@t-online.de)

photographsbytheauthor.

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

bahama,RumCay,andsansalvadorislandbanks),Cubaandassociatedcays,IsladelaJuventud,andlittleCayman,includingmostsatellitesandcayswithevenrudimentaryvegetation.thespeciesalsohasbecomeestablishedinJamaica,eithernaturallyorthroughhumanmediation.morerecentlyintroducedpopulationsareknowninGrandCayman,swanIsland,Grenada,st.Vincent,theGrenadines(Canouan),barbados,theAtlanticCoastofméxicoasfarasbelize,theIslasdelabahía(offhonduras),Aruba,hawaii(oahuandCoconutIsland,Kauai),taiwan,thesoutheast-ernUnitedstates,andCalifornia(orangeCounty;hendersonandpowell2009andreferencestherein).Atleastsomeintroducedpopulationsorigi-natedinthesoutheasternUnitedstates. Within a30-minuteperiodon2march2010, I observed threemaleandtwofemaleAnolis sagrei inandaroundthephilipsburgharborinst.maarten.st.maartenisamajorcommercialcenterintheeasternCaribbean;anumberofexoticspecieshavebeenrecordedthere,andsev-eral(e.g.,Cubantreefrogs,Osteopilus septentrionalis;GreenIguanas,Iguana iguana)havebecomeestablished(powelletal.2005). AlthoughIdidnotobserveanyjuveniles,theabundanceofobserva-tionsinsuchashortperiodissuggestiveofabreedingpopulation.IdidobservenativeAnguillabankAnoles(Anolis gingivinus)associatedwithornamentalvegetationinandaroundtheharbor.Isawnoevidenceofcompetitiveinteractions,buttheapparentlyrestrictedrangeofA. sagreiissuggestiveofarecentarrival.Whetherthenewlyestablishedpopulationremainslargelyrestrictedtoseverelyalteredhabitats,ashasbeendescribedonGrenada(Greeneetal.2002,Germanoetal.2003)andst.Vincent(hendersonandpowell2005,treglia2006,tregliaetal.2008)remainstobedetermined.theoriginofthepopulationonst.maartenisunknown.

Literature CitedGermano,J.m.,J.m.sander,R.W.henderson,andR.powell.2003.herpetofaunal

communitiesinGrenada:Acomparisonofalteredsites,withanannotatedchecklistofGrenadianamphibiansandreptiles.Caribbean Journal of Science39:68–76.

male(top)andfemaleCubanbrownAnoles(Anolis sagrei)fromthephilipsburghar-boronst.maarten.photographicvouchershavebeendepositedinthemilwaukeepublicmuseum:mpmp745(male)andmpmp746(female).theidentityoftheselizardswasconfirmedfromphotographsbyRobertW.henderson.

122 IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010 DACostA,GoDbeeR,AnDAUstIn

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.

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

powell,R.,R.W.henderson,andJ.s.parmerlee,Jr.2005.Reptiles and Amphibians of the Dutch Caribbean: St. Eustatius, Saba, and St. Maarten.st.eustatiusnationalparksFoundation,Gallowsbay,st.eustatius,netherlandsAntilles.

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.

Knight anoles (Anolis equestris) on Grand cayman Island

matDaCosta-Cottam,KristanD.Godbeer,andtimAustin

Departmentofenvironment,CaymanIslandsenvironmentalCentre,GrandCayman(mat.cottam@gov.ky)

KnightAnoles(Anolis equestris)arenativetoCuba,buthavebecomeestablishedinsouthernFlorida(KingandKrakauer1966)andhawaii

(mcKeown1996).secondaryintroductionsfromFlorida,primarilywithlandscapingvegetation,havebeendocumentedforseveralspecies(e.g.,Cubantreefrogs,Osteopilus septentrionalis,andCubanbrownAnoles,A. sagrei)onWestIndianislands(hendersonandpowell2009,andreferencestherein). on8June2007,wefoundaKnightAnoleonthegroundsoftheRitz-Carltonhotelonsevenmilebeach,GrandCaymanIsland(photo-graphicvoucher,milwaukeepublicmuseummpmp748,photographbyK.D.Godbeer,confirmedbyA.C.echternacht).subsequently,anaddi-tionalindividualwascollectedinthesavannaharea.bothpresumablyarrivedfromFloridawithdecorativeplants.todate,wehavenoevidencethatsuggeststhepresenceofabreedingpopulationinthewild.

Literature Citedhenderson,R.W.andR.powell.2009.Natural History of West Indian Reptiles and

Amphibians.UniversitypressofFlorida,Gainesville.

King,W.andt.Krakauer.1966.theexoticherpetofaunaofsoutheastFlorida.Quarterly Journal of the Florida Academy of Sciences29:144–154.

mcKeown,s.1996.A Field Guide to Reptiles and Amphibians in the Hawaiian Islands.Diamondheadpublishing,lososos,California.

AKnightAnole(Anolis equestris)fromFlorida,thepresumedoriginoftheanimalsfoundonGrandCayman. St

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b o o K R e v I e W

Timber Rattlesnakes in Vermont and New York: Biology, History, and the Fate of an Endangered Species.2007.JonFurman.Universitypressofnewengland,lebanon,newhampshire(www.upne.com/1-58465-656-5.html).207pp.,12colorillustrations,8halftones.paperback–Isbn:978-1-58465-656-2.$24.95

AnnouncedbytheCenterfornorthAmericanherpetology(CnAh)in2008,thisauthoritativeandwell-illustratedbookisanimportantacqui-sition for all ofuswhowork to advance the conservationoftimberRattlesnakes(Crotalus horridus)throughoutthewiderangeofthespecies.ReadersmaywellwonderwhyIfeelthatabookgroundedinthebiologyofthisimpressivespeciesintwonortheasternstatesissorelevantelsewhere.simplyput,conservation—especiallyofvenomoussnakes—isadelicatebalanceofscience,sociology,andpolitics.Furmanhascarefullyresearchedandthoroughlydocumentedtheinterplayofthesethreeelementsastheyrelatetothesundry(andincreasing)anthropogenicthreatsfacedbytimberRattlesnakepopulationsintwoeasternstatesthatonlyinrelativelyrecenttimeshaverecognizedthespeciesasendangeredandhavemovedtotrytoreversedeclines(orevenextirpations)resultingfromearlierflawedmanage-mentpractices. A talentedwriter, Furman explores the troublingdecline of thenortheasternpopulationscausedbybountyhuntingfromthe1890stotheearly1970s.hisfriendshipswithcontemporaryresearcherssuchasRandystechert,Williambrown,andothers,whoseindividualandcombinedresearchandconservationworkwithCrotalus horridushavebeencrucialtoreversinghistoricaltrendsofpersecutionofthespecies,addsmuchtothebook.Vignettesofthelegalbattleswagedonbehalfofthesesnakes,alongwith Furman’s observations on thepersonalities involved and their var-ied approaches to preventing furtherhumandamage todens andbirthingareasoffereffectivemodelsofappropri-ateconservationmeasures. AnunusualaspectofthebookisthetimeFurmaninvestedininterview-ing themostnotedCrotalus horridusbounty hunters of the area. Despitethegreattollthesepeopletookonthesnakes,fewweremotivatedbyanimos-ity toward them. Rather, they werepeopleekingouta living ina largelyagriculturalareaduringdifficulteco-nomictimes,andtheysawthepoorlyconceivedstatebountiesonrattlesnakesas anotherway toearn income.oneortwoofthehunterswentsofarastostudywildandcaptiveCrotalus horridusbehavior,ecology(althoughnotusingthatterm),andphysiology,thebettertolocatesnakesandplytheirtrade.eventheinterviewsinvolvingpreparationof

rattlesnakeoilasananti-inflammatory,atopicthatinitiallybroughtasmirktomyface,provedvaluableinunderstandinganotheraspectofwhythehunterspursuedtheirquarry.so,delvingintothepharmacologicalliteratureinDecember2009,Ifoundthatcertainsnakeoilsarehigherineicosapentaenoicacid(omega-3)thanmanyfishoils(Kunin1989,Graber2007)!

overall,thebookissolidlyanchoredinbiology, sociology, and history of science.AlthoughpartsofFurman’soralhistoryinter-viewswithold-timebountyhuntersoftenareunpleasantforherpetologiststoread,theyareimportantforunderstandingthesociologicalrealitiesthathistoricallyhavehamperedcon-servation of any venomous snake species.Ifherpetologistsaretosuccessfullyworktochangepublic attitudes about rattlesnakes,suchunderstandingiscrucial(pisaniandFitch1993).

conservation of venomous snakes is a Delicate balance of science, sociology, and Politics

Geo

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Amaledark-phaseCrotalus horridusfromWarrenCounty,newYork.

Literature CitedGraber,C.2007.snakeoilsalesmenwereonto

something.Scientific American,1november2007(availableonlineat:www.scientificam-erican.com/article.cfm?id=snake-oil-sales-men-knew-something).

Kunin,R.A.1989.snakeoil.Western Journal of Medicine151:208.

pisani,G.Randh.s.Fitch.1993.A surveyofoklahoma’s rattlesnake roundups.Kansas Herpetological Society Newsletter(92):7–15.

George R. PisaniKansasbiologicalsurvey

lawrence,Ks66047

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Copperheads in the Northeastern united states

SmItH Et Al.(2009.Herpetological Monographs23:45–73) studied the spatial and reproduc-tiveecologyofapopulationofCopperheads(Agkistrodon contortrix) in a basalt trap-rockecosystem in the central Connecticut RiverValley,aregionthatconstitutesthenortheast-ernextremeofthisspecies’geographicrange.Adultmales(n=20)andfemales(n=15)weresurgically implanted with radio-transmittersandtrackedevery48hduringtheactivesea-son(April–october)forthreeconsecutiveyears(2001–2003).Fromlateautumntoearlyspring(novemberthroughmarch),whensnakeswerehibernating and thus inactive, tracking wasreducedtoonceperweek.Activityrangesizeandmultiplemovementparametersdifferedsig-nificantlybysex.throughouttheactiveseason,maleshadgreateractivityrangesizesandshowedgreatermovementthanfemales.thistrendwaspronouncedduringthematingseason,whichwasrestrictedtolatesummerandearlyfall(lateJulythroughseptember).IncontrasttomostpopulationsofA. contortrixfrommoresouthernandwesternlocalities,theauthorsobservednosexualactivity(e.g.,courtship,coitus,andmale-malefighting)inthespring.Individualsofbothsexesshowedannualfidelityto:(i)Activityrangelocation,(ii)activityrangesize,(iii)movementdistances,(iv)particularfeaturesoftheiractiv-ityranges (e.g., refuges),and(v)hibernationsites.malesandfemalesshowednodifferenceinpreferredseasonalhabitats.Inbothsexes,shiftsinhabitatassociationsduringtheactiveseasonincludedmigrationsfromover-winteringsiteswithinbasalttraprockslidestohigher-elevationopendeciduousforestduringthesummerforag-ingandreproductiveseason.parturitioninthefieldwasrecordedineightinstancesandwasalwaysclosetooneofthetwohibernaculausedbytheindividualsinthestudyarea.theauthorsdedicatedthispapertohenrys.Fitch.

secretive snakes in a grassland Community

Fitch(1999)summarizedhisfiftyyearsofinten-sivefieldworkonthesnakefaunaoftheFitchnaturalhistoryReservation(FnhR)andadja-centstudyareas,commentingupontherela-tivescarcityoftwosmall,secretivespecies,thesmoothearthsnake(Virginia valeriae)andthebrownsnake(Storeria dekayi).locationofapopulationofV. valeriae(pisani2005)asacom-ponentofadiversesnakefaunathatincludesS. dekayionlandadjacenttoFitch’smainstudyareasprovidedanopportunitytoaccumulateconsiderablenewinformationonthesespeciesinasimilarsnakecommunity.PISAnI(2009.Journal of Kansas Herpetology32:20–36)con-ducted an intensivemark-recapture studyofthiscommunityfromseptember2006through

november2008.theongoingstudyincludedobservationsonfeeding,defense,homerange,morphology,andpopulationsizesalongwithrelevantmicrohabitatcharacteristics.Dataaccu-mulated indicate considerably greater use ofavailablegrasslandhabitatbytheseandsyntopicsmallsnakespeciesthanhadbeenrecognized.Anextensivereviewof literatureandunpub-lisheddissertationsprovidedinsightstoaspectsofthecommunityinterrelationsofsmallver-mivoroussnakesinnortheasternKansas.theseinterrelationsareconsideredin lightof land-managementandotheranthropogenicfactorsaffectingtheabundanceofearthworms.

Eavesdropping lizardsmadagascarspinytail Iguanas (Oplurus cuvi-eri cuvieri)usetheirearstoeavesdroponthemadagascarFlycatcher’s(Terpsiphone mutata)alarmcallstoprotectthemselvesfrompredators.Ryo Ito AnD AkIRA moRI (2009.Proceedings of the Royal Society B, Biological Sciences277:1275–1280)conductedastudyintheAmpijoroadrydeciduousforestanddiscoveredthatthelizardsuseauditorysignalsforpredatordetection.theiguanashavewell-developedears,butarenotvocalliketheflycatchers.eavesdroppingisthefirststepforthelizard;itthenmovesitsheadwhilekeepingitsbodystilltohelpidentifythepredatorandevaluatetherisk.thelizardsandtheflycatchersdonothavedirectinteractionsnordo they compete for resources,but theydosharepredatorssuchasraptorsandsnakes.evidence suggesting that Galápagos marineIguanas(Amblyrhynchus cristatus)alsoeavesdroponheterospecificalarmcallsledtheresearcherstobelievethatthismightbeawidespreadphe-nomenoniniguanidlizards.

Repeated Adaptations of garter snakes to deadly Prey

Wheredothegeneticvariantsunderlyingadap-tivechangecomefrom?Arecurrentlyadaptivealleles recruited by selection from standing

madagascar spinytail Iguanas (Oplurus cuvieri cuvieri)eavesdroponthealarmcallsofmadagascarFlycatcherstoalertthemselvestopotentialpredators.

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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

smoothearthsnakes(Virginia valeriae)andbrownsnakes(Storeria dekayi)exploitavailablegrasslandhabitats innortheasternKansastoamuchgreaterextentthanhadbeendocumented.

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Unlike populations of Copperheads (Agkistrodon contortrix)frommoresouthernandwesternlocalities(likethissnakefromKansas),apopulationfromthenortheasternextremeofthespecies’rangeexhibitednosexualactivityinthespring.

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pacificnewts,suchasthisCalifornianewt(Taricha torosa),containdeadlytetrodotoxinsthatarelethaltomanypotentialpredators.however,atleastthreespe-ciesofGartersnakesinthegenusThamnophishaveindependentlyevolvedaresistancetothosetoxinsandreadilypreyonnewts.

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IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010 125

loggerhead turtles Nesting in Pakistan

RenownedturtleexpertnicolasJ.pilcher,whowasinKarachiforaseminaronthreatstothetur-tlesinpakistan,hasconfirmedthatathirdspecies,theloggerhead(Caretta caretta),wasalsonestingontheseshores.Untilnow,theonlytwospe-ciesrecordednestinginthecountrywereGreenturtles(Chelonia mydas)andasmallernumberofoliveRidleys(Lepidochelys olivacea).byfarthemostwidespreadnestingisbytheGreenturtle,fromtheeasternshoresofsindhprovinceallthewaytothewesternshoresofbalochistan.Recentfindings by the IUCn pakistan team, undertheauspicesofthebalochistanpartnershipsforsustainableDevelopmentproject, havedocu-mentedathirdspecies,theloggerhead,nestingatDaranbeach,some11kmsoutheastofJiwani. positiveidentificationofadultsandhatch-lingswasmadebyDr.nicolaspilcher,Co-ChairoftheIUCnmarineturtlespecialistGroup,andalong-timeturtleresearcherandconserva-tionist.somesixtynestswererecordedduring2009,andtheworkwillexpandinthecomingmonths todetermine the extentof thisnest-ing. loggerheads are known to nest in largenumbers inoman,onmasirahIsland,andasmallnumbernestinYemen,butthisisthefirstrecordofloggerheadnestinginpakistan.thislatestdiscoveryexpandsthenestingrangeforloggerheadsandraisestheirsurvivaloutlookinaclimatewherecriticalnestinghabitatsarebeingrapidlylosttodevelopment.thediscoveryalsoboostsknowndiversityofwildlifeinpakistan. pakistanusedtohostsubstantialnumbersofoliveRidleysalongtheshoresofsindhprovince,butmosthaveceasedtonest.Commercialfish-eriesarethemainreasonforthisdecline;withover1,900activetrawlersoperatingjustoffshore,turtleshavebeenaccidentallylosttofishingnets.trawlingforshrimpandfishisknownasoneofthemajorcausesofseaturtlemortality. luckilyfortheloggerheads,sindh-basedcommercialfisheriesdonotgenerallyoperateasfarawayasthewesternendofbalochistan,sotheturtleshaveavoidedthethreats.turtlescanbesavedfromdrowninginfishingnetsthroughtheuse ofturtleexcluderDevices (teDs),whicharecleveradaptationstonetsthatallowfishandshrimptoenterthenetbutturtlesto

escapethroughaspecialopening.theIUCnintendstoworkwithpartnersinthecountrytohelpintroduceandpromoteteDssothatfish-ingwillhavelessimpactonturtles.thismayalsoresultinthereappearanceofoliveRidleys. Forthepastseveraldecadesmarineturtleconservationprogramshavebeenunderwayinpakistan.mostnoticeableistheworkunder-taken in sindh province, on sandspit andhawkesbay,wherethousandsofturtleshavebeenprotectedthroughhatcheryenclosures.

IUCN 14February2010

New specialist group for Chameleons

ChameleonsareprimarilyfoundinmadagascarandAfrica,withafewspeciesdistributedintheneareastandalongthecoastofsoutherneurope.Althoughsomespecieshavebroaddistributions,manyhaveextremelysmallranges.Importanthabitatsforchameleonsareheavilyimpactedbyhumanactivities,andsomerestricted-rangespe-ciesareparticularlyvulnerable.becauseoftheiruniqueappearance,and, in somecases, theirexceptional ornamentation, chameleons areoneofthemostsought-afterreptilesinthepettrade.tensofthousandsofwildchameleonsarelegallyexportedeveryyear,which,withanillegaltradeatunknownlevels,iscauseforconcern.AChameleonspecialistGroupwasestablishedinFebruary2010toaddressissuesofsustain-abilityandconservationoftheseuniquereptiles.Itsfirstmajorobjectiveistoconductaconserva-tionassessmentofallchameleonspeciesfortheIUCnRedlist.

IUCN SSC e-bulletin march2010

n e W s b R I e f s

VeiledChameleons(Chamaeleo calyptratus)arebutoneofmanyspeciesexploitedinthepettrade.AnIUCnChameleonspecialistGrouphasbeenestablishedtoaddressissuesofsustainabilityandconservationofthesespectacularreptiles.

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genetic variationwithinpopulations,movedthroughintrogressionfromotherpopulations,ordotheyariseasnovelmutations?FElDmAn Et Al.(2009.Proceedings of the National Academy of Sciences 106:13415–13420) examined themolecularbasisofrepeatedadaptationtothetoxinofdeadlypreyinthreespeciesofGartersnakes (Thamnophis) to determine whether

adaptationevolvedthroughnovelmutations,sievingof existing variation, or transmissionofbeneficialallelesacrossspecies.Functionalaminoacidsubstitutionsintheskeletalmusclesodiumchannelarelargelyresponsibleforthephysiological resistance of Garter snakes totetrodotoxin found in their newt (Taricha)prey.phylogeneticanalysesrejectedthehypoth-

esesthattheuniqueresistanceallelesobservedinmultiplespeciesofThamnophiswerepresentbeforethesplitoftheselineages,orthatallelesweresharedamongspeciesthroughoccasionalhybridizationevents.theauthors’resultsdem-onstratedthatadaptiveevolutionoccurredinde-pendentlymultipletimesinGartersnakesviathede novoacquisitionofbeneficialmutations.

neWsbRIeFs

nestingloggerhead(Caretta caretta)havebeendocu-mentedatDaranbeach,nearJiwani,pakistan.

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126 IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010

The patented ventilation system (US PAT# 7,481,183) provides

optimal ventilation and keeps the front glass free of moisture.

Different canopies available for all Exo Terra terrariumsThe digital meters simply

slide into place on the Exo Terra canopy

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Double door to prevent escape

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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,sanJose,CA

sandy Binns services Coordinator

InternationalReptileConservationFoundation,sanJose,CA

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©2010bytheInternationalReptileConservationFoundation,Inc.Allrightsreserved.nopartofthisjournalmaybereproducedinanyformorbyanyelectronicormechanicalmeans,includ-inginformationstorageandretrievalsystems,withoutthepublisher’swrittenpermission.IRCF, Reptiles & Amphibians, Conservation and Natural History (Issn1098-6324)ispublishedquarterlybytheInternationalReptileConservationFoundation,anonprofit,tax-exemptorganization,3010magnumDrive,sanJose,CA.periodicalpostagepaidatsanJose,CA.

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

Additionalcopiesareavailableuponrequestat$6.00eachpluspostage.

*theAdobepDFisoptimizedforwebpublishinganddoesnotprovidethequalityandresolutionofthearchivalprintedversion,especiallynoticeableinphotographsandcomplexgraphics.

JoIn onlIne: www.IRcf.org

membeRsHIP QuesTIonsinfo@IRCF.org,ortollfree1-877-472-3674(U.s.only),orwriteto:IRCF,3010magnumDrive,sanJose,CA95135

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

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

aDveRTIsInG PolIcYWeadvertiseonlynon-livingproducts(exceptfeederinsects).Foradvertisingratesandoptionscontactsandybinns,AdvertisingDirector,atsb@IRCF.orgor3010magnumDrive,sanJose,CA95135.

thesecondinstallmentofouryear-longtributetohenrys.Fitchappearsinthisissue.Athirdinstallmentwill

beintheseptemberissueandthefourthandfinalportionwillappearinDecember.Wewouldliketothink

thatDr.Fitchwouldhaveenjoyedthesenotesandarticlesaboutthenaturalhistoryoftheanimalstowhichhe

devotedmuchofhisprofessionallife.

AlsointhisissueisthesecondinclusionofourInvasivespeciessection,documentingspeciesofamphib-

iansandreptilesinplaceswheretheydonotoccurnaturallyanddonotbelong.Alienspeciesareamajorand

growingconcernfortheconservationofnativeandendemictaxathroughouttheworld.Althoughinteresting,the

accountsthatappearinthissectiondocumentatragedyimposedonnaturalbiotas.WeaskthatreadersofReptiles &

Amphibiansnotcontributetothisproblembyreleasingcaptiveanimalsintonatureorbypatronizingirresponsible

breedersandmerchants.

theeditorsofReptiles & Amphibians

Editors’ Remarks

128 IRCFReptIles&AmphIbIAns•Vol17,no2•JUn2010

Order online at:www.IRCF.org/LBB

Published by the IRCF

Award-winning conservation biologist Fred Burton has released a fascinating new book about saving one of the most endangered reptile species on earth. “The Little Blue Book: A short History of the Grand Cayman Blue Iguana” is the true story of how a noble and charismatic iguana is rescued from the brink of extinction. An engaging read and a beacon of hope for the conservation of reptiles.

Blue Iguana Story in Print

180 pages$19.95 plus p&p Maps & rare photographsRichly illustrated in full colorActual history and natural historyBlue cloth hardcover with dust jacketMore details online at www.ircf.org/LBB

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