IRCF Reptiles & AmphibiAns · IRCF ReptIles & AmphIbIAns • Vol 17, no 1 • mAR 2010 In thIs Issue male calling sites of the Australian toadlet (Uperoleia laevigata) are less densely

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male calling sites of the Australian toadlet (Uperoleia laevigata) are less densely shaded, more likely to be on bare ground, and farther from ponds than those of U. fusca (see article on p. 35).

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southern pacific Rattlesnakes (Crotalus helleri) are one of six species of rat-tlesnakes that partition habitats at Joshua tree national park in California (see article on p. 42).

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back Cover: George R. pisani

Dr. Fitch’s classical study of the Copperhead (Agkistrodon contortrix) is frequently cited by her-petologists as the single most influential publica-tion that triggered their interest in snake ecology (Fitch, h.s. 1960. Autecology of the Copperhead. University of Kansas Publications of the Museum of Natural History (13):85–288).

Front Cover: thomas Kennedy

Fitch’s Anole (Anolis fitchi), named in honor of henry s. Fitch, was formally described by ernest e. Williams and William e. Duellman in 1984 (Anolis fitchi, a new species of the Anolis aequa-torialis group from ecuador and Colombia. University of Kansas Publications of the Museum of Natural History (10):257–266).

IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 1tAbleoFContents IRCF REPTILES & AMPHIBIANS • VOL15, NO 4 • DEC 2008 189TABLE OF CONTENTS

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

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

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

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

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

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

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

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

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

H U S B A N D R Y

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

P R O F I L E

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

C O M M E N T A R Y

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

B O O K R E V I E W

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

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

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

Back Cover. Michael KernTotat et velleseque audant mo

estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus

aut dolor apicto invere pe dolum fugiatis maionsequat eumque

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

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

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

special Issue: a Tribute to Henry s. fitch

P R O F I L E

henrys.Fitch........................................................................................................ Henry S. Fitch as told to Alice Fitch Echelle 2

R E M E M B R A N C E S

henryFitchandthepracticeofnaturalhistory..........................................................................................Harry W. Greene 9Reminiscencesofhenrys.Fitch............................................................................................................William E. Duellman 11henryFitchathomeandinthetropics................................................................................................Robert W. Henderson 12IntheFieldwithhenryFitch........................................................................................................................ David M. Hillis 13naturalhistoryobservationsofhenryFitch............................................................................................. Raymond B. Huey 14memoriesofhenryFitch........................................................................................................................ Michael V. Plummer 14henryFitchasamentorandteacher.......................................................................................................... Richard A. Seigel 15henryFitch:thetwilightofanIncredibleCareer....................................................................................... George R. Pisani 16

F E A t u R E A R t I C L E S A N d N O t E S

DietspecializationbythescarletKingsnake,Lampropeltis elapsoides(Colubridae).............................................................................................................................Harry S. Greene, Edmund J. Zimmerer, William M. Palmer, and Michael F. Benard 18

AsurveyofGravidsnakesatseveralsitesinsouthernWisconsin...............................................................Joshua M. Kapfer 22DaytimeAmphibiansurveysinthreeprotectedAreasintheWesternGreatlakes.....Kenneth D. Brown and Erik A. Beever 26maleCallingsitesintwospeciesofAustraliantoadlets(Anura:myobatrachidae:Uperoleia)attwopondsin

newsouthWales......................... Francis L. Lemckert, Georgina Lemckert, Campbell Lemckert, and Frances A. Lemckert 35huntingtamacoré,Uranoscodon superciliosus.............................................................. Zilca Campos and William Magnusson 38Alligatorsnapperstuckinthemud:evidenceofAestivation....................................................................... Steven G. George 39timberRattlesnake(Crotalus horridus)swimsthemississippiRiver............................................................ Steven G. George 40ADefensiveDisplaybyasmoothearthsnake(Virginia valeriae)............................... Andrew M. Durso and Kevin P. Durso 41sharingtheResource:sixspeciesofRattlesnakesinJoshuatreenationalpark,California......................Harold F. De Lisle 42observationsofAmplexusandovipositioninOllotis[Bufo]occidentalisintheRíosalado,pueblo,mexico..........................

...................................................................................................................... Geoffrey R. Smith and Julio A. Lemos-Espinal 46DiscoveryofGoniurosaurusGeckos(squamata:eublepharidae)innorthwesternGuangdong,China...................................

................................................Zu-Sheng Yi, Zhen-Chang Li, Wei-Liang Wen, Mian Hou, Wen-Hua Lu, and James Lazell 47snakesUsingstumpholesandWindfalltree-associatedsubterraneanstructuresinlongleafpineForests............................

...................................................David A. Steen, Amanda D. Steen, Scott Pokswinski, Sean P. Graham, and Lora L. Smith 49

I N V A S I V E S P E C I E S

IsthenorthernAfricanpython(Python sebae)establishedinsouthernFlorida?..................................................................................................................................................Robert N. Reed, Kenneth L. Krysko, Ray W. Snow, and Gordon H Rodda 52

v ConseRVAtIonReseARChRepoRts:summariesofpublishedConservationResearchReports.............................. 55v nAtURAlhIstoRYReseARChRepoRts:summariesofpublishedReportsonnaturalhistory.............................. 57v neWsbRIeFs........................................................................................................................................................................ 58v obItUARY:Johnthorbjarnarson(1957–2010).................................................................................................................. 60v eDItoRIAlInFoRmAtIon............................................................................................................................................. 63v FoCUsonConseRVAtIon:RescueDarwin’sFrogs..................................................................................................... 64

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thisissueofReptiles & Amphibiansisdevotedlargelytothememoryofhenrys.Fitch(hsF)whopassedawayon8september2009,just

shortofhis100thbirthday.Althoughwesuspectthatmanyofourread-ersarealreadywellawareofDr.Fitch’scontributions,forthosewhomayberelativenewcomerstoherpetology,weprefacethistributeissuewithsomerelevantbackgroundinformation(seealsothe“profile”belowandtheremembrancesbeginningonp.9). As superintendent of the natural history Reservation at theUniversityofKansas(nowtheFitchnaturalhistoryReservation),hsFaccumulatedmorethan32,000capturerecordsof18speciesofsnakesovera50-yearspanoffieldworkfrom1948through1997.Recapturesofmarkedindividualsyieldedinformationongrowth,dailyandseasonalmovements,longevity,populationdensity,andmore.thesedataresultedinnumerouspublicationsprovidingmanyofthemostdetailedaccountsofsnakenaturalhistorythathaveeverbeenpublishedor,forthatmatter,everwillbepublished.theseincludetheclassicsAutecology of the Copperhead(1960)andA Kansas Snake Community: Composition and Changes Over 50 Years(1999).Dr.FitchcontinuedtoconductfieldworkinKansaswellintohis90s,especiallywithtimberRattlesnakes.Fortheseaccomplishments,hsFisappropriatelyconsideredthe“father”ofsnakeecology.

thesefeatsalonewouldmeritthistributeissue,buttheyrepresentonlyafractionofDr.Fitch’sscientificcontributions.hisherpetologicalworksalsoincludelong-termstudiesofseverallizardspeciesinCalifornia,Kansas,andtheAmericantropics.hespentconsiderabletimeinmexico,elsalvador,nicaragua,CostaRica,ecuador,andtheDomincanRepublic,wherehestudiedtheecologyandbehaviorofanoles(andalsodescribedsev-eralnewspecies)andothersmalllizardspecies,anddocumentedthecom-mercialexploitationofiguanas(CtenosauraandIguana)forconservationpurposes.Furthermore,hemademajorcontributionstoourknowledgeofplantsuccession,spiders,birds,andmammals. biologistswiththededication,enthusiasm,energy,longevity,andbreadthofknowledgeofhenryFitcharerare.thoseofuswhohadtheopportunitytocollaboratewithhim,orjusttohavespenttimewithhiminthefieldorataprofessionalmeeting,considerourselvesamongthefortu-nate.WeareindeedpleasedtodedicatethisissueandportionsofthenexttwoissuesofReptiles & Amphibiansasatributetothelifeandaccomplish-mentsofDr.henryFitch.

theeditorsofReptiles & Amphibians

Iwasborn25December1909attheFitchfamilyhomeinUtica,newYork,andtwoweekslaterwasnamedhenrysheldonFitch(aftermy

grandfather,henryAugustusFitch).myfather,ChesterFitch,graduatedfromWilliamsCollegeinmassachusettsandbrieflyattendedharvardmedicalschool,preparingforacareerasamedicaldoctor,butheabruptlyswitcheddirections in favorofanoutdoor lifeasanagriculturist.mymother,AliceticknorCheneryFitch,wasfrombelmont,asuburbofboston,massachusetts,whereherfamilyhadlivedsincethe1600s.shehadgonetofinishingschool,withtraininginmusic,poetry,andotherculturalpursuits.Inthosedays,youngwomenrarelytrainedforaprofessionoutsideofthehome.mymotherenjoyedtheoutdoors,anditwasquiteanexperi-enceforhertomovefromthesuburbsofbostontofairlywildcountryinthewest.AslongasIcanremember,sheusedtotakelonghikes,oftenbyherselforwithmeandmysiblings,margaret,Ruth,andChester,born,respectively,september1908,December1916,andmarch1919. WhenIwasoneyearold,myparentsmovedfromnewYorktosouthwesternoregonwheremyfatherhadboughta“ranch”of116acres,mainlyapearandappleorchardatthesouthendoftheRogueRiverValleyinthefoothillsofthesiskiyous,neartheoregon-Californiaborder.Fromourorchardtothesouth,therewasscruboakinthefoothillsandfirandpineinthemountains,risingmorethan6,000feettothecrestofthesiskiyous,andinmyearlyyearsIrangedfarandwideoverthewildcoun-tryandbecameinterestedinwildlifeandespeciallyreptiles.thecommonsnakesinthevicinityofourorchardweretheGophersnake,theWesternYellow-belliedRacer,andtheCommonGartersnake;theonereallycom-monlizardwasthepacificFencelizard,Sceloporus occidentalis.WealsohadWesternskinksandalligatorlizardsonourland. Iattendedaone-roomschoolwithgradesonetoeight,oneteacher,andkidswhowerebackwoods types.WhenIgraduatedandwent to

medfordhighschool,Ifoundmyselfsomewhatretardedinmyacademicqualifications,butaftergraduatingfrommedford,oneofthelargerhighschoolsinthestate,Iwasatleastaswellpreparedastheaveragehighschoolgraduate.myzoologicalinterestswereinnateanddidnotdependonanyoneperson,butnaturalhistorywasoneofmyfather’smanyinterests,andheencouragedmyowninterestbyconversationandbyhavingmanybooksonthesubject.IwasfascinatedbyanykindofwildanimalIsawandespe-ciallyreptiles.Iremembergrabbinglargebullsnakes,becausetheywerecommonontheranch.WhenIhandledabullsnake,Iwasenthralledandalittleafraid,andoftenmyhandswouldbebleedingfromthebites.therealbonuswasinseeinghorrifiedadultsscatter.Itwasquiteafeelingofpowerforafive-year-old.Iamsureitstimulatedmyinterestinsnakes.Feelinglikeasnakecharmer,Icouldimpresspeopleandnotquiteunderstandwhytheyweresoafraid.Wehadfewnearneighbors.oneofthem,earlschuchard,wasmybuddy,buthedidn’tapproveofmyinterestinsnakes,andheoncetoldme,“myDadsaysthatoneofthesetimesyouwillgrabholdofthewrongkindofsnake,andthat’llbetheendofyou!” IenrolledattheUniversityoforegonasazoologymajorin1926whenIwas16,butIdidnotexcelasanundergraduate.Iwasdisillu-sionedbythecollegecoursesthatIhadinbiologyandzoology,becausetheZoologyDepartmentattheUniversityoforegonhadastrongmedicalschoolorientation.thosewhodidn’tmakethegradetobecomedoctorseitherhadtochangedirectioncompletelyorteachbiology.therewasnoprofessorinthedepartmentwhohadanyinterestinthenativefaunaor

Henry s. fitch1

henrys.FitchastoldtoAliceFitchechelle

1originallypublishedinCopeia2000(historicalperspectives,p.891),copyrightedmaterialreprintedwithpermissionoftheAmericansocietyofIchthyologistsandherpetologists(AsIh).

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whocouldidentifyatoad,amouse,orasnake;thus,Ihadnostimulustobeazoologistandnorolemodelforazoologicalcareer.still,Ipersistedasazoologymajorwithoutanyclearideaofmyfutureprofession.InfactIdid,forawhile,contemplateswitchingtogeologyasamajorandtookanumberofcoursesintheGeologyDepartment.AsfarasIwasconcerned,thesecourseswerefarmorestimulatingthananybiologycourseIwasabletofindattheUniversityoforegon. IknewthatIwantedtobeabiologistbuthadnoconceptofthecareeropportunities,whichinthosetimeswereverylimitedcomparedwiththoseavailablenow.professorearlpackard,apaleontologistattheUniversityoforegon,knewmyfatherandmadegeologyfieldtripswithsummerclassestoourpartofthestate.ItookhiscourseattheUniversityoforegon,andhestronglysuggestedthatIgototheUniversityofCaliforniaforgradu-atework.WhenIfirstenrolledatU.C.berkeley,IchoseJosephGrinnell,thedirectorofthemuseumofVertebrateZoology,asmygraduateadvisor.WhilestillattheUniversityoforegon,IreadthemassivevolumesonthereptilesofthepacificCoastandGreatbasinbyJohnVanDenburgh,whowastheherpetologistattheCaliforniaAcademyofsciences.Inhisratherlongspeciesaccounts,hequotedfrompublicationsofGrinnell,andespe-ciallythoseofCharlesl.Camp,whohadbeenagraduatestudentunderGrinnellandwhohadwrittensomepapersthattouchedonnaturalhis-tory.Camp’smaincontributiontoherpetologywashisClassification of the Lizards,andhismaininterestwasmorphology.however,ononeoftheUniversityofCaliforniafieldtrips,hevisitedtheturtlemountainsoftheColoradoDesertandwroteapaperonthedesertlizardswithquiteafewbehavioralnotesandsometaxonomy,andIwasimpressedbythis.IplannedtoenrollunderCampforgraduatestudybutwassurprisedanddisappointedtofindthatCamphadbecomeapaleontologistintheGeologyDepartmentandthathenolongerworkedwithrecentanimals.ItooktwoofCamp’scoursesinmyfirstyearatberkeley:theVertebrateskeletonandelementaryVertebratepaleontology.Ienjoyedbothcoursesandfoundthemprofitable. GraduateschoolwasmuchmoreinterestingandexcitingthanmyundergraduateworkattheUniversityoforegon.WhenIcametothemuseumofVertebrateZoology(mVZ)in1931,ithadjustmovedfromanovercrowdedandinadequatebuildingonthenorthsideofthecampusintothebrand-newlifesciencesbuilding.theentirewestendofthebuild-ingwasoccupiedbythemuseumwithitslargevertebratecollection.thereweregraduatestudentsdoingfascinatingfieldstudiesofdifferentkindsof

animals,someofwhichweretotallynewtome,andIbeganmeetingpeoplewithbasicinterestsoverlappingmyown.myfirstdayinthemuseumImetastudentwhowasjustfinishingamaster’sdegreestudyingmountainbeavers,primitiveaplodontidrodentsthatIhadneverheardofbefore.hehadsomeincaptivity,andIwasfascinatedbythem.AldenmillerhadjustgottenhisdoctoraldegreetheyearIcame.DuringmyfirstsemesteratmVZ,Grinnellmetwithmeandthreeothernewstudentsinweeklyorientationsessions;hepresentedeachofuswithmiller’spublishedthesis(freshoffthepress)onCaliforniashrikesandsuggestedthatweusethisasamodelforadetailedstudyofavertebrate.Wewereallimpressedandinspiredbymiller’sstudy.Grinnellhimselfwasprimarilyanornithologistandsecondarilyamammalo-gist,butheknewverylittleaboutreptiles.hehadpublishednaturalhistorynotesonreptiles,butwhenIbeganstudyingalligatorlizards,hecouldnothavetoldmehowtodistinguishsexintheselizards. Grinnell,uptothisstageinhiscareer,hadasomewhatnegativeatti-tudetowardgraduatestudentsandwasreluctanttospendtimeonthem,becausehewasveryactiveinresearchonbirdsandmammalsandalwayshadfieldprojectsinprogress.overtheyears,however,hisinterestingraduatestudentsgrewstronger.Ididnotseemverypromisingatfirst,Iamsure,buthesensedthatIwashighlyinterestedinanimalsingeneral,andparticularlyinreptiles,andwasquitetolerantofmyshortcomingsinotherdirections. IalwayshaveadmiredsomeofGrinnell’smethodswithregardtograd-uatestudents.WhenIfirsttalkedwithhim,Isupposedthathewouldhavesuggestionsformygraduateresearch.Instead,heaskedmewhatIwasinter-estedinandsuggestedthatIlistasmanypossibleprojectsasIcouldthinkofandcomebacktoseehiminafewdays.thenexttimeItalkedwithhim,hesuggestedthatIcutthislistdowntothethreethatseemedthebestofthelot,takingintoconsiderationforeachprojectpracticality,predictedtimetocompletion,potentialscientificvalue,questionstobeanswered,travelandfundingrequirements,andprospectsforsuccessfulcompletion.FollowingGrinnell’ssuggestions,Inarrowedthepossibilitiesbyseveralstagesandfinallysettledonalligatorlizardsformymaster’sresearch. Among thosewhodetermined the courseofmycareer, I shouldmentionespeciallyprofessore.Raymondhall,whobroughtmetotheUniversityofKansas(KU)totakechargeofthenaturalhistoryReservation(nhR)andtoteachecology.IfirstmethimwhenhewasayoungassistantprofessorandcuratorofmammalogyatmVZ.hewasoneofGrinnell’searlystudentsandwassohighlythoughtofthatheworkedintoapermanentpositioninthemuseum.AftermyfirstacademicyearofgraduatestudyatmVZ,Ienrolledinasummercoursetaughtbyhall.Atthetime,hewasstudyingthemammalsofnevada,andhepublishedatomeonthisworkafewyearslater.therewereabouteightornineofusinthesummercourse,andthefieldworkconsistedmainlyoftrapping,especiallysnap-trapping,smallmammalsandpreparingthemasstudyskins.Wealsocollectedothervertebrates,includingafewbirds,andforme,especially,reptilesandsomeamphibians.thiswasmyfirstexperiencewiththehighdesertherpetofaunaoftheGreatbasin.thelizardsandsnakesweremostlynewtome,anddur-ingthecourseofthisfieldtrip,wevisitedmanylocalities,coveringmuchofthestate.samplingbylive-trappinginthehighmountainsandonthedesertflatswasagreatlearningexperienceforallofusintheclass. therewere,ofcourse,manyotherinterestingpeopleIgottoknowwhileingraduateschool.onepersonwhoturnedouttobemygoodfriendduringthecourseofgraduateworkwasDonhatfield,whogotamaster’sdegreeatmVZ.hehadagiftforwriting,becameinvolvedinhollywoodmovies,andevidentlyhadacareertherethatwasnotmainlybiological.Also,therewasWardRussell,hall’sfieldassistant,whodevisedthedermes-tidbeetlemethodofcleaningvertebrateskeletons. IjoinedAsIh(AmericansocietyofIchthyologistsandherpetologists)intheearly1930s.Inthosedays,gettingtothenationalmeetingcouldinvolveasmuchastwoweeks’absencefromtheworkplace,andwithmylimitedfunding,thecostwasprohibitiveuntilthewesterndivisionofthesocietymetinberkeleyinJune1934.Atthismeeting,IfirstmetCarlhubbsandhiswifeandchildren.theywereonacollectingtripcoveringmanyofthewestern

Postscript.2—henry Fitch was interviewed by Alice Fitchechelle,herhusbandtony,andtheirchildrentysonandlena.theinterviewwasconductedon6september1998,theFitch’s52ndweddinganniversary,atthefamilyhomeontheFitchnaturalhistoryReservation,lawrence,Kansas.Dr.Fitchhasreceivedmanyhonors.In1950,hereceivedtheGeorgemercerAwardoftheecologicalsocietyofAmericafortheyear’sout-standingecologicalpaper,“ecologyoftheCaliforniagroundsquirrel on grazing lands” (American Midland Naturalist39:513–596).hispublicationsspantheperiodfrom1933to1999andarestillinprogress.In1997,theAmericansocietyofIchthyologistsandherpetologistsestablishedthehenrys.FitchAwardforexcellenceinherpetology,tobemadeannu-allybythesocietytoadeservingherpetologistforfieldresearch.Dr.Fitchwasthemajoradvisorof18mastersand14doctoralstudents.severalnamesarewellknowntoherpetologists:Johnlegler,Russellhall,Dwightplatt,michaelplummer,Richardseigel,RobertWebb,andRoberthenderson.

—margaretm.stewart,historian

2this“postscript”wasamendedtothis“perspective”whenitwasoriginallypublishedin2000.

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states.hubbswasinterestedtoknowthatIwasworkingongartersnakes,andhegavemesomelocalitiesforThamnophis elegans vagrans thatwerefarthereastthantherangewasknowntoextendatthattime. GeorgemyerswastheichthyologistatstanfordUniversity,andIknewhimfromseveralvisitstothemuseumthere.myerswasahot-shotichthy-ologistwhogothisph.D.atstanfordandrevitalizedichthyologythereafterJordanpassedon.myerswasfriendlywhenIfirstmethim.Atstanford,Victortwittywasstudyingwesternnewts,andfoundthat“Triturus torosus”wasacompositeofthreedistinctspeciesthatoverlappedinsomeareasbutwerereallyverydifferentincolor,habitat,andhabits.thedifferenceswerequitesharplydefinedinthelarvae,too.twoofthespecieswerelimitedtoCalifornia,butthethirdwasmuchmorewidespread,andthatwastheonewehadinoregon.Ifoundthattwitty’sTriturus (nowTaricha)similans hadbeennamedlongbefore,asT. granulosus,andIpublishedanoteonthisin1938.Afterthenotehadbeensubmitted,IproudlymentionedmydiscoverytomyerswhenhewasvisitingatmVZ,buthetoldme,“I’mworkingonTriturus,”andheseemedveryresentfulthatIhadinfringedonhisterritory.Afterthat,hewasconsistentlyhostile.Whenmydissertationwaspublishedhewroteascathingreview.therewasnothinggoodinitatall,accordingtohim,andheevencriticizedmyusingtheGermanwordArtenkreis forthegartersnakes(inthoseyearsbeforeWorldWarII,manypeoplehadabiasagainstanythingGermanic).subsequently,CarlhubbswroteareviewforAmerican Naturalist thatwasmorepositiveand,inmyopinion,moreinsightful.therewasmutualantipathybetweenhubbsandmyers,representingmichiganandstanford,themainichthyologicalcentersinthecountryatthattime. InevermetDavidstarrJordan,butmybrother- in-law,Gaetonsturdevant,tookacourseinichthyologyfromJordanatstanfordinthelate’20sorearly’30s.earlheraldandRobertRushmillercametomVZforexperienceworkingwithGrinnellmylastyearthere(1937). From1934on,IwasemployedasateachingassistantinGrinnell’scoursesandtaughteconomicVertebrateZoologyinthefallsemesters.Itwasacourseforforestrymajors,anditemphasizedanimalssuchasbea-versandwoodpeckersthatareimportantinforestecology.Inthespringsemesters,IassistedinZoology113,acourseinvertebratenaturalhistoryforzoologymajors.thiswasacourseinbirdsandmammals,withasmat-teringofherps,thatwastaughtmainlybyGrinnell,withhalltakingthemammalpartofit.ItwasthebestcourseIhadevertakeninmystudentcareer.Grinnellgaveexcellentlectures.Iassistedinitoveraseriesofyearsandlearnedmoreeachtime. mysummerswerespentinthefield.severaltimesafterthesummerof1931,whenIworkedinnevadawithhall,IwentonmuseumfieldtripsasamemberofanmVZteam.onthesetrips,theobjectivewastocollectascompleteasampleaspossibleofthevertebratefaunaofaspecificlocalityorarea.WespentpartofasummercollectinginthevicinityoflehmanCaveinnevadatosamplethemodernfaunaasabasisforcomparisonwiththepleistocenecavedeposits.everysummerIwasalsobusywithmyfieldresearchinseveralwesternstates,butmostlyinCalifornia,oregon,andnevada,withmuchlesstimeinUtahandWyoming.WhenIwascollect-inggartersnakesformydissertation,IoftencollectedpocketgophersforGrinnell.heneededthemfromveryspecificlocalitiesandpaidmebythegopher,notverymuchpergopher,butenoughtofinancemyownfieldwork. Aftertheconferringofmydoctorateinmay1937,Iworkedinthemuseumpreparingmydissertation forpublication in theUniversityofCaliforniapublicationsinZoology.Duringnovember1937,IlivedwithJeanm.linsdaleandfamilyatthenewlydedicatedhastingsnaturalhistoryReservationinmontereyCounty,California,andwasduetocomebackandspendmoretimeinthespringof1938.IwasthefirstofmanylinsdalefieldassistantsatthehastingsReservation.ItbecameroutineformVZgraduatestudentstospendtimewithlinsdaleathastings,benefitingfromhisguidanceintakingfieldnotes.buttherewerecomplaintsfromsomethatlinsdale’sstyleofbitingsarcasmoftentraumatizedstudents.linsdale’scareerendedtragicallywithdeterioratinghealth,lossofmemoryandeyesight,andprematuredeath(inthe1960s).IdidnotreturntoworkwithlinsdaleasplannedbecauseI

hadqualifiedforagovernmentjobandwashiredbythebureauofbiologicalsurvey(whichlaterbecametheU.s.FishandWildlifeservice).thebureauhadstartedoutasaresearchorganizationwithasmallcadreofleadingnatural-istsunderC.hartmerriam.bythetimeIwashired,thesurveyhadexpandedandchangeddirection,withpestcontrolasamajorpriority. thebureauneededafieldbiologisttocollaboratewiththeForestservice at the Forest and Range experiment station’s san JoaquinexperimentalRangeinthesierrafoothillssouthofYosemitenationalpark.Iwenttoworkonthisjobearlyin1938,studyingtheecologyofrangerodents.Inorderoftheirimportanceoncattlerange,theserodentsweretheCaliforniaGroundsquirrel,thesanJoaquinpocketGopher,andthetulareKangarooRat.AlsotheAudubonCottontailwasofsomeimportance.Inaddition,therewereseveralkindsofnativeratsandmice,includingNeotoma fuscipes, Peromyscus boylei,andPeromyscus truei,thatwerecommoninthearea,nottomentionMicrotus californicus,whichwasratherlocalizedbutabundantinthefewplacesthatweremoistenoughforit.Groundsquirrelswerethefocusofmyfieldwork.Ihadbeenfamiliarwithgroundsquirrelsinoregonsinceearlychildhoodandhadshotandtrappedmanyofthem.AtthesanJoaquinRange,theyweresuperficiallythesamebutmuchwilierandhardertotrapthantheircounterpartsinsouthernoregon.theywereexceptionallyalertandsuspiciousofanystrangeobject,includingsteeltrapsplacedinburrowentrances,andgenerallyavoidedtrapsbycirclingorjump-ing.Inunderstandingthisdifference,Ibecameawarethatduringthesnakeseason,fromearlymarchthroughnovember,thegroundsquirrelswereatconstantriskofbeingbittenbyrattlesnakes—theironemostimportantnaturalenemy.thesepopulationsthathadbeenexposedtorattlesnakepre-dationformillionsofyearswerebehaviorallyquitedifferentfromgroundsquirrelsfrommorenorthernareaswhererattlesnakeswereeitherabsentorweremuchmorerecentandlessabundant.Ihadbeeninterestedinsnakesfrommystartthere,andmoreandmoreIbecameinvolvedwithrattlesnakesaspredatorsonthelocalrodents.Asaresult,IstartedmarkingandstudyinglocalpopulationsofthepacificRattlesnake. Jessenelson,superintendentatthesanJoaquinRange,disapprovedofanaturalhistoryapproachandthoughtthatIshouldconcentrateinsteadonso-calledpestcontrol.Itgottothepointwherenelsonwouldassignafieldworkerthechoreoftimingmydayinthefield—howmuchtimewasspentonsnakes,especiallywhenwehadacrewout.soonthedataseemedtoindicatetomyemployersthatIwasspending36hoursaday(myowntimeandthatofcrewmembers)onsnakesratherthanonrodentcontrol,andIwasorderedtostopthesnakework.Afterthat,Irarelybroughtlivesnakestotheheadquarters.butwheneverIcaughtoneinthefield,Iwouldprocessitthere,“bootlegging”thispartofmyresearch. IwasbittentwicebyrattlesnakesinthecourseofmyworkattheexperimentalRange.thebiteswerenotlifethreateningbutweretraumaticexperiences,andasaresponse,Ichangedmycatchingandhandlingtactics.Forthefirstbite(springof1938),Iwenttoadoctor’sofficeandhadashotofserum.Iwasallergictotheshot,brokeoutinarash,andsufferedalmostasmuchfromthetreatmentasfromthebite.IwasbittenwhenIdroppedthesnakeintoabag;thesnakecouldstrikefasterthanIcouldwithdrawmyhand.thesecondbite(springof1940)happenedalmostthesameway.Itwasafirst-yearsnakerecentlyemergedfromhibernation,anditsshotofvenomseemedevenmorepotentthanthatofthelargersnakethathadbittenmein1938.Ihadchangedhands,fromthetimeofmyfirstbite,andneveragaingraspedasnakewithmyrighthand,whichwasleftfreetowrite.sincethesecondbite,Ihaveneverletgoofapoisonoussnakewithoutfirstholdingdownthehead,havinglearnedthehardway.besidesashotofserum,acceptedtreatmentforabiteatthattimewastocutatthesiteofthebiteandthenapplysuction.Forthefirstbite,Ihadtogetbacktomycarandthendrivebacktoheadquarters.thecarwasaboutone-thirdmiledistant,andIjoggedbacktoitanddrovetoheadquarters,andmyfriendharoldbiswelldrovemetomaderafortreatment.thesecondbitehappenedbackattheheadquarters.Idroppedthesnake,anditbitmeonthefinger,andreflexively,Ijerkedbackandslungit20feet,whichmay


havecausedmorevenomtobeinjected.IwasworkingwithbenGlading,andhedrovemeintomadera. Iwasdraftedinthespringof1941,onfourdays’notice,andIthinkpartlyormaybeentirelybecauseofmyage(32),IwasputinthemedicalCorpsandwasassignedtoastationhospital.Duringthatsummer,IwassenttoWilliambeaumontGeneralhospitalinelpaso,texas,andhadathree-monthcourseoftrainingasanarmypharmacist.soonaftermyreturntoCampsanluisobispo,California,inseptember1941,IwasreleasedfromthearmyonthegroundsthatIwastoooldtoserve,beingover26,andthiswaswhentheone-yeartermofdraftees(exceptforthose26orolder)wasextendedindefinitely.theterm-extensiontriggereddrafteeresentmentandriotinginarmycampsacrossthecountry,butIwasgladtobeoutofthearmyandwentbacktomyoldjobatthesanJoaquinRange. myreliefwasshort-livedbecauseIwasrecalledtoactivemilitarydutysoonafterpearlharborwasbombed.IwasatCampsanluisobispoforawhile,thenassignedtoahospitalatsawtelle,California,subsequentlyassignedtothe348thstationhospitalatCampbeale,California,andfinallywasshippedoverseasinlate1942ontheVictoryship,KoKomo.theshipcarriedabout5,000ofusacrossthestormynorthAtlanticinquartersthatwerecrowdedanddirtyandthatreekedofsickodorsbecauseoflackofadequateventilation.Wewerepartofalargeconvoyandhaddestroyerescort.theholdoftheshipwaspartitionedintomanycompart-mentsthatcouldbeusedasairspacetokeeptheshipafloatafterpossibletorpedostrikes,andtheunannouncedandfrequenttestingoftheelectroni-callyoperatedslidingdoorsthatsealedoffourcompartmentswasagrimandpersistentreminderoftheexpendabilityoftheindividualintimesofwar.myunitwassentfirsttoCardiff,Wales,andafteraperiodofmonthsthere,weweretransferredtollandudnoonthenortherncoastofWales,latertonearnottinghamandtotwoorthreeotherplacesinengland.AfterD-Day,wewereatprestwick,scotland,afirststopforflyingcasualties,andwewereoftenroutedoutofbedduringthemiddleofthenighttocarrythewoundedintoourhospitalortransferthemontoanotherplaneortrain.Finally,aftertheinvasion,weweresenttoFranceandeventuallytooccupiedGermanyatbremerhavenwhereIspentaspring,summer,andfall.bythattime,thewarwaswon;therewasapointsystemforreleasingveterans,andIgotoutofthearmysoonerthansomeofmycolleaguesonthebasisofmyadvancedage.IwassentbacktoCampbealenearmarysville,California,whereIwasreleased,andafteranightofhitchhiking,Iarrivedbackhomeatmedford,oregon,innovember1945afteralmostfiveyearsinthearmy. Armylifeduringthewarwasnotveryconducivetopursuitofzoologi-calinterests.Whenwehadtimeoff,Ioftenwatchedbirds.Alongthecoastnearllandudno,therewerelargecoloniesofnestingseabirdsonalongrockypeninsula,anditwasagreatplaceforbird-watching.earlier,whenIwassenttobeaumontGeneralhospital,Idrovemycartoelpasofromsanluisobispo.Ihadweekendsfreeandexploredfarandwide.becauseIhadneverbeforedonefieldworkinthatpartofthecountry,thereweremanykindsofherpsthatwerenewtome,andIsentmycollectionsbacktomVZ. Whilestationedatvariousplacesinengland,Iwasabletomakeweek-endtripstolondon,whereIfrequentedthezoo,thebritishmuseum,andmanybookstores.Atthattime,londonwasbeingbombedintermittently.theinfamous“buzzbombs”wereahazard,andIrememberhearingmanyexplosions,butnonewaseververyclosetowhereIhappenedtobe. AssoonasIgothometomedford,oregon,afterthewar,IwrotetotheFishandWildlifeserviceindicatingthatIwasreadytoreturntomyjob.theU.s.presidenthadpromisedthatallGIswouldbeentitledtotheirformerjobsuponbeingreleased,butIgotadiscouragingreplyfromtheFishandWildlifeservicesayingthatmyoldjobnolongerexisted.theygavemethenameandaddressofthedirectorofRiverbasinsurveys.thesesurveys,initiatedbythesoilConservationservice,wereinventoryingver-tebratesofeconomicimportanceinthecentralstates.Italkedwithapersonwhohadrecentlyreturnedfromthewarandgonetoworkforthem,andhewasveryencouraging.butIwasconsiderablyirkedbecauseIwasnotbeingallowedtoreturntomyformerjobashadbeenpromised,soIwrotealetter

ofcomplainttotheFishandWildlifeservice.ItoldthemthatIwasnotatallinterestedinworkingonRiverbasinsurveysinapartofthecountrythatwasunfamiliartome,andthatIwasdisappointedthatthegovernmentwasnotfulfillingitsresponsibilityafterhavingpromiseddrafteesthattheirjobswouldbewaitingforthem.Itseemedtomethatthegovernment,especially,shouldliveuptoitspromise.Intheletter,ImentionedthatIwasnotapplyingfortheRiverbasinsurvey,butinsteadIwasreturningtothesanJoaquinexperimentalRange(sJeR)onmyowntosalvagewhatIcouldofmyresearchthere.Idon’tknowwhatbecameofthatletter,butitmusthavecausedsomestir.Afteraboutthreeweeks,Ireceivedareply:theyhad“found”somefundsforsalaryandwouldbegladtohavemegobacktothesJeRandfinishupmyprojectsthere.IreturnedtothesJeR,spentallof1946thereandalsothespringof1947,dividingmytimebetweenfieldworkandwriting.on6september1946,ImarriedVirginiaRubypreston,whomIhadmetatapartysoonafterreturninghomefromthewar.WewereallowedtoliveinalittlehousenearthesJeRheadquarters.Icompletedaseriesofpapersforpublication,andproductionwasinhighgear.butdissensiondevelopedbetweenmybossesintheFishandWildlifeserviceandtheForestservice,resultinginmytransfertolouisiana. Inmay1947,VirginiaandIdroveourowncarandagovernmentcarfromthesanJoaquinRangetoAlexandria,louisiana,andlivedinahous-ingprojectatnearbyleesville.Duringouryearinlouisiana,IworkedinthenationalForest,20–40milesfromleesville,studyingquail,mourningdoves,armadillos,cottonrats,anddeer.myworkinlouisianawouldhavecontinuedexceptthatIwasinvitedbye.RaymondhalltoapplyforthepositionofecologistattheUniversityofKansas(KU).hallhadbeenthemammalogistatmVZatberkeley,butwhenhelostouttoAldenmillerfor

henryFitchinArmypharmacyschool,WilliambeaumontGeneralhospital,elpaso,texas,August1941.theuniformwasaWorldWarIcavalrysuitbecausethatwasallthatwasavailableatthecampatthattime.

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thedirectorship,hereturnedtohisalmamaterasdirectoroftheUniversityofKansasnaturalhistorymuseumandaschairmanofthe(then)ZoologyDepartment.heknewmewell fromthetriptonevadaandfromourassociationatmVZ.IhadalsoworkedforhimonhismajorstudiesofAmericanweasels.hewasawareofmyworkwithlinsdaleatthehastingsReservationandhadmeinmindforthejobonthenewlycreatedreservationattheUniversityofKansas.thislandhadbeenownedsince1910bytheUniversityofKansas,andhallhadpersuadedthechancellorthatitsbestusemightbeasanaturalhistoryreservation.IcametoKUandgaveaseminaronmyworkatsJeRandgotthejob.thiswasaverystrategictimetostartinuniversityteachingbecausetheGIbillofRightshadbeenpassed,anduniversitieswerefullofreturningveteranswhosehighereducationwasbeingfinancedbythegovernment.AtKU,atleasthalfthecoursesinthedepart-mentwerebeingtaughtbygraduatestudents.Istartedmyteachingcareerwiththetitleofinstructor,aswascustomarythen,inthefallof1948.Iwaspromotedtoassistantprofessoraftermyfirstyear,andfinallytofullprofes-sorin1958.FromJuly1948throughFebruary1950,welivedoncampusinthesunnysidehousingprojectnearwhereAllenFieldhousestandsnow,andon1march1950movedintothenewresidenceonthereservation. IwasbroughttoKUtobesuperintendentofthereservationandtoteachecology.Uptothispoint,KUdidnothaveanecologist.IthinkprofessorWorthyhorrtaughtacourseinplantecology,andmr.mcnairhadtaughtanimalecologyafewyearspreviouslybuthadsincedeceased.ecologywasjustbeginningtocomeintoitsown;Ihadneverhadacourseinecologymyself.Itaughtitduringfallsemesters,andforthefirstfewyears,Ihadlargeclassesofabout30–35students.In1968,whenwereturnedfromsabbaticalinCostaRica,myteachingdutieswereswitchedfromecologytonaturalhis-tory,becauseKUwasacquiringseveralecologistsofdifferenttypes.tome,ithasbeenmuchmoresatisfyingtoteachnaturalhistorythanecology. Asamammalogist,hallwasanxioustohavesomemammalresearchdoneonthereservation.Iwasmammal-orientedaftermyworkatthesanJoaquinRangeandstartedsomelarge-scalestudiesofsmallmammalswithlive-trappingandmarking,andthatwashow,foryears,Ispentamajorpartofmytime.AlsoIstudiedlizards,includingFive-linedskinksandGreatplainsskinks,nowalmostgonebecauseofhabitatchanges,butatearlierstagesofsuccession,theywereabundantandconspicuous.Afterseveralseasons,Idevelopedalive-trapforcatchingsnakes,andputoutlongerandlongertraplinesforthesnakesthatcametohibernateathilltoprockledges.In1957,Ilearnedhowtotraptheminthefields,wheretheydisperseinsummer,byputtingupdriftfences,andthiswaslaborintensive;asubstantialportionofmyfieldworkhereonthereservationhasconsistedoflive-trappingsnakes. everypartofthereservation’ssquaremilehaschanged,totheextentthatitishardlyrecognizableasthesameareaIfirstsawmorethan50yearsago.Fromalmostanystandpoint,itwouldlookentirelychanged,butthewoodedparthaschangedrelativelylittlecomparedtotheoriginallyopenareas.thewoodlandshavechangedthroughthedyingoutofthelargeAmericanelmsthatwerethedominanttrees.thesewerereplacedbyashandavarietyofothertreespecieswiththeclimaxspeciesconstantlygainingandspreadingtoplaceswheretheyformerlyhadbeenabsent.osageorange,honeylocust,andmulberryarepioneerinvadersofthelandthatwasformerlycultivatedorthatwasovergrazedpasture,andontheseformeropenareas,denseweeds,brush,shrubs,andseraltreesbecameprominentasstagestowardaclimaxforest. everyanimalspecieshaschangedindistributionandabundance,andingeneral,thegrasslandspecies,especiallythoseofshortgrass,havedisappeared.IthasbeenalongtimesinceIhavefoundabullsnakeonthereservation,butinthefirstfewyears,theywerecommoninthepastureareas.tallgrassspeciesliketheYellowbelliedRacerandtheprairieKingsnakearestillhere,buttheyarebecomingmuchmorescarce.evensomeforestspecies,notablythetimberRattlesnake,havedisappearedcompletelyfromoursquaremile.Althoughitshabitatisforest,itrequiresopensunnyplacestobask,andthecontinuouscanopythathasdevelopedhaseliminatedbaskingplaces.Despiteconsiderableanthropogenicmortality,mostofthetimberRattlers,perhaps20,caughtinthelastfewyearshavebeenfromtheKU-ownednelsontract

adjoiningthereservationtothenorthorfromfarmsimmediatelyadjacenttothenelsontract.Irememberonlyonerecordinrecentyearsfromthereservation,andthatwasneartheheadquarters.thatoneadultmaletimberRattlermusthavewanderedfarfromhishomerange. Ihadalwayswantedtodofieldworkwithreptilesinthetropicsandfinallywasabletodosoatage55when,in1965,ItooktheorganizationfortropicalstudiescourseinCostaRica.Duringthecourse,wecoveredmostofthecountryandbecamefamiliarwithitsfauna,andIlaidthegroundworkforlaterstudyoflocallizardpopulations.In1967–1968,onsabbaticalfromKU,IreturnedtoCostaRicaonanationalscienceFoundationgrantwithatruckandcamperandmywife,daughterAlice(20),andsonChester(14).ouroldersonJohnwasatthattimebasedinhawaiiandworkingforthesmithsonianInstitutionontheirpacificbirdproject.Withthehelpofmyteam,AliceandChester,Iestablishedtran-sects(north–southandeast–west)spanningCostaRicawith14studyareaswithindividuallymarkedpopulationsof15lizardspecies.sixofthesespe-cieswererepresentedattwoormoreofthestudyareasrevealingtheeffectsonreproductivecyclesofcontrastingclimates,fromrainforestandcloudforesttoxericscrub.eachareawasrevisitedatsix-weekintervalsin1968,andsamplingofthesesameareaswascontinuedthroughtheearly1970s. Anothermajorprojectwasacomparativestudyofanoles,about50species,frommexicotoecuador,mostlyonthemainland,butincludingseveralintheDominicanRepublic.habitat,seasonalscheduleofactivity,breedingseason,dewlapdisplays,andsexualsizedifferencewerefoundtobecloselyrelated. AthirdprojectwasanaturalhistoryandconservationstudyofGreenIguanas(Iguana iguana)andctenosaurs(Ctenosaura similis)innicaraguaandotherCentralAmericancountries.InmanylatinAmericancountries,theselargelizardsconstituteanimportantfoodsource,butoverhuntingandhabitatlosshaveeliminatedthemorcauseddrasticreductionthroughmuchoftherange.Appropriateconservationmeasureshavethepotentialtorestorepopulationsofthesehardyandprolificlizards,withatremen-douseconomicbenefittothelocalpeople.Withmyformerstudent,bobhendersonofthemilwaukeepublicmuseum,Ibeganfieldworkin1976;morethan1,000ctenosaursand343GreenIguanaswereexamined,mea-sured,andweighed,mostlyinthemarketsofnicaraguantownsandvil-lages.Atthattime,nicaraguawasstillcontrolledbythesomozadictator-ship.ourstudywassponsoredbythebancoCentralofmanagua.theInternationalFundforAnimalWelfaresponsoredseveralmoretripstonicaraguainthe1980s.Duringthesevisits,IworkedwiththeGovernmentof Reconstruction (sandinista) and their conservation organization,InstitutonicaraguensedeRecursosnaturalesydelAmbiente(IRenA),andaFive-Yearplanforiguanaconservationandrestorationwasinstituted. thethreemajorthrustsofmytropicalresearch,describedabove,over-lappedintimeandstudyareas.thefieldworkspanned20yearswithatleastonetripannuallytothecountriesinvolved,includingmexico,Guatemala,elsalvador,honduras,nicaragua,CostaRica,panama,ecuador,andtheDominicanRepublic. Iretiredin1980andamenjoyinglifewhilecontinuingsomeofmyformeractivities,theonesthatImostrelish,andthatincludestrappingsnakes.Inthe1980sand1990s,Iwasinvolvedwithrattlesnakeroundupsinoklahoma(WesternDiamondbacks)andKansas(prairieRattlers).likemostherpetologistsandconservationists,Iamopposedtotheroundups,buttheydooffergreatopportunitiestolearnmoreabouttheecologyofthespeciesandhowtoconservethem. WhenIretired,IwastoldbytheKUadministratorsthatVirginiaandIcouldcontinuetoliveintheresidencehereonthenaturalhistoryReservationaslongasthatwastotheadvantageoftheuniversity.Wetrytobeuseful,andanyonewhohasanyinterestinnaturalhistoryorecologyisencouragedtovisitandusethearea. mywifeVirginiahadnobiologicalbackgroundexceptforahighschoolcourseinbiology,butshehasalwayshadanavidinterestinnaturalhistory.snakehuntswereprominentduringourcourtship,andshereaduponherps


andbecamewellversedinherpetology.Duringourearlyyearsonthereserva-tion,Virginiaoftenaccompaniedmeinthefieldandhelpedinmanyways,includingmakingplantsurveysandassemblingspidercollections.Whiletendingtraplinesforbothsmallmammalsandsnakes,Virginiarecordedthefielddata,makingmyhandlingofliveanimalsmucheasierandmoreefficient.shealsotypedmostofmymanuscripts,andwehavealwaysreadproofstogether,evenonourweddingnight.Virginiahasnotdoneasmuchfieldworkwithmeinrecentyearsbecauseofhealthproblemsandsusceptibil-itytopoisonivy,chiggers,andticks.however,shestillhelpsmeinpreparingliveanimaldisplaysforvisitingschoolchildrenandinnumerousotherways. ourthreechildren,John(theoldest),Alice,andChester(theyoung-est),allenjoyedgrowinguponthereservationandhelpedmeinvariousways.Johnwasaspecialhelpinpolicingtheareaduringhuntingseasons.Alice,evenfromthetimewhenshewasinjuniorhighschool,wasagreathelptomeinrecordingfielddataonthousandsofsnakesandotherani-mals.ChesterwasespeciallyhelpfulinobtainingglasslizardswhenIwasstudyingtheminthe1970s.heorganizedhisfriendstohelpwiththehunting,payingasmallfeeforeachlizardcaught,whilestillmakingabitofaprofitforhimself.JohnnowteachesenvironmentalcoursesatFloridaGulfCoastUniversity.helivesinnaples,Florida.Aliceisaresearchasso-ciateintheZoologyDepartmentatoklahomastateUniversity.sheandherhusbandtonyechellehaveworkedtogetherasaresearchteamforthepast30years.Chesterrunsarentalmanagementserviceinlawrence,andheandhiswifeDeannaliveaboutahalf-milefromthereservation. Inretrospect,mymainprofessionalaccomplishmentshaveinvolvedlong-termfieldstudiesthatusuallyentailedindividualmarkingofliveani-malsinnaturalpopulations(lizards,snakes,rodents)andcollectingdemo-graphicaldata.thesefieldstudieshaveextendedover64years:Sceloporus occidentalis atberkeley,California,andatmyformerhomeinJacksonCounty,southwesternoregon,mid-1930s;snakes(especiallyCrotalus viri-dis andPituophis catenifer)androdents(especiallyOtospermophilus beecheyi andDipodomys ordii), late1930sandearly1940s,at thesanJoaquinexperimentalRange,maderaCounty,California;snakes(18species)androdents(especiallyMicrotus ochrogaster andPeromyscus leucopus)attheUniversityofKansasnaturalhistoryReservation,1948–1999;lizardsof15speciesat14localitiesofcontrastinghabitatsonnorth–southandeast–westtransectsinCostaRica,1967–1973. myfieldstudies,asoutlinedabove,havedemonstratedthatmark-recaptureprocedures,extendingoverperiodsofyears,areusefulforunder-standingspecies’demographiesanddemonstratingthaterroneousecologicalimpressionsmayresultfromashort-termstudyoroneconfinedtoaspecificlocality.(JournalsofmyfieldworkareslatedtobedepositedattheKansasecologicalReservesofficewheretheywillbeaccessibletofutureworkers.)

my1940publicationon the“ordinoidesArtenkreis”ofWesternGartersnakesisnowlongforgotten,andtaxonomyhasundergonedrasticchanges,butIconsiderthatpapertobebyfarthemostscholarlyandimpor-tantofallmystudies.FromfieldworkovermuchofthewesternUnitedstatesandexaminationofallavailablemuseumspecimens,Iwasabletoshowthatthemorphologicalcharactersuponwhichclassificationwasbasedwerehighlyadaptiveandwerecloselylinkedwithbehavior,habits,andhabitat. economicconstraintshavepreventedmefromattendingAsIhmeet-ingsregularlyorfrequently,butIhaveattendedthemwheneverfeasibleoverthepast64years.Inrecentyears,IhaveenjoyedmakingjointcampingtripstosomeofthemeetingswithtonyandAlice.Attendingthemeetingshasalwaysbeenanexhilaratingexperienceforme,becauseAsIhmembershavealwaysbeenleadersinthefield.overtheyears,thecostofbeingaherpetologisthasescalated,andIstronglyrecommendthatAsIhdowhatitcantocounterthistrend,forexample,byholdingdowncostsofregistra-tionfeesandhousingattheannualmeetings.Itwouldbewellalwaystoholdmeetingswherecampingfacilitiesareavailable. Ihaveseengreatchangesinherpetologyandherpetologistsovertheyears.Inthe1920sand1930s,therewereonlyafewindividualswhowereactiveinherpresearchintheUnitedstates.thefieldwasdominatedbythosesuchasstejneger,barbour,Ruthven,andKlauber.studieswerealmostentirelyinsystematics.Informationonlifehistoriesandecologyaccumulatedmainlyintheformofnotesandwasanecdotal.Acommontypeofpublicationwasanannotatedcountylist.Inthe1930s,whenIfirstattendedAsIhmeetings,studieswereorientedtomorphologyandsystem-atics,almostexclusively.themajorchangehasbeenashiftofinteresttobehavior,demography,andecology.Fieldworkintheearly1900sconsistedmostlyofcollectinganimalsthatcouldbepreservedforlaterstudyinthelab.now,herpetologyisthrivingasneverbefore,andtheincreasedinterestisverygratifying.Aphenomenonofthepresentgenerationisthefemaleherpetologist;therearenowmorewomenthanmeninsomeherpetologyclasses,andahighproportionofresearchpublicationshavefemaleauthors.WhenIwasgrowingup,Ineversawawomanreacttothesightofaliveherpotherthanwithhorrorandrevulsion.ofcourse,therewereafewpio-neersevenintheearly1900slikeDorisCochranofthenationalmuseumandhelent.GaigeatthemuseumofZoology,Universityofmichigan. myfatherencouragedmyearlyinterestinreptilesandhelpedmebuildalarge,outdoorscreencageinwhichIkeptmanydifferentkindswhichcoexistedmoreorlessharmoniously.theseweremostlylocalspecies,butIobtainedthenameofatexasdealer,the“snakeKing,”andpurchasedavarietyofkindsfromhim,includingIndigosnake,berlandier’stortoise,Collaredlizard,andctenosaur.Also,ImadecontactwithaGermanherpetologist,Wernerschröder,andbymailsenthimseverallocalkindsinexchangeforeuropeanspecies.sincethen,herpetoculturehasbecomeaverypopularhobby.somespeciesarebestknownfromobservationsinzoosorprivatecollections.thisinterestiscommendable,butthereisdangerthatrareandendangeredspecieswillbeadverselyaffectedbyoverzealousandillegalcollecting. ForaslongasIcanremember,interestinreptilesandamphibianshasbeenadominantinfluenceinmylife,andotherinterestshaveseemedrelativelyminor.Findingandcapturingherpshasofteninvolvedstrenuousexercise,andIalwayslikedthat.Distancerunningwasoneofmyhabits.AttheUniversityoforegoninmysophomoreandjunioryears,Iwasonthecrosscountryteamandranthemileandtwo-mileintrack.Iespeciallyenjoyedtennis,sincewehadatenniscourtattheranchwhereIgrewup.Inmyhighschoolyears,Iroutinelyplayedtenniswithmyfatherandinlateryearsplayedwithmysister,Ruth.Also,Ienjoyedscrubsoccerandbasketballandwouldoftenassembleneighborhoodkids,allyoungerthanI,withmyyoungersiblings,forgameswiththreetosixorsevenparticipat-ing.In1923(or’24?)whentherewasalocaltennistournament,Iwonthejuniorchampionship(underage16)ofmedford.AsanadultinKansas,Icontinuedthegamingtradition,withintrafamilyteamsofmychildren,theirfriends,graduatestudents,andeventuallymygrandchildren.Inmorerecentyears,pingponghaspartlyreplacedthemorerigorousgames.

henry Fitch on the Fitch natural history Reservation holding a Red-sidedGartersnake(Thamnophis sirtalis parietalis);23may1991.

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Henry Fitch and the Practice of Natural History

harryW.Greene

DepartmentofecologyandevolutionarybiologyCornellUniversity,Ithaca,newYork14853([email protected])

texasAlligatorlizardswerefirstdescribedfromtheDevil’sRiverin1858andacenturylater,whenIencounteredtheminphotographs,

therewasstillalmostnothingknownaboutthosesnaky,bright-eyedrep-tiles.Asamissouriyouth,hungryforwilderplacesandimaginingmyselfatrailblazingnaturalist,Iporedoveraccountsinhobartsmith’sHandbook of Lizards oftworelatedWestCoastanguids—especiallyfieldstudiesbyhenryFitch,whoreferredtothemas“unusuallyintelligent”andsawasouthernAlligatorlizardholdoffYellow-billedmagpiesbyhissingandthreateningwithopenjaws,tailcurledforwardlikeashield.someday,Ithought,I’llroamthemexicanborderlandsandlearnsomethingexcitingaboutGerrhonotus infernalis! Fitch’sphotoalsocaughtmyattentionamongthe“influentialsau-rologists”profiledinsmith’sHandbook becauseheworeaWorldWarIcavalryhatandlookedintense,asifdistractedfromsomeimportanttask.hispublicationsinourlocalcollegelibraryprovidedaUniversityofKansasaddress,soIwroteannouncingmyupcomingherpetologicalcareerandask-ingquestionsaboutproposedtexasfieldwork.howeverpretentiousthatletter,rightbackcamehenry’scordial,handwrittenexplanationofhowtosexalligatorlizards:“bygraspingthebaseofthetail,gentlytwistingit,andexertingpressurewiththethumbventrally,onecancauseahemipenistobeexposed.Failinginseveralsuchattempts,onemaybereasonablysurethespecimenisafemale.” Iknewventralmeantundersideandpeneswereforcopulation,butcouldn’thaverealizedahighschoolinternshipwithhenrywouldsetmycourseorthathewouldauthoralmost200publications,morethan4,000pagesonplants,snails,spiders,anddiversevertebrates.thisunassumingmanstartedgraduateworkatberkeley’smuseumofVertebrateZoologyin1931,whenthediscoveriesofCharlesDarwinandAlfredRusselWallacewerestillrelativelyfresh,andtookhisfirstacademicpositionin1948,fiveyearsbeforeJamesWatsonandFrancisCrickunraveledDnA.Decadeslater,afterhenrysummarizedhalfacenturyoffieldworkatasympo-

siuminhishonor,agraduatestudentwrylynotedthatthankstohimshecouldn’tcallfouryearsofhornedlizardpopulationresearch“longterm.”Applausetypicallyoccursafter presentations,buthisarrivalatthepodiumthatdayprovokedastandingovationbeforehebeganspeaking. Withhenry’spassingIwanttohonorhisimpactonbiologyandmepersonallybyreflectingonaconundrum.AlthoughDarwin,Wallace,andcountlessothershavebeendrawnintonaturebyorchids,beetles,orwhateverseizedtheirfancy,andthisissurelytruernowthaneverbefore,acclaimtypicallycomesfromgeneralizingratherthangatheringfacts.ernstmayr,forexample,wasrenownedforevolutionarytheoryandlesswidely

Henry s. fitch (1909–2009): field notes on a Wonderful life

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1Originally published in Herpetological Review 40:393–400 (2009). Reprinted with permission.

henrysheldonFitchpassedawayon8september2009,justabitshyofhis100thbirthday(25December).Inlieuofatraditionalobituary,andbecauseanautobiographicalaccountofhisliferecentlyappeared(echelleandstewart2000),severalformerstudentsandcolleagueseachdescribetheirpersonalmemoriesofthemandeservedlyreferredtoasthe“fatherofsnakeecology.”Althougheachoftheremembrancesbringsadifferentperspective,threecommonthreadsrunthroughallofthem,namely:henryFitch’senergy,enthusiasm,anddecency.

henryFitchandRainbowtrout;ca.1920atKlamathlake,oregon.

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sofordescribingmorethan400newspeciesandsubspeciesofbirds.Atfirstglancethen,henry’scareer-longfocusonorganismsseemsanachronistic,thewidespreadesteeminwhichhe’sheldabitsurprising.Inaforthcomingbook,Tracks and Shadows: Field Biology as Art,I’vesetouttoilluminatethatstatureaswellasmoregenerallyassesstheenduringvaluesofnaturalhistory.hereI’lldrawoncorrespondenceandinterviewsofmyteenagementor,withthegoalofaddressingaquestion:whydidhedoit? oneofourextendeddialogswassoonafterhenryhadfallenandspentachillynightstrandedinacreek,andexceptfortheincident’snotori-etyheseemedsurprisinglyunfazed—searchdogswerehopelesslyconfusedbecausehisscenttrailwaseverywhere ontheReservation,andhe’dbeenconsciouswhenhelicopteredtoahospital.DaughterAlicewasvisitingherfolks,andwetalkedallafternoonandlateintoevening.Almost80,withravenblackhair,henry’swifeVirginiaservedfriedchicken,mashedpota-toeswithgravy,cornonthecob,andhome-maderolls,andshefairlyspar-kledasconversationmeanderedfromourfirstvisitandthewhereaboutsofformergraduatestudentstodetailsofFitchfamilylife.WhenIsmiledathermentionof“youthfulindiscretions,”Virginiasaidshemarriedyounganddivorcedtheotherguy.“then,”sheexclaimed,grinningatmeandhuggingherhusbandfrombehindhischair,“Imetthiswonderfulguy!” FromtimetotimeIcheckedalistofquestions,andalthoughincor-respondencehenryhadbeenenthusiasticaboutmybookproposal,hisanswersweren’teffusive.I’dknownthiswasn’tgoingtobeeasy,iffornootherreasonthanconstitutionalreticence—asRandyReisererwroteinadissertationacknowledgmentofhisundergraduateadvisor,“Inevermetanyonewhocansaysomuchwithsofewwords,orindeed,withoutanyatall.”butIwantedtounderstandwhyhenrydoesthework,keepscatchingstillmoresnakes,andwhatthepracticeofnaturalhistorymeanttohim,sofinallyIblurtedoutsomethingabouthavingmyownproblemsshrinkinthefaceofgrandeuranddiversity.“soundsgoodtome”wasallhesaid,withasoftchuckleandmaybeahintofirony. Ialsohopedtolearnhowhenryknewwhattorecord,givenhebegangatheringdatainthe1930sforwhichtherewerenoguidingtheories.hispaperstypicallysetforththeecologyoftargetspecies,withinsightswovenamongempirical findings—thethesisworkonalligator lizards,e.g.,addressedadvantagesofviviparitybynotingthat“eggsleftinthegroundareexposedto…egg-eatingreptiles,mammals,andinsects,andtoextremesoftemperatureanddangerofdesiccation,whilethosecarriedbythefemaleprobablystandabetterchanceofdevelopingintoindependentlysuccessfulyoung.”In1949he’dlaidoutinEcology detailsofwhat towritedown,butalmostnothingastowhy particularinformationwouldinterestotherbiolo-gists.Andin1966RobertmacArthurandericpianka’sbrilliantpaperonoptimalforagingwouldinspirewidespreadmeasurementofparametersthathenryhadbeenrecordingfordecadeswithnoconceptualprompting. soIkeptcomingatthequestionsfromvariousdirections,hopingVirginiaandAlicewouldjumpinwithsomethingdefinitiveornudgehimfordetails.myqueryaboutgodresultedinaslightpauseand“Ihavenoreligiousbeliefsalthoughraisedinthatenvironment.naturalhistorydoesitforme.”Askedaboutfavoritehabitats,henryattributedhispreferencefordeserts,“becausetheyareopenandhaveinterestinganimals,”toanevadafieldtripduringgraduateschool.Atthementionoffavoritespecies,heresponded“alligatorlizards,Copperheads,andgartersnakes,becausetheyhaveinterestingnaturalhistories.”byevening’sendthebestIcouldgetwas“myinitialinterestinzoologywasinnate”and“Iwrotedowneverythingthatinterestedme.” twoyearslaterIwasbacktrudgingupahillontheReservation.henrywasaudiblywindedaswecrestedthefamiliarlimestoneridgeandexplainedwithoutatraceofself-pitythathe’dloststaminabuthopedtocompleteonemorefieldseason.otherwiseheseemednodifferentthanmylastvisitandat91hishairwaslightbrown.hewalkedslightlystooped,inworkbootswithvisiblythinsoles,andwaswearingkhakipants,aberkeleyherpetologycourset-shirt,andabaseballcapdecoratedwithvariousuniver-sityinsignia.thetatteredcottonbagstuffedthroughhisbelt,custommade

byVirginia,wasforcarryingsnakesbacktothehouse.heusedasmooth,sturdystickwithanailheadprotrudingonthebottomtosteadyhimself,holdontotreelimbs,turnovercoveritems,andprobemattedgrassforthelongnarrowtinpieceshe’dlaidouttoattractsnakes. Asourconversationturnedtocurrentprojects,henryspokewithquietfatherlyprideofapaperwithAliceaboutchangesintreediversityoverthepast50yearsattheReservation.theirfindingswerethought-provoking:oncelargelyprairie,perhapsthebest-knownsquaremileinnorthAmericahadlostathirdofitsfaunasincehearrivedbecauseoffireprevention,lackofgrazing,andforestencroachment.onthebrightside,therewerestillbobcatsandtimberRattlesnakesinthevicinity,andablackbearwasseennearhererecently.Aswereturnedtothehousehepointedoutalargecedarbythedriveway,plantedmanyyearsagoasatinyfamilyChristmastree. thatnighthenrysatinthefrontrowformycampuslectureonorganismsasthecentralfocusofbiology,duringwhichIhelduphiscap-stoneopus,A Kansas Snake Community,andintroducedmyteenagehero.Ipraisedhenry’scontributionstoecologyandsystematics,thensaidhisgreatestlegaciesareimmediateproductsoftheworkitself—tensofthou-sandsofobservationsarchived,manymuseumspecimenscollected—andthatinthescholarlytraditionofhisberkeleyadvisor,JosephGrinnell,he’dbridgedDarwin’ssynthesiswithtwenty-firstcenturyscience.Asithap-pens,thoseindividualorganismshestudieddemonstratedsuchthingsassubstantialshiftsinCopperheaddietsoverthedecades,aspreypopulationsrespondedtothehabitatchangeshedocumented. henry’saccomplishmentsamountedtoseveralbetter-than-averagecareers,ItoldtheK.U.crowd,givenhisCaliforniaandlouisianawork,decadesattheReservation,andhistropicalexpeditions.Infact,althoughmainlyknownasaherpetologist,hispublicationsonmammalswouldeclipsethoseoftheaverage“mammalogist.”DuringthelectureIshowedphotosofislandCottonmouthsthateatfishregurgitatedbyseabirdsandhave exceptionally large young, exemplifying, Ipointedout,unusualandunexpectedopportunities forresearchandenlargementoftheory.Afterwardhenryaskedaboutthenumberofyounginislandsnakelitters,saidhe’denjoyedmytalkandourhike,thenaddedwithacharacteristicgrinandswingofthechin,“oh,andthanksfortheplug!” Workcanbeajob,acareer,orapassion,andforhenrytheprac-ticeofnaturalhistorywasallthree.WhenIcomplainedaboutfundingheresponded,“Ihavealwaysspentmytimeonwhateverinterestedme—withorwithoutgrants—andhavegreatlyenjoyedallmyprojects,especiallythefieldwork.”InhiseightieshewasquotedinabookonKansas

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henryandVirginiaFitchin1975attheUniversityofKansasnaturalhistoryReservation(nowtheFitchnaturalhistoryReservation).

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personalities,“Iwouldn’tchangeathing.peoplewhoworkwithanimalsinthefield,whethersnakesorbirdsorrodentsormonkeys,finditdeeplysatisfyingandwouldn’ttradeitforanyotherkindofcareer—eventhoughitmaynotbeveryfinanciallyrewarding.”Andin1995hewroteAlice,“IfasayoungpersonIcouldhavedreamedofmyfutureandtheworldIwouldliketosee,itwouldhavebeenaboutthesameasthelifeIhavehad.Gettingaph.D.,havingaloving,supportivewife,childrenlikeyouandJohnandChester,grandchildrenliketyson,lena,andben,livingontheReservation,teachingnaturalhistory,studyinganolesandpitvipers,andmakingtwodozentripstoninecountriesinthetropicsforherpetologicalresearchhaveallbeengreatexperiences.” onevisit,aftertheFitcheswalkedmetomytruck,IdrovebacktolawrenceonasultryKansasnight.AhugemoonshonethroughfogandorangelightningflashedoversurroundingfieldsasIponderedmyadmira-tionandaffectionforhenry.What,Iwondered,makeshimtick?Certainlyhemarchedtohisowndrummer,unmindfuloffads,whichmakesitallthemorefascinatingtocontemplatehisaccomplishments,aswellashowthatstanceaffectedhislifemorebroadly.maybeverbalfrugalityreflectedlimitedinterestinanalyticthought,personallyandprofessionally,whichifnothingelseprotectedhimfromthepettinesssocommoninuniversities.maybehewasalwayssomuchwithinhimselfthathesimplydidn’tpaymuchattentiontotheorizing.Andmaybethoselikehenrywhogodeeplyintonatureaschildren—hewascatchingsnakesasafive-year-old—areespeciallypronetoimmersionasadults.Wehavetobeoutthere. JustweeksbeforehenrydiedheaskedAliceandherhusbandtonyechelle,iftheymightvisitalocalcreekandcatchwatersnakes.Whenshereplied,“Well,whatthendad?”hesaidsimply,“We’llmarkandrecapturethem.”evidentlytheanswerstomyquestionsareequallystraightforward:henrywasalwaysdrivenbypassionatecuriosityandapenchantfordetail,accentuatedbyparentalencouragement,andthoseattributescombinedingradschoolwithaframeworkforunderstandingbiologicaldiversitythatharkedbacktoDarwinandWallace.thatwasenough.hisapproachworked,againstformidableoddsattimes,andhewasnotinclinedtodootherwise.AspecialgiftforfieldbiologyandquietbutstubbornconfidencemusthavebeenobvioustoGrinnellin1931,whenanunusuallyshybutpromisingnewstudentarrivedatberkeley,freshoffanoregonpearranch.thosetraitswereundiminishedtotheend,andhenry’slong,happylifewasinseparablefromthequesttounderstandnature.

Reminiscences of Henry S. FitchWilliame.Duellman

biodiversityInstitute,UniversityofKansaslawrence,Kansas66045([email protected])

ForthepasthalfcenturyIwasfortunatetohavebeenacolleagueandfriendofhenrys.Fitch.Duringthattime,weservedondoctoral

committeesofoneanother’sstudents,co-advisedsomegraduatestudents,andteam-taughtagraduatecourseinreptilebiology.Althoughwenevercollaboratedonapublishedpaper,weeachnamedaspeciesofAnolis fortheother.henryavoideduniversitypoliticsandonlyreluctantlyattendeddepartmentalmeetings.manyofhiscolleaguesmusedthathenrylivedinhisownlittleworld.butthatworldwasmuchlargerthantheythought.hewasathomeontheUniversityofKansasnaturalhistoryReservation(subsequentlynamedforhim),whereheconductedintensivestudiesontheecologyandbehaviorofreptiles.overtheyearshewitnessedthesuccessionofhardwoodforestonthereservation,whilemethodicallysearchingthissquaremileofland,capturingandrecapturingthousandsofsnakes,andlogginghundredsofpagesofnotes.hisdedicatedeffortsculminatedwiththepublicationin1999ofA Kansas Snake Community: Composition and Changes Over 50 Years.

however,henryhadalonganddistinguishedpublicationrecordbeforethisfinale.WhileattheUniversityofCaliforniaatberkeleypriortomovingtoKansasin1948,hepublishedaclassicworkonalligatorlizardsin1935andahighlyperceptiveworkonwesterngartersnakes(Thamnophis)in1940.twoofhisbest-knownworksareonthenaturalhistoryofreptiles,especiallythatontheFive-linedskinkin1954andhisexhaustivestudyoftheCopperheadin1960.Inadditiontothesesystematicandecologicalstudies,Fitchprovideduswithimportantsyntheses—reproductivecyclesinlizardsandsnakes(1970)andsexualsizedifferencesinreptiles(1981).Consequently,hispublicationsarecitedextensively.Anumberofyearsago,whileFitchwasstillanactivememberofthedepartment,thechairmantookituponhimselftotallycitationstopublicationsbyallmembersofthedepartment.Forseveralconsecutiveyears,henryFitchwasthemostcited. Duringthewarmermonthsoftheyear,themajorexceptiontocon-ductingfieldstudieswerethebasketballgamesonthe“sandlot”bytheFitch’sresidence.Gameswouldinvolveallmembersofthefamilyandany-onewhohappenedtobevisitingthereservation.henrydisplayedhisusualdoggeddeterminationfromhisfieldworktothebasketball“court,”andonequicklylearnedtoavoidhiselbowsunderthebasket. In1967IintroducedhenrytothetropicalrainforestinAmazonianecuador,wherehewastheonlymemberofthefieldpartywhowouldworkinthefieldduringthetorridafternoons,allthewhilelamentingtheappar-entabsenceofsnakes.Customarilyhewentintohiscabinandemergedafewminuteslaterwithatowelwrappedaroundhismiddleanduntiedsneakersonhisfeet.togettothedribblingbamboospoutlooselyreferredtoastheshower,hehadtocrossthedirt“courtyard,”thehometerritoryofaveryaggressivegoose,whichtookparticulardelightinnippingathenry’sbuttocks.oneafternoonweheardhenryexclaim“ouch,”ashestoodnakedsnappinghistowelatthegoose.hewascompletelyunawarethathewasthe“floorshow”inthemiddleofcamp. hereIlearnedthathenryhadpoornightvisionandconsequentlywasprimarilyadiurnalbiologist,buthewasconstantlyamazedthatwefoundsomanysnakesatnight.onlyaftermuchcajolingdidheaccompanyustwiceonnocturnalforaysduringamonthintheforest.however,IliketothinkthatIinfluencedmuchofhenry’ssubsequentworkinthetropics,whereheconductednumerousstudiesonthesystematicsandecologyofanolesandonpopulationsandconservationofiguanas. henryFitchwasoneofthelastremainingnaturalists.hisbreadthofknowledgewasmatchedbyveryfewofhiscontemporariesandscarcelyimaginedbyhisyoungercolleagues.hiscarefulworkonnaturalhistoryiswellworthemulating.ourknowledgeofanimalsinnaturewouldbefargreaterifmanymorebiologistsaroundtheworldfollowedinthefootstepsofhenrys.Fitch.

henry Fitch was modest and unassuming — but very competitive inbasketball.

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Henry Fitch at Home and in the Tropics

RobertW.henderson

sectionofVertebrateZoology,milwaukeepublicmuseummilwaukee,Wisconsin53233-1478([email protected])

Iwas,herpetologically speaking, incrediblygreenwhenIentered theUniversityofKansasasanundergraduatein1967.theattractionofKU

wasitslonghistoryofherpetologicalfieldworkintheneotropics,andIwasawareoftheambitiousresearchprogramofWilliamDuellmanandhisstu-dentsin,atthattime,ecuador.Iwasnot,however,awareoftheexistenceofhenryFitch.IdidknowthatIwantedtostudythewaysnakeslived,butIhadabsolutelynoideaofhowtodoitorthatanyonewasalreadydoingit.eventually,whilebrowsingthroughthelibraryintheherpetologydivision,IcameacrossAutecology of the Copperhead.Ireaditfromcovertocover,realizedthiswasthekindofresearchIwantedtodo,andIhadtomeetDr.Fitch,whojusthappenedtoteachatKU. theFitcheslivedonthegroundsofKU’snaturalhistoryReservationandIbecameafrequentvisitor,oftenmakingtheroundsofcoverboards(sheetsoftin)inQuarryFieldtocollecttheprairieRing-neckedsnakesonwhichDr.Fitchwasthenworking.IhavefondmemoriesofthewarmhospitalityofVirginiaFitch,energeticbasketballgamesplayedonthebaregroundattheReservation,theFitch’syellowVWbugwiththesmilinghappyfacestucktotheroof,andofmanyseeminglycarefreehourstramp-ingovertheReservationandencounteringosageCopperheads,easternYellow-belliedRacers,andprairieKingsnakes.DespitethetimewespenttogetherwhileIwasanundergrad,IsomehowconvincedDr.Fitchtotakemeonasagraduatestudent,andthusbegana30-yearcollaboration. Althoughmyprimaryinterestwasinsnakeecology,andDr.Fitch’slong-termsnakepopulationstudiescontinuedtobethefocusofhisKansasfieldwork,wenevercollaboratedonaresearchprojectwithanophidiancon-centration.ourtropicalfieldworkalwayshadlizardsasourprimaryobjec-tive.mexicowasthesceneforourfirstfieldtriptogether,primarilyinthestatesofChiapasandoaxaca,tocollectecologicalandmorphologicaldataonvariousspeciesofAnolis.thiswasfollowedbytravelsinnicaraguaandbelizeforworkontheconservationofIguana andCtenosaura,andthenmoremexicanfieldwork(whereonenighthesuggestedwebatheinthepacificatabout10pm;Iwasterrified,certainsomecreaturewouldlopoffmylegswhileItriedtogetclean).Asidefromthemanyhoursspentinatruckorjeep,onceatasitetherewaslittledowntime.onweekendsinnicaragua,wedidnothaveaccesstothejeepanddriverprovidedforiguanawork,butDr.Fitchknewthatanolesawaitedussomewhere,sooffwewentwithlocaltransportation,neverquiteknowingwherewewereheadedorifwe’dgetbacktoourquartersinmanagua.Itwasexhaustingfun. IbecameenamoredoftheWestIndiesinthelate1970s,andwaspleasedwhenDr.FitchjoinedmeintheDominicanRepublic(DR)foraprojectwithAnolis bahorucoensis in1985.IknowhehadagoodtimeintheDRbecausehekeptcommentingonthenumbersoflizardsencounteredeverywhereallthetime.hecameupwithonewonderfulideaafteranotheronhowbesttostudythiselusiveanole.helaterreturnedtotheDRtostudytheinvasiveAnolis cristatellus anditsimpactonnativeanolesinthelaRomanaarea.Cumulatively,wespentmonthstogetherdoingfieldworkinthetropics.Dr.Fitch’senergyandenthusiasmforlookingforlizardsandsnakesneverwanedand,althoughIwasmuchyoungerthanhe,itwasnotalwayseasytokeepupwithhim.hehadnoqualmsaboutsleepingincrummyhotelsand,despitehisdiabetes,hecouldbeprettymalleablewhenitcametofood(althoughweatespamandboiledpotatoesevery nightforsixweeksinmexico).IsenseDr.FitchneverlosttheenthusiasmIlastobservedmorethan20yearsago,andthat’sbeenborneoutbyhisincred-ibleproductivitywellintohis90s.Ashenotedinaninterviewconductedbyhisdaughterin2000(echelleandstewart,2000),“interestinreptiles

andamphibianshasbeenadominantinfluenceinmylife,andotherinter-estshaveseemedrelativelyminor.”IrecallhimtellingmeaboutoneofhisKUcolleagueswhowentoffonatwo-weekfishingtrip.heintimatedthathewouldnotbeabletostanddoingthat,andseemedgenuinelyperplexedthathiscolleaguecouldforegoresearchforthatlong. Dr.Fitchwastheidealcollaborator:healwaysmetdeadlines(andexpectedthesamefromme),andwasgenerousinsharingauthorship.I’msureImusthavetriedhispatiencemanytimes(whileastudent,oftenshow-ingupathishomeunannouncedtodiscusssome“important”issue),buthenevervisiblylostit.evenaminorfalling-outwasquicklyforgotten.hedid,however,atonetimeinformmethatIwasnoJohnlynchormartyCrump! Whilehewasalive,henryFitchwasaninspirationtomeandIcon-sidermyselfamazinglyfortunatetohavebeenoneofhisstudentsandcol-laborators.heremainsmyherpetologicalinspirationandI’mconfidentthathisincrediblebodyofworkwillcontinuetoinspireresearchersformanydecadestocome.WespokeinfrequentlyonthephoneduringhislastyearsontheReservation(andbeforemovinginwithhisdaughterandson-in-lawinstillwater,oklahoma),butoneofourlastconversationswasmyfavorite.Idon’trecalltheexactwording,butyou’llgettheidea.RWh:so,youmustbe,what,91or92now?hsF(chuckling):no,93.RWh:Well,how’reyoudoing?hsF:notsogreat.Ihaveabadhipandmybackispainful,sogettingaroundisdifficult.RWh:I’msorrytohearthat.nofieldwork,Iguess.hsF:Ah,wellIjustcamebackfromradio-trackingsixtimberRattlesnakes.RWh(shakinghisheadinaweandadmiration):Amazing. IalwayshopedheknewhowmuchhehadinfluencedmyeffortsandIwaslookingforwardtosendinghimacopyofaforthcomingbookon

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henryFitchandaboaConstrictor;February1972inChiapas,mexico.

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thenaturalhistoryofWestIndianamphibiansandreptiles.Idoubtthathewould’vebeenuptoreadingit,butIdidwanthimtoknowthatIhadwrittenit(withbobpowell).perhapsmyego(orinsecurity)hopedhe’datleastthink,“bob’sdoneoK.”

In the Field with Henry FitchDavidm.hillis

sectionofIntegrativebiology,UniversityoftexasAustin,texas78712([email protected])

IreceivedaphonecallfromAliceFitchechelleinthefallofmyjunioryearatbaylorUniversity.IknewAlice,henryFitch’sdaughter,throughher

husbandtonyechelle.tonywasteachingagraduatecourseinsystematicsatbaylor,andhadgivenmepermissiontoenrollinhisclassasanunder-graduate.thatcourseopenedmyeyestotheexcitingpossibilitiesofphylo-genetics.tonyandAliceoftenletmecomealongontheirmanyfieldtripstocollectfishesthroughouttexas,soAliceknewthatIwasenthusiasticaboutfieldwork. overthephone,Alicedescribedhowherfatherhadreceivedasmallgrant to study the impactofhumanexploitationof iguanas for food,andwasplanningasemester-longfieldtripthroughmexicoandCentralAmerica.Ashenrywasnearingretirement,Alicewasworriedabouthimmakingthetripalone,andsheknewthatIhadapassionforanythingtodowithherpetology.shewantedtoknowifIwasinterestedintakingoffasemesterfromschoolandaccompanyinghenryinthefield.“Iwouldlovetodoit,butI’llneedtothinkaboutit,”Isaid,asIconsideredhowdelay-ingmygraduationbyasemestermightaffectmyfutureplansforgraduateschool.Ihungupthephoneandthoughtforabout30seconds,thencalledAliceback.“I’lldoit…whendoweleave?”thatwastheextentoftheback-groundandtheplanningforwhatwastobemymosteducationalsemesterasanundergraduate. IwasalreadyquitefamiliarwithhenryFitchthroughhisbooksandresearcharticles.IhadmethimtheprevioussummerashevisitedAliceandtony,whenIaskedhimtosignmycopyofhisbook,Reproductive Cycles of Lizards and Snakes.Irememberthathewaspleasedtoseethatmycopyofthebookwaswornandobviouslyheavilyused.henryhadbeenembar-rassedandmodest,actingasifhefelthonoredtobeaskedbyanover-eagerundergraduateforhisautograph.Icouldn’tbelievethatsuchafamousper-soncouldbesomodest. henrydrovedownfromKansasandpickedmeupintexasinearlyJanuary,inanoldInternationalharvesterpickupwithawell-worncamperinthebed,whichwastobeourhomeforthenextfourmonths.Wecrossedintomexicothenextday,andIlearnedveryquicklythattherewaslittlejustificationforhenry’smodestdemeanor.Iaskedquestionsaboutevery-thingwesaw,andrarelydidImanagetostumphim.Ateverycampingsite,Icaughtpracticallyeveryherp,fish,insect,spider,crustacean,andmam-malthatIcouldfind,andhenrytoldmesomethingaboutthemall.hefrequentlyalsowouldexplainfundamentalconceptsofgeology,climate,ecology,andjustabouteveryotheraspectofnaturalhistoryasweslowlymovedsouththroughmexicointoCentralAmerica.hisimpromptulec-turesstimulatedmetowritelongentriesinmyfieldjournaleveryevening. thefundedpurposeofthetripwastostudyiguanaexploitation,andwedidthat.buthenrywasfascinatedwithanoles,anditbecamecleartomethathenrywasmuchmoreinterestedinstudyingthesystematics,ecology,andbehaviorofanolesthanhewasinstudyingiguanas.thatwasfinewithme;Ididn’tcarewhatwestudied,aslongasitwasrelatedtoher-petology.Wetraveledfairlyslowly,sothatwehadplentyoftimetocollectanolesandotherherpsateverycampsite. Ihadneverbeforemetanadultwhowassocompletelyimmersedinthestudyofnaturalhistory,sowehititoffimmediately.henrywasasenthusiasticaboutbeinginthefieldasIwas,atleastbyday.Iwassurprised,

however,thathenrydidnotaccompanymeonmynightlyforaysintotheforestsaroundourcampsitesafterdark.hisdiabeteshadaffectedhisnightvision,andsohedidnotseewellatnight.buthewasalwaysenthusiasticaboutalltheherpsIwouldfindandbringbacktohiminthecamperafterdark,andwewouldoftenstayuplatetalkingaboutourlatestcapturesandwhatinterestingresearchproblemstheymightsuggest.tohenry,everyspeciesprovidednewquestionsthatwerewaitingtobeanswered.Icouldn’timagineamoreexcitinglife. AsmuchasIwasenjoyingmyself,therewereafewaspectsofhenry’sapproachtofieldworkthatcausedmesomeaggravation.First,heinsistedondoingallthedriving,eventhoughI’dbeenaskedbyAlicetoaccompanyhenryonthebasisthathewouldneedsomehelp.Givenhispoornightvision,hisnight-drivingprovedquitefrighteningtoeveryoneontheroadexcepthenry,andmorethanonceIwasconvincedwewoulddriveoffoneofthemanyprecipitousroadsidesinthemountainsofCentralAmerica.eventually,Iinsistedthathehadtoletmedriveatnight,orelseIwouldonlygetinthetruckwithhimbyday.Wedroveonlybydayfromthenon;IwasstillnotpermittedtodrivetheoldInternationalharvesterpickup. Asecondminoraggravationwaswhatcangenerouslybedescribedashenry’sratherparsimoniousnature.Weboughtandcookedallourownfood,whichwasfinewithme,exceptthathenrywasastraightmidwesternmeat-and-potatoesman,andwealmostneverhadanymeat.soweatemostlypotatoes.Afterthefirstfewweeksofasteadypotatodiet,Iwasget-tingdesperateforsomethingIcouldtaste.Istartedbuyingafewpeppersandspicesfromthemarketsonthesly,andcatchinglandcrabs,freshwaterprawns,andfishfromthestreamsaroundourcampsites.soonIwascook-inguptwomeals:boiledpotatoesforhenry,andaspicyconcoctionofwhateverIcouldcatcharoundourcampsitesforme.henrypolitelytriedmyattemptsatculinarydiversityafewtimes,buthealwaysreturnedtopotatoesfordinner.sometimes,forexcitement,hewouldputalittlesaltandpepperorbutteronthem,orevenaddascrambledeggwhenhewasfeelingextravagant. otherthangasoline(whichcostabout30centsagalloninmexicoatthetime),Ithinkhenrywasspendinglessthanadollaradayforallourotherlivingexpenses.sowhenwereachedtheGuatemala–elsalvadorbor-deronasaturdayinFebruary,andhenrydiscoveredthattheweekendfeeforcrossingtheborderwas50cents/person,butthattheweekdayratewas25cents,weofcoursehadtoturnaroundandcampoutinGuatemalauntilmondaymorningtosave50cents.AtourGuatemalancampsiteneartheborder,Ifoundsynbranchideelsinasmallstream,andIhadablastfigur-ingouthowtocatchthem(andIstillhavefondmemoriesofSynbranchus stew).bysundayIhadtohangmyclothesouttodry,andsomelocalswhohadbeendrinkingdecidedtotakeadvantageofthesituation.theygrabbedmyclothesandran,andIbrieflygavechase,untiloneofthemturnedandpulledoutamachete.Ireturnedtocampandsuggestedtohenrythatwemightwanttofindanewcampingspot.henrydidnotseemveryinterestedindoingsountilrocksstartedrainingdownonusfromthecliffaboveourcampsite.Wepulledoutjustasalargerocksmashedourwindshield.WhenIlastsawthatpickupmanyyearslater,thatbrokenwindshieldhadneverbeenrepaired. onmondaymorningafterthewindshield-smashingevent,wearrivedattheborderat8Am,justwhentheweekendratewassupposedtochangebacktotheweekdayrate.butafterthecustomarysprayingofthetruckwithDDt(aritualthatwaspracticedonbothsidesofeveryCentralAmericanborder),theborderofficialchargedus50centsaperson.henryobjected,andpointedtotheclockonthewall,whichsaid8:05Am.theborderofficialcalmlywenttotheclock,changedthetimeto7:55,andchargedustheweekendrate.ItwasoneofthefewtimesinfourmonthsIsawhenrymadaboutanything.Forthemostpart,hewouldrollwithwhateverlifedishedout,andmakethebestofeverysituation. ourstudyofhumanexploitationofiguanasandctenosaursreachedapeakduringlent,whenthemarketsofCentralAmericabecamewellstockedwiththeselargelizards.lizardisnotconsidered“meat”bymanypeoplewhogiveupmeatforlent,soiguanasandctenosaursweresoldand

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eatenbythethousandsduringthistime.butafterlent,wewerebackintothemountains,lookingformorespeciesofanoles.bythen,wehadhatchedaprojecttostudyintra-andinterspecificvariationofanoledewlaps,andwerebusycollectingseriesofasmanyspeciesaswecouldlocate. onemightthinkthatitwouldhavebeenaphysicalmismatchtopairayoungstrappingundergraduatewithaprofessornearingretirementonanextendedfieldtrip.Itwas…Ihadahardtimekeepingupwithhenry.hewasn’tfast,butheneverstopped.hecouldwalkalldayupsteepmountain-sidesinpursuitofthenextspeciesofanole,andwedidsojustabouteveryday.someyearslater,asagraduatestudentattheUniversityofKansas,IwouldtakepubliceducationclassesofschoolchildrenouttotheKUnaturalhistoryReservationwherehenryandhiswifeVirginialived.evenwellintohis“retirement”years,henrytookgreatprideinleavingkidsinthedust.AshecruisedthroughtheReservation,hewouldgrabsnakesashewalked,checkingtheirventralclipsforindividualidentification.“Aha!It’sfemalenumber1675!Ihavecapturedthisracermorethan300timesinherlife…Iamgladtoseethatsheisstillalive.”thekidswerealwayswide-eyedwithamazement,astheystoodtheretryingtocatchtheirbreathbeforehenryrushedofftothenextsnake. Asitturnedout,Aliceneednothaveworriedabouthenry’s“advancingage”whenwesetoffforourCentralAmericanfieldtrip.henryremainedactiveinthefieldforanotherthreedecadesafterthattrip…afullcareerformanypeople.henrycollectedmoredataafterhe“retired”thanmostpeoplecollectintheirlifetimes.Idoubtanyonewilleverbeabletorepeatthekindoflong-termautecologicalstudiesthatheperfected.butmorethanthespecificknowledgethatIlearnedfromhenry,whatIadmiredmostabouthimwashischild-likeenthusiasmfornatureandeverythinginit,andhisunquench-ableneedtoaskquestionsabouteverythingheencountered.severalgen-

erationsofherpetologistsbenefitedfromhiswisdom,hiskindness,andhispassion.mentionhisnametoanyonewhoevermethim,andyouwillgetasmileandastory.Ican’tthinkofabetterlegacyforagreatnaturalistwhosqueezedsomuchoutofsuchalongandproductivelife.henry,wewillallmissyou,butwewillsmileeverytimeyournameismentioned.

Natural History Observations of Henry Fitch

Raymondb.huey

Departmentofbiology(box351800),UniversityofWashingtonseattle,Washington98195-1800([email protected])

thesecondWorldCongressofherpetologywasheldinAdelaide,Australiain1994.onesymposiumwasonthebiologyofsnakes,and

henryFitchwasapresenter.Formanyintheaudience,thiswastheirfirsttimetohearthelegendinperson.soashenrywalkedtothepodium,theaudience’sinitialmood(oratleastmine)wasoneofexcitement,anticipa-tion,andrespect. henrygaveanamazingtalk.thethemewaslong-term(reallylong-term!)demographic trendsofallof thesnakesontheKansasnaturalhistoryReserve.theresultswerestunningbutdepressing:thedensityofessentiallyeveryspecieshaddeclinedovertime. theaudience(largelysnakebuffsofthefirstorder)quicklybecamesomber.Addingtothepoignancyofthemoment,weallrecognizedthatthisgrimresultwasbeingdeliveredbyakindandgentlemanwhosedeeploveforsnakesandtheirnaturalhistoryhadinspiredhimtocarryoutalife-longstudyofthesesnakes.buttheclearconclusionemergingfromallofhisimmenseworkwasthathisbelovedsubjectsweredecliningtoextinction.sometimes,lifedoesn’tseemfair. eitherinhistalk,orinthequestionperiodafterwards,henrynotedthataprimarycausewasapolicyofsuppressingfiresontheReserve:asaresult,successionwasunchecked,suchthathabitatsonthereservewereincreasinglyforestedandincreasinglyunfavorabletosnakes. someoneasked,“Whydon’tyoulightafire?”henrythoughtforamoment,andthenrepliedquietly,“Ican’tdothat.”mydistinctimpressionatthetimewasthathereallydidwanttodojustthat,butthathecouldn’tdosoethically,givenhispositionattheReserve. butIalsorememberdistinctlyfeelingatthetimethathenrywassend-ingasubliminalmessagetousintheaudience:“Ifyouwanttostartafire...” Forme,henry’stalkwascertainlyamongthemostmemorableonesoftheentireCongress.moreover,itscentrallessonstillhauntsmythoughts.Weshoulddosciencebecausewelovetheprocess,notbecauseweneedtolovetheresults.henryFitchcouldnothavelovedtheresultsofhiswork,butthere’snodoubthelovedtheprocess.

Memories of Henry FitchmichaelV.plummer

Departmentofbiology,hardingUniversitysearcy,Arkansas72149([email protected])

henryFitchwasoneofthemostgracious,kind,andgentlemenIhaveeverknown.Igreatlyrespectedhimforhislonglistofprofessional

achievements,butIalsorespectedhimforthemanhewasandhowhetreatedothers.hewasalwaysthegentleencouragertomeandIoftenthinkofhisexamplewhenIgetinanexasperatingsituationwithmystudentsorcolleagues.Ineverheardadegradingwordspokenagainsthenrytheman

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henryFitchexamininganiguanainamarketinelsalvador,February1979.

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byanyone.AnyandallwhohappenedtocomebythenaturalhistoryReservationwereheartilywelcomedbyhenryandhislovelywifeVirginia,andthatseemedtohappenfrequently. myformalassociationwithhenrybeganinJuly1972whenIcametoKUtobeginaph.D.program.IfounditdifficulttobelieveIhadtheoppor-tunitytoworkwithsuchaprofessionalgiant.Irememberstandinginaweaswespokeaboutpotentialresearchprojectsandthinkingthatsincehenrywas64atthetime,Ijustmightbehislaststudentbeforeheretired(wrong!Ionlymissedbyabout30years).henrywasthemanasfarassnakeecologywasconcernedandIassumedmydissertationwouldbecomeyetanothermono-graphofaReservationsnakespecies.butthenhesuggestedwegodownonthe“Kaw”(KansasRiver)tolookforsoftshellturtles.Iwasfascinatedbythesandbars,thesoftshells,andhenry’snonstopfountainofknowledgeaboutthem–talkaboutinformationoverload!butwhatIremembermostaboutourinauguralsoftshelltripwasthis27-yr-oldkidintheprimeoflifetryingdesperatelytokeepupwithanaging64-yr-oldwalking(morelikerunning)onthesoftsandashetalked.Ilookedforared“s”onhischest. henryoncehadaguestattheReservation,agentlemanfromthebombaynaturalhistorysociety,andbecauseIndiaisnearthecenterofsoftshelldiversity,henryaskedmetotakethegentlemanoutontheKawandshowhimourAmericansoftshells.bythattime,Ihadcapturedhun-dredsofApalone mutica andIassuredhenrythatitwouldbenoproblemseeingnumeroussoftshells.Anyfieldbiologistcouldprobablyguesswhathappened.Afterworkinghardforacoupleofhours,wesawmaybetwoorthreejuvenilesoftshells.Ilearnedthatdaythatoneshouldnevermakesuchrashstatementsregardlessofhowconfidentheisaboutseeinganimalsinthefield.henrywasapologetictothegentlemanandIwasembarrassed,buthenryneversaidanotherwordaboutittome. henryfrequentlyencouragedhisstudentstodo“interesting”sideprojectsalongwiththeirthesisordissertationwork.hesuggestedaprojectonsoftshellglandsinthesummerof1972thatwouldfitin“nicely”withmyecologicalph.D.work.beingaswampedandoverwhelmednewgradstudent,Iwasn’tterriblyinterestedinhissuggestionatthetime,butIneverforgotit(probablybecauseofwhosuggestedit).Wellhenry,youwouldbepleasedtoknowthatIfinallydiddotheprojectanditwaspublishedin2009justbeforeyouleftus.sorryI’msoslow;itonlytook37years.thanksforthetreasuredmemories.

Henry Fitch as a Mentor and TeacherRichardA.seigel

Departmentofbiologicalsciences,towsonUniversitytowson,maryland21252([email protected])

mostherpetologiststodayknowhenrys.Fitchonlyasanameonhisclassicpapersandmonographs(e.g.,Autecology of the Copperhead,

A Kansas Snake Community).Inthisremembrance,IwouldliketogivemyperspectiveonhenryintherolesIknewhimbest,asamentorandteacher. In1979,allIknewabouthenryFitchwashisoutstandingpublica-tionrecord.Ihadbeenacceptedtotheph.D.programattheUniversityofKansasforthefall1979semester,and,throughaseriesofletters,henryhadtentativelyagreedtoacceptmeashisdoctoralstudent.however,hehadcautionedmethathewasretiringin1980andthatImightwanttoreconsidercomingtoKUashisfinalstudent.thus,inJune1979,mywifenadiaandIdrovetoKansastomeethenryandhiswifeVirginiaforthefirsttime.littledidIknowthatthisinitialmeetingwouldleadtoa30yearrelationshipwithhenryandVirginiaandthatmyappreciationandrespectforthemwouldgofarbeyondanythingImighthaveexpected. Aswedroveontheunpavedentranceroadleadingtohenry’shouseontheKUnaturalhistoryReservation(nowtheFitchnaturalhistoryReservation),Isawnumerousmetalcoverboards(“shelters”inhenry’ster-

minology)anddriftfencesscatteredatwhatappearedtoberandomintervalsalongtheroad.Assoonaswereachedthesmall,incrediblymodesthousewheretheFitcheslivedontheReservation,henryandVirginiawereoutthefrontdoortogreetus.Almostimmediately,Igainedaninsightintohenry’scharacter:Knowingwhattocallyourpresumptivemajorprofessorisalwaysadelicatepropositionforanewstudent(“Dr.Fitch?”“professorFitch?”),buthenrydealtwiththatbyholdingouthishandandintroducinghimselfas“henryFitch,”andwewereonafirstnamebasisfromthenon.Virginiawentevenfurtherandgavenadiaandmeahugehug,invitingusinforlunch. Feelingmuchmoreatease,weonlygottotheFitch’sfrontporchwhenIsawthattherewerealargeseriesofjars,cans,andsnakebags,allholdingvariousliveherps.Askinghenrywherethesecamefrom,heproceededtotellusthatwastoday’scatchandthentellmuchmoreabouttheecologyandnaturalhistoryofhis“finds”thanfouryearsoffieldworkandreadinghadprovidedmesofar.Iwasstruckespeciallybythedetailednoteshenrytookoneachfindandhowmuchdatahewasextractingfromeachindividual. Afterlunch,henrysuggestedthatwedo“around”ofhistrapsandshelters,andthetwoofussetoffupthehillsidebythehouse.Withinmin-utesIdiscoveredthatthis69-year-oldmanwasinbettershapethanmostgradstudents,ashewentupthehillonwhatfeltlikeatrot.Aswewent,hebegantotellmeadetailedhistoryofwhatfeltlikeeverytreeandcritterwesaw.phrasessuchas“IamhearingaYellow-billedCuckoo”werethrownoutcasually,leadingtotwothoughtsIdarednotexpress:“Ithoughtthiswasaherpetologist”and“IamgladYoUarehearingthis,sinceIsurelydon’t!” IwasthrilledwhenwecametothefirstseriesofsheltersatwhatwasknownasQuarryField,sincethepacefinallysloweddownandhenrysaidthiswasthebestplacetoseeCopperheads.sureenough,thereweretwogor-geousCopperheadscurledupunderthefirstshelterweflippedandtheywererightinfrontofhenry.problemwas,therewerealsofourRingnecksnakesrightinfrontofme,and,whileIhadeyesonlyfortheCopperheads(anddid

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henryFitchusedfunneltrapsanddriftfencestocapturemanyofthesnakesonthenaturalhistoryReservation.

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notseetheRingnecks),henrydidexactlytheopposite.thus,IwentfortheCopperheadsinfrontofhenrywithmyhookandhenrybentdowntogettheRingnecksinfrontofme(thinking,Iamsure,“whoisthisdummywhodoesnotseesnakesrightinfrontofhim?”).Aswebouncedoffofeachother,henrystumbledforward,puttinghisfootrightinfrontoftheCopperheads,oneofwhichimmediatelystruckhisboot!Icouldseetheheadlinenow:“Famedherpetologistkilledbyvenomoussnake;newgradstudenttoblame.” Fortunately,thesnakemanagedonlytoclipthefrontofhenry’sboot,andwequicklycapturedbothCopperheadsandatleastsomeoftheRingnecks.Afterflippingafewmoreshelters,wehadatotaloffourCopperheads,threeofwhichweremarkedindividuals,oneaslongasnineyearsago.Ifoundthisnothingshortofamazing,havingonlyrecentlyreadthathighrecapturerateswereimpossibleforsnakes,andsaidasmuchtohenry.heshookhisheadandsaid:“IfIhadjustafewmorefencesandmoreshelters,I’dhavea100%recapturerate.”Iglancedathenry,tryingtodecideifhewassayingthisinjestorifhewasanglingforacompliment.Iquicklyrealizedthatneitherofthesewastrue;hegenuinelyfeltthathehadsimplynotworkedhardenoughandneededtodomoretosatisfyhisownstandards.Whatagreatexampleforanewph.D.student! Afewyearslater,Igottoseeasecondexampleofhenry’sgenuinehumilityregardinghisfieldworkwhenthenotedlizardecologistlaurieVittvisitedtheReservation.Knowingthatlauriewasespeciallyinterestedinliz-ards,henrymadesuretochecktheshelterswhereheknewslenderGlasslizardscouldbefound,andwequicklygotseveralofthem.Whenlauriesaidsomethingtotheeffectof“areyougoingtopublishanythingonthese?”henryindicatedthatthesamplesizewasstilltoosmallforasolidpublica-tion.laurieasked:“howmanyofthemhaveyoufound?”thinking(Iamsure)thattheanswerwouldbeahundredorso.henry’sresponseflooredus:“About1,500sofar,”hesaidrathercasually.Whenwetriedtoconvincehenrythat1,500glasslizardswasnothingshortofphenomenalandfarmorethananyoneelsehad,hejustshookhisheadandsaidheneededmoredata!bytheway,henrydidpublishamonographontheselizardsin1989;thesamplesizewas2,216individualscaptured3,353times!(Fitch1989). Inadditiontodoingfieldresearchwithhenry,IwasalsoluckyenoughtobehistAforthelasttwocourseshetaughtatKU,VertebratenaturalhistoryandAnimalsofKansas.bothcourseswerecombinedlecture/fieldtripformatsandmymainrolewasdrivingstudentstoandfromthefieldsitesandhelpinginthefieldanywaythatIcould.Althoughhenry’slecturesweredetailedandcomprehensive,thefunpartofbothclasseswasthefieldtrips.studentsenjoyedtryingto“challenge”henrybybringinghimwhateveroddinsect,snakeskin,ormammaldroppingtheyfound,thenhavinghimactasalivingversionofWikipediaandproceedtolecturethemonevery-thingthatwasknownaboutthespeciesunderquestion.oneofmyfavoritememorieswaswhenastudentfoundanewlyhatchedFive-linedskinkandaskedhenryhowmuchthetinylizardweighed.henryheldthelizardforafewsecondsandthenreplied:“1.15grams.”theclassimmediatelyburstoutlaughingattheabsurdlypreciseanswer.so,thewholeclasswalkedovertotheold,dilapidatedbuildingthathenrycalledhis“lab”andweproceededtoweightheskinkonanoldtriple-beambalance.sureenough:1.15grams!somewhatawed,oneofthestudentsasked,“howcouldyoupossiblyknowthat?”henry’sresponsewastypicallylow-key:“Whenyouhaveprocessedover5,000ofsomething,youknowtheirweightsprettywell!” Givenhenry’slow-keyapproachandhumility,itwouldbeeasytoconcludethathewasnotcompetitiveandthathewouldlethisgradstu-dentsgetawaywiththings.neitherconclusionwouldbetrue.IrecallquitewellthefirsttimehenrycametomystudysiteinnorthwesternmissouriwhereIwasdoingmark-recapturestudiesonsnakes.Aftercatchingourfirstsnakeoftheday,Imarkeditusingscale-clipping(nopIttagsinthosedays)andproudlyshowedittohenry.“oh,”hesaid,“Iguessyoudon’twanttobeabletorecognizethissnakeifyoucaptureitagain?”thiswashenry’swayoftellingmeIwasnotmarkingproperly,andthemessagewasdeliveredloudandclear.tothisdaywhenIscaleclipsnakes,Ifollowhenry’smethodsandcantellyou,theyworkextremelywell.

henry’scompetitivenaturemaybeillustratedbysomethingonlyaselectfewgottoexperience,somethingcalled“Fitchianbasketball.”henryalwayshadagrass/dirtbasketball“court”setupoutsidehisfrontdoorandduringthespringof1980,hiscurrentstudentsgotatasteofhowcompetitivehenryFitchcouldbeattimes.therewereeightofusthatday,threewomen(nancyZushlag[henry’smaster’sstudent],mywifenadia,andJimKnight’swife,Karin),andfivemen(henry,myself,andthreeofhenry’sotherstu-dents:larryhunt,luismalaret,andJimKnight).Whenthetimecametoplay,henrystartedexplainingthe“rules”;first,thereweretwohoops,oneat10feet,theotherat8feet.the10-foothoopwasfortheguys,the8-foothoopforthewomen.next,therewereoddbutveryspecificrulesaboutthemenandwomentakingtheballoutseparately,whetherthewomencouldbeguarded,andhowthepointsweretallied.Whatweallfoundmostamus-ingwashownewrulessuddenlyappearedwheneversomeonescoredagainsthenry’steam.mypersonalfavoritewas“nojumpshotsfromthecorner,”whichjusthappenedtobemybestplacetoshootfrom.needlesstosay,weallspentmoretimelaughingthanwespentplaying,aswatchinghenrymorphintothiscompetitivejockwassomethingnoneofushadseenbefore. basketballfinished,wethengottoseeafinedemonstrationofhenry’scharacter.henrywasduetoretirethatyearanditfelltoourgroupofgrad-uatestudentstofindanappropriatewaytocelebratehenry’smanyachieve-ments.WiththehelpofJosephCollins,billDuellman,andmanyothers,weorganizedasymposiumatthe1980herpmeetingsinmilwaukee,withmanyofhenry’sformerstudentspresentingpapers.Wealsoplannedtopublishavolumebasedonthatsymposium,whichappearedin1984.Afterlunch,wesprangallthisonhenry,includingaplaquemadeespeciallyfortheoccasion.henrywasdeeplymoved(Virginiawasintears),butIcouldtellthatwhilehewasclearlytouched,partofhimwassayingtohimself:“Ineedtogetoutthereandchecktheshelters…” nodiscussionofhenrycouldpossiblybecompletewithoutmention-ingVirginia.herprideinhenry’saccomplishmentsandherirreplaceableroleinhislifecannotbeoverestimated.Watchingherbeamfromeartoearduringoursymposiumhonoringhenryin1980wasajoytowatch.Forme(andnadia),Virginiawasmorelikeagrandmotherthanthewifeofmymajorprofessor.Fromgivingusfurniturewhenwewerestarvinggradstu-dentstogivingoursonhisfirsttricycle,herwarmthanddevotiontohenryandhisstudentswastrulyremarkable. AsIthinkofhenrynow,Iseehimheadingofftodoanother“round”attheReservation.mayhealwayshaveasmanyshelterstocheckashecouldeverwant,andmayhistrapsalwayshavemanymarkedsnakes.Whenhismorningroundsareover,he’llbeheadedbackhome,whereVirginiahaslunchwaiting.

Henry Fitch: The Twilight of an Incredible Career

GeorgeR.pisani

Kansasbiologicalsurvey,UniversityofKansas2101ConstantAvenue,lawrence,Kansas66047([email protected])

IfirstmethenryFitchin1968,forty-oneyearsago,whilevisitingafriendinlawrence.I’dreadmanyofhispapersinthecourseofmystudies,

andwhenImethimagainin1970,Iwasagainimpressedbytwothingsbeyondhisvaststoreofknowledge.Atagefifty-ninehecould,whilemak-ingaroundinthefield,walkthelegsoffmanypeoplefaryounger.And,forsomeonewhosemanypapershadessentiallyestablishedthefieldofsnakeecologyaswenowknowit,hewasincrediblyunassumingandreserved(exceptwhenplayingthe,ummm…rules-modified,basketballgamesthatthenwereaReservationfeatureevent). manyherpetologistsaccompaniedthisremarkablemanintothefieldinthecourseofhislong,distinguishedcareer.Itwasmydistinctprivilege

RemembRAnCes

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tobeamongthematvarioustimes,andtoworkwithhimduringthefinalfewyearsofthatcareer.ourcloseprofessionalrelationshipreallybeganinthemid-1980swithourcollaborationontimberRattlesnaketelemetry.Ibuiltaneffective(albeitawkwardlymaneuverable)antenna.Usingthatandsometransmitters(immensebycurrentstandards!)assembledbyDr.tonyshirer,andwhichhenrypushedgentlydownthethroatsoflargerattlers,wespentasummergatheringmovementdatathatretrospectivelyturnedouttobemorealearningexperienceaboutthetechniquethanaboutthesnakes.henryassimilatedallthis,andnearly20yearslater,whenwehadfarbetterequipmentandfarmorebackgroundknowledgeonthesubtletiesofusingit,wewereabletoinitiateanongoingstudyofthisspeciesinnortheastKansas. In1987henryandIreceivedfundingfromoklahomaDept.ofnaturalResourcestostudyrattlesnakeroundupsandtheireffectsonthepopulationsofWesternDiamondbackRattlesnakes.Itwasnosecretthathenry’slong-termenthusiasmforrattlesnakestudywasn’tsharedbyhiswifeVirginia,perhapsoutofrecognitionthathisreflexeswereslowing,asdoeveryone’swithtime,hamperingtheavoidanceresponseessentialforsuchwork.It’sonethingtododgethefeintofa2-footCopperhead,butquiteanothertododgethelongstrikeofa5-footCrotalus.Virginia’sreactionwassubduedapprehension.butaftersomediscussionweagreedtohandlethecaptivesnakesaftercoolingthem,areassuringmodelthatVirginiaaccepted,andwhichprovedeffectiveforsafelygatheringthemor-phologicaldatawesought. henry’senthusiasmforfieldworkandforlearningalltherewastolearnaboutsnakeecologyneverdiminished,thoughby2002,timeplainlywashavingitsinexorableeffectoflimitingboneandmuscle.Intheselateryearsofhiscareer,Ihadretiredandsowasabletohelphimmakeroundsinthefieldusingoneofthefieldstation’s“Gator”AtVs.Idrovewhilehenry’sgazewasfixedfirmlyonourdestinationoftheday,whereverithappenedtobe.by2003,hewasgamelytryingtomaneuverusingcrutchesonalocalledgethatservedasatimberRattlesnakeden.IemphasizedtohimthatIdidn’tplantoberememberedastheguywhobroughthimbackfromthefieldwithasevereenvenomationorabrokenleg,andsoheagreed(perhapsacquiescedisamoreaccurateterm!)withachangeofmodel—I’ddotheroughandtumblestuffandhe’dwaitinthe6-wheelertoshareintheresults,andwe’dcollaborateonpapersthat,ofcourse,requiredhisincrediblestoreofknowledge.or,iftopographyallowed,he’dhuntclosetothe‘GatorwhileIventuredfurther.thatmodelworkedwellthroughtheremainderofthetimehewasabletoliveindependentlyonFnhR,andotherpersons,youngerthaneitherofus,pitchedintohelphenrymakeproductiveroundsinthefieldsafely.notableamongthesearescottsharp(ahighschoolteacherinanearbydistrict)andhisfamily,andKUunder-gradsmikeZerwekhandJoeybrown.

Aslateas2006,the“henryandGeorgeteam”(whichIoncepointedouttohimhadanaverageageof79,arealizationhegreatlyenjoyed)stillmadejointrounds,theseinmysmoothearthsnakestudyareanotfarfromFnhR.thiswasaspecieshe’dbarelyseen,withjust3FnhRrecords.hewasfascinatedbythefactthatthespecieswassoclose,yethe’dseensofew.WhenonourfirsttripintothatareaIcaughtthefirstofseveralwesubsequentlyfoundintallgrasshabitat,hisreactionwastolookatitintentlyandsoftlyremark,“Well,I’llbedamned.”IthinkitwastheonlytimeIheardhimsaythat!I’mgladIsenthimthefinaldraftofthearticlesummarizingthatresearch.WhenIemailedittohisdaughterAlice,hisprimarycaregiverbythen,Iaskedherto“tellhenryhehastostickaroundtoseethisinprint.”plannedpublicationwasforDecember2009.Alicereadittohimandafterwardtoldmehowattentivehe’dbeenandhowhe’denjoyedlearningaboutthiselusivespecies. on8september2009,justafewmonthsshyofhiscentennialbirthday,henrysheldonFitchpassedaway,leavingforscienceoneofthemostout-standinglegaciesofecologicalstudyeverknown.bestknownforhisextensivelong-termstudyoftheherpetofaunaofwhatin1948wastheUniversityofKansasnaturalhistoryReservation(renamedin1986theFitchnaturalhistoryReservation[FnhR]),hispublishedstudies inanimalecologyextendingbackto1933alsoincludeawiderangeofbirds,mammals,andinvertebrates,aswellasthelocalsuccessionalfloraoftheirhabitats.harryGreene,interviewedbytheLawrence Journal-World,accuratelyreflectedonhenry’slegacybystating,“It’snotanexaggerationtosaythathenry’sthefatherofsnakebiology.”hisstudiesontheecologyandrelationsofthesemanyspecieswererefinedthroughhisextensivecareertoreflecthisuniqueinsightsregardingthewaytheyformcommunitiesofinteractingorganisms.Allofuswhocontinuetobuildonthislegacy,andthosewhofollowusandwilldothesame,oweatremendousthankstothismodestmanofgreattalent.

Literature Citedechelle,A.F.andm.m.stewart.2000.historicalperspective:henrys.Fitch.

Copeia2000:891–900.

Fitch,h.s.1989.AfieldstudyoftheslenderGlasslizard,Ophisaurus attenu-atus innortheasternKansas.University of Kansas Museum of Natural History Occasional Paper125:1–50.

Note.—AllofhenryFitch’spaperspublishedbytheUniversityofKansasareincorporatedinthebiodiversityheritagelibrary,wheretheycanbedownloadedatwww.biodiversitylibrary.org.thefamilysuggestsdonationstohonorprofessorFitchbedirectedtothehenryandVirginiaFitchmemorialFundattheKUendowmentAssociation:<www.kuendowment.org>.

henryFitchattheentrancetotheFitchnaturalhistoryReservation,1997.

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henryFitch(right)inanultralightplaneinmarch2004.hisnephewflewhimovertheareawherehehadgrownupnearmedford.thisisagoodindicationofhenry’sspiritofadventureandloveoftravel—characteristicsheneverlost.

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scarletKingsnakes(Lampropeltis elapsoides),nowrecognizedasdistinctfromthemuchmorewidespreadmilksnake(L. triangulum),occuratlow

andmoderateelevationsfromVirginiatoFlorida,thencewesttoKentuckyandlouisiana(ConantandCollins1998).thetwospeciesaresympatricwithlittleornohybridizationatseveralcontactzones,thebeststudiedofwhichareinwesternKentuckyandadjacenttennessee(Armstrongetal.

2001)andnorthCarolina(harperandpfennig2008).thesesecretivecoralsnakemimics(GreeneandmcDiarmid2005,harperandpfennig2008)rangeintotallength(tl)from130mmathatchingtoamaximumof576mm(Wrightandbishop1915,Williams1988),and,alongwithotherlam-propeltines,areofinterestfromdiverseperspectives(e.g.,Rodríguez-RoblesanddeJésusescobar1999,pyronandburbrink2009).

Diet specialization by the scarlet Kingsnake, Lampropeltis elapsoides (colubridae)

harryW.Greene1,edmundJ.Zimmerer2,Williamm.palmer3,andmichaelF.benard4

1Departmentofecologyandevolutionarybiology,CornellUniversity,Ithaca,nY14853([email protected])2Departmentofbiologicalsciences,murraystateUniversity,murray,KY42071([email protected])

3northCarolinamuseumofnaturalsciences,Raleigh,nC276994Departmentofbiology,CaseWesternReserveUniversity,Cleveland,oh44106([email protected])

basedon34naturalpreyitems,Lampropeltis elapsoideseatsprimarilyelongatesquamates(97%),especiallyskinks(74%)andcolubroidsnakes(15%).noontogeneticorgeographicvariationisevident;preyitemsareswallowedheadfirstandaverage19%ofpredatormass.thedietsubstantiallyoverlapsthatofjuvenilesofsomeotherlampropeltines,includingsympatricL. triangulum,butisunusuallynar-rowcomparedtoadultsofmostotherspecies.

scarletKingsnakes(Lampropeltis elapsoides)occuratlowandmoderateelevationsfromVirginiatoFloridaandwesttoKentuckyandlouisiana.Incontrastwithpublishedreports,thesesnakesarenotdietarygeneralists.

IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 19LAMPROPELTIS ELAPSOIDES

herein,we:(i)provideafirstcriticalanalysisofthefeedingecologyofL. elapsoides;(ii)refuteimplicationsthatitisadietarygeneralist(e.g.,“foodincludessmallsnakesandlizards,babymice,smallfish,insects,andearthworms”;ConantandCollins1998:375);and(iii)assessitspotentialinteractionswithothersympatricsnakes,especiallyRedmilksnakes(L. t. syspila)intheKentucky-tennesseecontactzone.WeareespeciallypleasedthathenryFitchprovidedunpublisheddataforthisprojectandenjoyedreadingourmanuscriptduringthesummerofhis100thyear.

Materials and MethodsWeexaminedstomachcontentsofpreservedL. elapsoides(formethodsseeGreeneandRodríguez-Robles2003)atArchboldbiologicalstation,Carnegiemuseumofnaturalhistory,andnorthCarolinastatemuseumofnaturalsciences,thenintegratedthosedatawitheJZ.’sfieldobserva-tionsfromKentucky;anecdotesprovidedbyJ.D.Groves(pers.comm.)andJ.D.Wilson(savannaRiverecologylabfiles);andcredibleliteraturerecords(Carr1940,basedonFloridamuseumofnaturalhistory1568;K.l.Krysko,pers.comm.;mount1963;brown1979;palmerandbraswell1995,excludingaWormsnake,Carphophis amoenus,whichfieldnotesindicatewaseatenaftercapture;lee2006).WeomittedrecordscitedbyWilliams(1988)thatwerebasedoncaptivesorunsupportedbyexplicitdata(brimley1905,Ditmars1907,brodeandAllison1958);weexcludedWrightandbishop’s(1915:167)reportof“ananglewormand…twokil-lifishes,”becausethosepreyareotherwiseunknowninthedietofanylam-propeltine(Rodríguez-RoblesanddeJésusescobar1999)andmoreplau-siblystomachcontentsfromaningesteditem(e.g.,easternGartersnake,Thamnophis sirtalis).WeassesseddietaryoverlapwithL. t. syspilabasedonfiverecordsfromKentuckyandtennesseeobtainedbyeJZand22recordsfromKansas(Fitch1999,pers.comm.).

Resultsthirty-fourprey items from32L. elapsoides (mean1.1 items/snake)include13Scincella lateralis(Groundskink,including1setof5eggsand1tail),6Plestiodon inexpectatus(southeasternFive-linedskink),1P. egre-gius(moleskink),2Plestiodonsp.,3unidentifiedskinks,1setof4lizardeggs,2Aspidoscelis sexlineata(six-linedRacerunner),2Diadophis punctatus(Ring-neckedsnake),1Tantilla coronata(southeasternCrownedsnake),1Thamnophissp.(gartersnake),1Virginia striatula(Roughearthsnake),and1nestlingrodent.theyencompass≥9preyspecies,including33(97%)somewhattoveryelongate,mostlysmooth-scaledsquamatereptiles;25(74%)wereskinks(mainly2species)and5(15%)werecolubroidsnakes.only2atemultipleitems,sowecannotevaluatewhetherindividualsspe-cializeonparticularpreytypes.

the34preyitemsarefromFlorida(8),Kentucky(7),louisiana(1),mississippi(2),northCarolina(12),andsouthCarolina(4),andlizardspredominatedthroughouttherange.FloridaL. elapsoidesate7 lizardsandamouse;northCarolinapreyincluded10lizardsand2snakes,and,althoughKentuckysnakesoccupydistinctivehabitat(Armstrongetal.

Redmilksnakes(Lampropeltis triangulum syspila)showasignificantdietaryshiftwithincreasingsize,shiftingfromreptilestomammalsastheygetlarger).

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preyingmainlyonslendersquamates,suchasreclusiveskinksandsmallsnakes,thedietofscarletKingsnakesappearstobeconstrainedbyasmallgape.southeasternFive-linedskinks(Plestiodon inexpectatus;top)andGroundskinks(Scincella latera-lis;bottom)areprominentlyrepresentedinstomachsamples.

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2001),alloftheirpreywereskinks.snakeswithpreywerefoundinJanuary(1),march(1),April(4),may(1),June(5),July(3),andseptember(3).theymeasured196–496mmintl(mean354mm,n=23),ofwhichthetwosmallestateS. lateralis,oneofthem(tl197mm)onlyatail;thelargestcontainedanadultP. inexpectatusandtheonlymammalwasina436-mmadult.prey/predatormassratioswere0.11–0.38(mean0.19,n=6),ofwhichthelargestwasanA. sexlineatainasmalladultsnake(tl379mm),andallfiveitemsforwhichdirectionofingestionwasrecordedwereswallowedheadfirst. scarletKingsnakesexhibitnoontogeneticchangeindiet;theythuscompletelyoverlapthedietofjuvenileRedmilksnakesandpartlythatofadults.elevenL. t. syspilawithinthesizerangeofL. elapsoides(<576mmtl)hadeaten8skinksand3smallsnakes,whereas17largeradults(600–890mmtl)hadconsumed4skinks(18%),alimblesslizard,and17mammals(77%)—asignificantdietaryshiftwithincreasingsize(reptilesversusmammals;Fisher’sexacttest,p<0.0001).

Discussionbypreyingmainlyonslendersquamates,L. elapsoidesofallsizesresemblejuvenilesofL. californiae(K.Wisemanandh.W.Greene,unpubl.data),L. triangulum(Fitch1999;m.F.benardandh.W.Greene,unpubl.data),

andL. zonata(GreeneandRodriguez-Robles2003),aswellasadultsof

GReeneetAl.

table 1.DataforindividualscarletKingsnakes(Lampropeltis elapsoides)andtheirprey.mR=massratio.

Locality Size Prey MR Direction Date SourcelA 353mm,9g 1Thamnophissp.,1g 0.11 headfirst Cm91903Fl 379mm,16g 1Aspidoscelis sexlineata,6g 0.38 headfirst AbsmI 403mm,18g 2 Diadophis punctatus,2g,3g 0.11 lee(2006) 0.17Fl 413mm,nA 1Plestiodon inexpectatus,nA AbsFl 429mm,13g 1skink,2g 0.15 headfirst 9/1936 Cm19840Fl 436mm,20g 1rodent,4g 0.20 AbsGA 1angleworm,2killifishes CUmV6242,W&bFl 196mm 1Scincella lateralis 1/1937 Carr(1940)Fmnh1568Fl 1Scincella lateralis Carr(1940)Fl ~10in UIDskinkscalesinfeces J.D.Groves(inlitt.)Fl 300mm Plestiodon egregius,tail mount(1963)sC 2Scincella lateralis brown(1979)sC 1 Scincella lateralis brown(1979)sC 348mm,12.5g 1Scincella lateralis 4/17/07 J.D.Wilson(sRel)nC,moore adultfemale 1Aspidocelis sexlineata,juv. 9/1/2004 nCsm04-2049nC,Richmond adultfemale 1Plestiodon inexpectatus 4/13/2002 nCsm02-364nC,Craven 197mm 1Scincella lateralis,tail 9/12/1968 nCsm33820nC,beaufort 483mm 1Scincella lateralis[5eggs] 7/1/1968 nCsm/pbnC,Craven 394mm 1Plestiodon inexpectatus 5/24/1968 nCsm33819/pbnC,hyde 496mm 1Plestiodon inexpectatus,adult 4/30/1960 nCsm972/pbnC,Cateret 465mm 1Plestiodon inexpectatus,adult 4/25/1970 nCsm9253/pbnC,bladen 294mm 1Scincella lateralis,adult nCsm15005/pbnC,brunswick 1Plestiodon inexpectatus p&b(1995)nC,brunswick 1Virginia striatula p&b(1995)nC,scotland 1Scincella lateralis p&b(1995)nC,scotland 1Tantilla coronata p&b(1995)KY 370mm 1setof4reptileeggs(6–7mm) 7/98 emZ1523KY 375mm 1Plestiodonsp. headfirst 6/98 emZ1525KY 290mmsV 1Scincella lateralis,4cmsV headfirst 7/98 emZ1527 (est.334tl)KY 295mm 1Plestiodonsp.,4cmsV 6/99 emZKY 290mm 1skink 6/05 emZKY 320mm 1Scincella lateralis 6/05 emZKY 315mm 1Scincella lateralis,4cmsV 6/06 emZ

preytypesinrelationtopredatorsizeforscarletKingsnakes(Lampropeltis elapsoides)andRedmilksnakes(L. triangulum syspila),basedonsamplesfromasympatriccontactzoneandelsewhereintherangeofeachspecies.

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someothersmallcolubroidsnakes(e.g.,Hypsiglena,Rodríguez-Roblesetal.1999).ComparedtootherLampropeltis,scarletKingsnakeshavepointedsnoutsandfusedheadscalessuggestiveoffossorialhabits(Wrightandbishop1915,Williams1988),andtheyaretypicallyfoundundercoverobjects,especiallybarkonoldfallenlogsandverticalstumps(palmerandbraswell1995,Reichling2008).Accordingly,thisspeciesmightsimplybeahabitatspecialist,itsnarrowdietconstrainedbysmallgapeandhighencounterratesforreclusiveskinkandsnakeprey. AdditionallinesofevidenceimplyscarletKingsnakesarespecializedpredators(forgeneraldiscussionsseeRodríguez-RoblesandGreene1999,brischouxetal.2009).With82%ofitsdiet lizardsand89%ofthoseitemsskinks,L. elapsoideshasanarrowerdietthananotherlampropeltine,Rhinocheilus lecontei(long-nosedsnake),withadietconsistingof65%lizardsand72%(48%oftotalitems)teiids(Aspidoscelis);respectivemeansfor55othersnakespeciesinwhichlizardsaremodalpreyare75%and47%(Rodríguez-RoblesandGreene1999).moreover,L. elapsoideshasthehighestincidenceofskinksintheadultdietofanynorthAmericanserpent(ernstandernst2003)anditsanteriorteethareenlargedlikethoseofotherskinkspecialists(Greene1989).Finally,throughoutitsrange,L. elapsoidesco-occurswithS. lateralisandtwotofourspeciesofPlestiodon,whereas,elsewhereinnorthAmerica,nomorethantwospeciesofskinksaretypi-callysympatric(mapsinConantandCollins1998,Jonesandlovich2009). Intermsofpreytakenbyothersympatricsnakes,L. elapsoidespartlyoverlapsColuber constrictor(Racer),L. getula(CommonKingsnake),L. extenuata(short-tailedsnake),andMicrurus fulvius(easternCoralsnake),ofwhichthefirsttwohavemuchbroaderdietsandtheothersaremod-eratelytoexclusivelyspecializedonsnakes(ernstandernst2003).AsmacArthur(1972)pointedout,onewaytotrumppotentialcompetitorsis

toeatthem,andM. fulviusindeedpreysonbothP. inexpectatus(Absfieldnotes)andL. elapsoides(KryskoandArbdelfattah2002).noneofthoseothersnakesspecializesonskinks,butwehavedemonstratedthatjuvenilesofL. t. syspiladoso,andadultsofthatspeciespartiallyoverlapthedietofsympatricL. elapsoides.AdditionalstudiesofscarletKingsnakesinacom-munityecologicalcontextwilllikelyprovidefurtherinsightsintospecializa-tion,speciation,mimicry,andotherevolutionaryprocesses.

AcknowledgmentsWethankK.l.Krysko,J.n.layne,ands.p.Rogersforaccesstoand/orinfor-mationonpreservedspecimens;h.s.Fitch,J.D.Groves,andJ.D.Wilsonforuseoftheirobservations;andJ.D.Grovesandl.J.Vittforinsightfulfeedback.

Literature CitedArmstrong,m.p.,D.Frymire,andeJ.Zimmerer.2001.Analysisofsympatric

populationsofLampropeltis triangulum syspilaandLampropeltis triangulum elapsoides,inwesternKentuckyandadjacenttennesseewithrelationtothetaxonomicstatusofthescarletKingsnake.Journal of Herpetology35:688–693.

blanchard,F.n.1921.ArevisionofthekingsnakesgenusLampropeltis.Bulletin of the U.S. National Museum114:1–260.

brimley,C.s.1905.notesonthefoodandfeedinghabitsofsomeAmericanrep-tiles.Journal of the Elisha Mitchell Society21:149–155.

brischoux,F.,X.bonnet,andR.shine.2009.Determinantsofdietaryspecializa-tion:Acomparisonoftwosympatricspeciesofseasnakes.Oikos118:145–151.

brode,W.e.andp.Allison.1958.burrowingsnakesofthepanhandlecountiesofmississippi.Herpetologica14:37–40.

brown,e.e.1979.somesnakefoodrecordsfromtheCarolinas.Brimleyana1:113–124.

Carr,A.F.,Jr.1940.AcontributiontotheherpetologyofFlorida.University of Florida Publications, Biological Science Series3:1–118.

table 2.DataforindividualRedmilksnakes(Lampropeltis triangulum syspila)andprey.

Locality Size Prey Direction Date Source

tn,montgomery 720mm mammalhair 6/97 emZKY,lyon 755mm 1Peromyscussp. 6/97 emZKY,trigg 770mm mammalhair 6/97 emZKY,marshall 775mm 1Plestiodonsp.80mm headfirst 6/00 emZtn,lake 890mm 1Plestiodonsp.70mm 10/03 emZtn,lake 800mm 1Peromyscussp.,youngwithhair 5/06 emZKs 228mm 1Diadophis punctatus 6/18/66 hsFKs 278mm 1Plestiodon fasciatus,1styr 4/86 hsFKs 283mm 1Plestiodon obsoletus,tailonly 9/21/66 hsFKs 291mm 1Plestiodon fasciatus,juv. 4/30/66 hsFKs 342mm 1Plestiodon fasciatus,adult 9/25/64 hsFKs 355mm 1Plestiodon fasciatus,juv. 9/17/73 hsFKs 406mm 1Plestiodon fasciatus,gravidad. 6/14/66 hsFKs 430mm 1Carphophis amoenus,adult 5/26/55 hsFKs 430mm 1set3Diadophis punctatuseggs 7/16/78 hsFKs 515mm 1Plestiodon fasciatus,ad.&5eggs 6/26/78 hsFKs 573mm 1Plestiodon fasciatus,adult 5/22/66 hsFKs 600mm 1Plestiodon fasciatus,1styr 5/18/60 hsFKs 626mm 2Microtus ochrogaster,juveniles 6/7/93 hsFKs 663mm 1Plestiodon fasciatus,adult 5/7/67 hsFKs 673mm,90g 4Microtus ochrogaster,juv.@15g 10/8/86 hsFKs 688mm 1Blarina hylophaga 5/23/93 hsFKs 730mm 1Blarina hylophaga 4/17/81 hsFKs 782mm 1Cryptotis parva 10/10/90 hsFKs 815mm 1Microtussp. 8/18/67 hsFKs 845mm 2Microtus ochrogaster,nestlings 7/23/92 hsFKs 852mm 1Microtus ochrogaster,adult 5/26/90 hsFKs 884mm 1Ophisaurus attenuatus,adult 8/1/61 hsF


Conant,R.andJ.t.Collins.1998.A Field Guide to Reptiles and Amphibians: Eastern and Central North America(3rded.).houghtonmifflin,boston,massachusetts.

Ditmars,R.l.1907.The Reptiles of North America.Doubleday,GardenCity,newYork.

ernst, C.h. and e.m. ernst. 2003. Snakes of the United States and Canada.smithsonianbooks,Washington,D.C.

Fitch,h.s.1999.A Kansas Snake Community: Composition and Change over 50 Years.KriegerpublishingCo.,malabar,Forida.

Greene,h.W.1989.DefensivebehaviorandfeedingbiologyoftheAsianmockViper,Psammodynastes pulverulentus(Colubridae),aspecializedpredatoronscincidlizards.Chinese Herpetological Research2:21–32.

Greene,h.W.andR.W.mcDiarmid.2005.Wallaceandsavage:heroes,theo-ries,andvenomoussnakemimicry,pp.190–208.In:m.A.Donnelly,b.I.Crother,C.e.Guyer,m.h.Wake, andm.e.White (eds.),Ecology and Evolution in the Tropics: A Herpetological Perspective.UniversityofChicagopress,Chicago,Illinois.

Greene, h.W. and J.A. Rodríguez-Robles. 2003. Feeding ecology of theCaliforniamountainKingsnake,Lampropeltis zonata(Colubridae).Copeia2003:308–314.

harper,G.R.andD.W.pfennig.2008.selectionoverridesgeneflowtobreakdownmaladaptivemimicry.Nature451:1103–1107.

Jones,l.l.C.andR.e.lovich(eds.).2009.Lizards of the American Southwest: A Photographic Field Guide.Rionuevopress,tucson,Arizona.

Krysko,K.l.andK.R.Arbdelfattah.2002.Micrurus fulvius(easternCoralsnake).predation.Herpetological Review33:57–58.

lee,J.R.2006.Lampropeltis triangulum elapsoides(scarletKingsnake).ophiophagy.Herpetological Review37:231.

macArthur,R.h.1972.Geographical Ecology.harperandRow,newYork.

mount,R.h.1963.thenaturalhistoryoftheRed-tailedskink,Eumeces egregiusbaird.American Midland Naturalist70:356–385.

palmer,W.m.andA.l.braswell.1995.Reptiles of North Carolina. UniversityofnorthCarolinapress,Chapelhill.

pyron,R.A.andF.t.burbrink.2009.neogenediversificationandtaxonomicstabilityinthesnaketribelampropeltini(serpentes:Colubridae).Molecular Phylogenetics and Evolution52:524–529.

Reichling,s.b.2008.Reptiles and Amphibians of the Southern Pine Woods.UniversitypressofFlorida,Gainesville.

Rodríguez-Robles,J.A.andh.W.Greene.1999.Foodhabitsofthelong-nosedsnake, Rhinocheilus lecontei, a “specialist” predator? Journal of Zoology (London) 248:489–499.

Rodríguez-Robles,J.A.andJ.m.deJésusescobar.1999.molecularsystematicsofnewWorldlampropeltinesnakes(Colubridae):Implicationsforbiogeog-raphyandevolutionoffoodhabits.Biological Journal of the Linnean Society68:355–385.

Rodríguez-Robles,J.A.,D.G.mulcahy,andh.W.Greene.1999.Feedingecol-ogyof theDesertnightsnake,Hypsiglena torquata (Colubridae).Copeia 1999:93–100.

Williams,K.l.1988.Systematics and Natural History of the American Milk Snake,lampropeltis triangulum (2nd rev. ed.). milwaukee public museum,milwaukee,Wisconsin.

Wright,A.h.ands.C.bishop.1915.AbiologicalreconnaissanceoftheokefinokeeswampinGeorgia.II.snakes.Proceedings of the Academy of Natural Sciences of Philadelphia67:139–192.

KApFeR

Fitch(1987a)indicatedthatmorphologicalmeasurements,particularlymeasuresofsnout-to-ventlength(sVl),providesomeofthemostuse-

fulinformationthatcanbeobtainedfromfield-basedresearchonsnakes.Unfortunately,publishedresearchthatfocusesonnaturalhistory,whichincludesmorphologicaldatasuchassnakesize,hasdeclinedsharplyinrecentyears(hendersonandpowell2009,mcCallumandmcCallum2006).Insomecases,naturalhistoryobservationsconsidered“anecdotal”areeventreatedwithscornbyresearchers.AsFitch(1987b)suggested,suchreactionstolifehistorystudiesareunfortunate,andinformationthatmaybeconsideredanecdotalstillhasavaluableroleinincreasingtheunder-standingofmanyspecies’ecologicalneeds.therefore,thisinformationshouldbepublished. Considerabledatahavebeenpublishedontheecologyandlifehistoryofseveralnatricinesnakespecies,particularlytheCommonGartersnakes(Thamnophis sirtalis;e.g.,Fitch1965,1999,2001).theseincludeinfor-mationonaspectsoftheirmorphology,suchassize(sVl)andweight.however,certainsmallfossorialspecies,suchasmembersofthegenusStoreriahave received lessattention. Inaddition,althoughthe sizeof“mature”femaleshasbeenreportedforseveralspeciesatsomelocationsintheuppermidwesternUnitedstates(e.g.,ohioandmichigan),suchinformationisrarefrompopulationsinWisconsin.Furthermore,fewifanydatafromWisconsinhavebeenpublished,withthepossibleexcep-

tionoftechnicalreportsthatarenoteasilyobtained.suchinformationisvaluablefordeterminingnumerousaspectsofthebiologyofthesespecies,suchassizeatsexualmaturityandmorphologicalcharacteristicsofregional

a survey of Gravid snakes at several sites in southern Wisconsin

Joshua M. KapferDepartment of Environmental Studies,Elon University,Elon, North Carolina 27244 ([email protected])

Photographs by the author except where indicated.

AnexampleoftypicaleasternGartersnake(Thamnophis sirtalis sirtalis)habitatinWisconsin.


populations.therefore,myobjectivewastosurveyforgravidnatricinesatseveralsitesinsouthernWisconsin,withparticularinterestintheunder-studiedmembersofthegenusStoreria, todetermineifthesizesofgravidsnakesencounteredconformedtopreviouslypublishedreportsofadultfemalesizeorsizeatmaturity.Inaddition,thegoalofthesesurveyswastoprovidebaselinenaturalhistorydatathatmayhavecurrentandfuturevaluetoherpetologists,ecologists,andconservationbiologists. Iconductedsurveysforgravidfemalesatfivesitesacrossthreecoun-tiesinsouthernWisconsinduringthelatespring-summerof2009.IchosethisperiodtoconductmysurveysbecauseitiswithintheseasonduringwhichWisconsinspeciesaregravid.Althoughsitesvariedinarea,ratioofwetlandtoupland,andplantcommunitycomplexity,allsurveylocationspossessedopenshallowmarshhabitatswithassociatedopenuplandhabitats(primarilygrasslandorfallowgrassland).eachsitewassurveyedviaartificialcoverobjects(Fitch1987a)intheformof¾-inchplywoodcutin3x4-ftsheets.boardswereplacedopportunisticallyinareasthatappearedsuit-ablefornatricinesnakes,withaparticularfocusonspeciesofThamnophis,Storeria,andNerodia.boardswerecheckedafter1700honlyondaysthathadbeenpartlysunnytosunny.thisinsuredthatthemicroclimateundereachboardwouldbewarmenoughtoattractgravidfemales,butnotsowarmastobeunsuitable.Graviditywasdeterminedbypalpationofthelowerthirdofthesnaketocheckforthepresenceofformedembryos.ImeasuredthesVlofgravidsnakesbygently,butfirmly,stretchingthemalongatapemeasureasdescribedinFitch(1999). Duetoresourcelimitations,thesurveyeffortateachsitevaried.Atsomelocations,coverobjectsweredeployedseveralyearspriortoinitiation

ofthestudy,whereasatotherlocations,boardsweredeployedinlatespring2009,immediatelypriortoinitiationofsurveys.Inaddition,eachsitewasnotsurveyedthesamenumberoftimes,andanequalnumberofcoverobjectswasnotimplementedequallyacrosssites.mudlakeandWestford,forexample,eachhadtencoverobjectsandwerevisitedsixtimesthrough-outthelatespringandsummer.Cedarburg,ontheotherhand,had15coverboardsandwassurveyedfourtimes.FrenchCreekhad10boardsandwassurveyedfourtimes,whereashoriconhad20boardsdividedamongfourlocationsandwassurveyedninetimes.however,becausetheobjectiveofthesestudies(i.e.,tosamplegravidfemaleson-siteandmeasuretheirsVl)didnotdependuponequitablesurveyeffortsacrosssites,thesevaria-tionsweredeemedacceptable. throughoutthecourseofthesesurveys,Iencounteredadultgravidfemalesofthreenatricinespecies:northernbrownsnake(Storeria dekayi dekayi),Redbellysnake(Storeria occipitomaculata occipitomaculata),andeasternGartersnake(Thamnophis sirtalis sirtalis;table1).however,Ididnotfindallthreespeciesateverysurveylocation.Gravidsnakesalsowerefoundatallsurveylocations.Forexample,attheFrenchCreeksite,IfoundonlyonegravidRedbellysnakeandnogravidfemalesofotherspecies.brownsnakeswereencounteredonlyatthemudlakesite,whereasgravideasternGartersnakeswerefoundatallsitesexceptFrenchCreek(table1).overall,gravideasternGartersnakeswerethemostfrequentlyencoun-teredsnakesacrossallsites(n=30).thesewerefollowedbynorthernbrownsnakes(n=21)andRedbellysnakes(n=5).becauseIdidnotmarkindividuals,Imayhavecollectedandmeasuredthesamesnakeonmorethanoneoccasion.

GRAVIDsnAKesInsoUtheRnWIsConsIn

table 1.samplesizes,meansnout-vent-length(sVl)inmmforeachspeciescapturedpersurveylocation.

Site Wisconsin Sample Mean SVL Species Name County Size ± SD (range)

S. dekayi Mud Lake Dodge 21 264 ± 25 (241–295)

S. occipitomaculata Westford Dodge 2 231 ± 22 (215–248)

S. occipitomaculata French Creek Marquette 1 196

S. occipitomaculata Horicon Dodge 2 225 ± 16 (214–237)

T. sirtalis Cedarburg Ozaukee 11 471 ± 72 (356–620)

T. sirtalis Horicon Dodge 10 546 ± 53 (469–603)

T. sirtalis Mud Lake Dodge 5 562 ± 36 (524–609)

T. sirtalis Westford Dodge 4 492 ± 39 (450–533)

Grasslandhabitatisoftenpreferredbybrownsnakes(Storeria dekayi)inWisconsin. brownsnakes(Storeria dekayi)werefoundatonlyoneofthesurveylocations.

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Ingeneral,thesVlofgravidfemalesofallspeciesencountereddidnotvarygreatlyfrompastreports(table2).onaverage,gravidbrownsnakeswere264±25mminsVlwithlittlevariationinindividualsVl.thisspecieshasbeenreportedtoshowsignificantgeographicvariationinsVl(King1997),whichalsohasbeennotedforThamnophis sirtalis(King1989).GravidRedbellysnakeswereslightlysmallerthanbrownsnakeson

average,butsizesvariedlittle(222±20mm).theaveragesVlofgravideasternGartersnakescapturedduringmysurveyswas514mm,makingthemthelargestsnakeencountered.Unliketheothertwospecies,however,considerablevariationinsVlwasobservedinthisspecies(sD=67mm). AlthoughRedbellysnakesVlsreportedfrompaststudiesaresmallerthantheaverageIrecordedforindividualsinWisconsin,mysamplesizewas

KApFeR

table 2.sizecomparisonsofadultfemalesnakesencounteredduringthisstudyandpastpublishedreports.measurementsofsnakesreportedtobegravidareindicatedwithasingleasterisk(*);publicationsreportingestimatedsizeatsexualmaturityareindicatedwithadoubleasterisk(**).

Location Sample AverageSpecies Source of Study Size SVL (mm)

S. dekayi thisstudy Wisconsin 21 264*

S. dekayi WilsonandDorcas(2004) northCarolina 9 245

S. dekayi minton(2001) Indiana 21 266

S. dekayi Fitch(1999) Kansas 22 256*

S. dekayi King(1997) ontario/ohio nA 230**

S. dekayi Kofron(1979) louisiana 30 170–175**

S. occipitomaculata thisstudy Wisconsin 5 222*

S. occipitomaculata Wilson&Dorcas(2004) northCarolina 7 181

S. occipitomaculata minton(2001) Indiana 10 198

S. occipitomaculata brodie&Ducey(1989) newYork 42 209*

S. occipitomaculata semlitsch&moran(1984) southCarolina 37 157

S. occipitomaculata blanchard(1937) michigan 71 194

T. sirtalis thisstudy Wisconsin 30 514*

T. sirtalis minton(2001) Indiana 30 531

T. sirtalis Fitch(1999) Kansas 1592 639

T. sirtalis Rossmanetal.(1996) Various nA 420–550**

T. sirtalis mitchell(1994) Virginia — 515

T. sirtalis larsenetal.(1993) Canada 23 745*

T. sirtalis King(1989) lakeerieIslands 345 683

T. sirtalis hebard(1950) Washington 32 408

Redbellysnakes(Storeria occipitomaculata)werefoundinhabitatsuchasthisdur-ingsurveys.

Redbellysnakes(Storeria occipitomaculata)wereencounteredatmoresitesthanbrownsnakes(S. dekayi),butinlowerdensities.

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IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 25GRAVIDsnAKesInsoUtheRnWIsConsIn

small(n=5),andIrecommendcautionwheninterpretingthis.however,tomyknowledge,fewpublishedreportshavelargesample-sizesofthisspecies.Forexample,WilsonandDorcas(2004)reportedcapturingsevenfemalesoverfiveyearsofsurveysinnorthCarolina;brodieandDucey(1989)reportedasampleof42gravidfemalescapturedovertwoyearsinnewYork;semlitschandmoran(1984)captured37females;andblanchard(1937),whocommentedspecificallyonhowinfrequentlytheyareencountered,ana-lyzed71femalesforwhichhehadacquireddataover12years.Inaddition,althoughmysamplesweresmall,despitebeingspreadacrossthreesitesintwocounties,littlevariationwasobservedinsVl(sD=20mm). Rossmanetal.(1996)reportedthatfemale Thamnophis sirtalis matureat420–550mmandalloftheaveragesVlsthatIobservedpersitefitwithinthisrange(table2).Atonelocation(Cedarburg),Iexaminedtwogravidindividualsthatwerebelowthisthreshold(397mmand356mm).Fitch(1999)reportedthatthesmallestgravidfemale,amongthehundredsthatheanalyzedover50yearsatasiteinKansas,measured490mminsVl.AtleasttengravidindividualsthatIencounteredatallsurveyloca-tionswereshorterthanthis. Insummary,thegravidsnakesofallspeciesencounteredduringmysur-veysweresimilarinsVltopreviouslypublishedreports.Unfortunately,thesmallsamplesizesandrelativelyshortsurveyperiod(i.e.,oneseason)ofthisworklimitscomparisonswithotherstudies.moredataarenecessarytomakedefinitiveconclusionsaboutthesizesofgravidsnakesinthisregionofnorthAmerica.

AcknowledgementsIthanknaturalResourcesConsulting,Inc.andtheWisconsinDepartmentofnaturalResources-bureauofendangeredResourcesforsupplyingcoverboardsthatwereused,inpart,toobtainthisinformation.IthanktheU.s.FishandWildlifeservice,theWisconsinDepartmentofnaturalResources,andtheUniversityofWisconsin-milwaukeeFieldstationforgrantingmeaccesstotheirpropertiesforsurveywork.Roberthenderson(milwaukeepublicmuseum)graciouslyreadandcommentedonanearlierdraftofthismanuscript.

Literature Citedblanchard,F.1937.DataonthenaturalhistoryoftheRed-belliedsnake,Storeria

occipitomaculalta(storer),innorthernmichigan.Copeia1937:151–162.

brodie,e.D.andp.K.Ducey.1989.AllocationofreproductiveinvestmentintheRedbellysnake,Storeria occipitomaculata. American Midland Naturalist122:51–58.

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

Fitch,h.s.2001.FurtherstudyoftheGartersnake,Thamnophis sirtalis,innortheasternKansas.Scientific Papers, University of Kansas Museum of Natural History19:1–6.

Fitch,h.s.1987a.Collectingandlifehistorytechniques,pp.143–164.In:R.A.seigel,J.t.Collins,ands.s.novak(eds.),Snakes: Ecology and Evolutionary Biology.theblackburnpress.Caldwell,newJersey.

Fitch,h.s.1987b.thesinofanecdotalwriting.Herpetological Review18:68.

Fitch,h.s.1965.AnecologicalstudyoftheGartersnake,Thamnophis sirtalis.University of Kansas Museum of Natural History15:493–564.

hebard,W.b.1950.RelationshipsandvariationintheGartersnake,genusThamnophis,ofthepugetsoundregionofWashingtonstate.Herpetologica6:97–101.

henderson,R.W.andR.powell.2009.thestateofnaturalhistory:AperspectivefromtheliteratureonWestIndianherpetology.Herpetological Review40:273–275.

King,R.b.1997.Variationinbrownsnake(Storeria dekayi)morphologyandsca-lation:sex,family,andmicrogeographicdifferences.Journal of Herpetology31:335–346.

King,R.b.1989.bodysizevariationamongislandandmainlandsnakepopulationsnearlakeerie.Herpetologica45:84–88.

Kofron,C.p.1979.Femalereproductivebiologyofthebrownsnake,Storeria dekayi,inlouisiana.Copeia1979:463–466.

larsen,K.W.,p.t.Gregory,andR.Antoniak.1993.ReproductiveecologyoftheCommonGartersnakeThamnophis sirtalisatthenorthernlimitofitsrange.American Midland Naturalist129:336–345.

mcCallum,m.l.andJ.l.mcCallum.2006.publicationtrendsofnaturalhistoryandfieldstudiesinherpetology.Herpetological Conservation and Biology1:62–67.

minton,s.A.2001.Amphibians and Reptiles of Indiana.Revised2nded.IndianaAcademyofsciences,Indianapolis.

mitchell, J.C. 1994. The Reptiles of Virginia. smithsonian Institute press,Washington,D.C.

Rossman,D.A.,n.b.Ford,andR.A.seigel.1996.The Garter Snake: Evolution and Ecology.theUniversityofoklahomapress,norman.

semlitsch,R.D.andG.b.moran.1984.ecologyoftheRedbellysnake(Storeria occipitomaculata)usingmesichabitatsinsouthCarolina.American Midland Naturalist111:33–40.

Wilson,J.D.andm.e.Dorcas.2004.AspectsoftheecologyofsmallfossorialsnakesinthewesternpiedmontofnorthCarolina.Southeastern Naturalist3:1–12.

theeasternGartersnake(Thamnophis sirtalis sirtalis)wasthemostfrequentlyobservedsnakeduringsurveys.


Daytime amphibian surveys in Three protected areas in the Western Great lakes

KennethD.bowenanderikA.beever

U.s.nationalparkservice,GreatlakesInventoryandmonitoringnetwork,Ashland,Wisconsin54806([email protected])

WeperformedpilotmonitoringofamphibianpopulationsatApostleIslandsnationallakeshore(ApIs)in2006,picturedRocksnationallakeshore(pIRo)in2007,andsleepingbearDunesnationallakeshore(slbe)inboth2006and2007.Weperformeddaytimesurveys(usingmultiplemethods)atnumeroussitesineachofthethreeparks.Wedetected10amphibianandtworeptilianspeciesatApIs,nineamphibianandfourreptilianspeciesatslbe,andnineamphibianandonereptilianspeciesatpIRo.noonedaytimesurveytechniqueappearedtobesuperiortoanyother.ourworkresultedintwonewspeciesrecords(GraytreefrogandGreenFrog)forbasswoodIslandatApIs.

theGreatlakesInventoryandmonitoringnetwork(hereafterGlKnorthenetwork)wasformedbytheU.s.nationalparkservice(nps)

in1999andisoneof32networksofparksthatsharecommongeographyandmanagementpriorities.thepurposeofGlKnistoinventoryandmonitornaturalresourceswithinninenationalparkunitsinthenorth-ernGreatlakesecoregion,includingIndiana,michigan,minnesota,andWisconsin(Routeandelias2007).In2000,GlKnbeganabiologicalinventoryinnetworkparks(Route2000),and,in2002,thenetworkbeganplanninga“Vitalsigns”monitoringprogram.Vitalsignsaredefinedasaselectgroupofattributesthatareparticularlyrichininformationneededforunderstandingandmanagingnpsareas(Route2004).Vitalsignswerechoseninpartbasedonhowtheyreflectthehealthofparkeco-systemsandhowtheyrespond(orarehypothesizedtorespond)tonaturaloranthropogenicstressors.AprioritizedlistofVitalsignswasfinalizedin2004andamphibianpopulationswereoneoftheVitalsignschosenforearlyprotocoldevelopment(Routeandelias2007). AmphibianpopulationswerechosenasaGlKnVitalsignforsev-eralreasons.manyspeciesofamphibiansneedbothaquaticandterrestrialhabitatsforlifecyclecompletion,andthereforeprovideabiologicallinkbetweenlandandwaterandthestressorsofeach(stebbinsandCohen1995,semlitsch2000).Concordantly,amphibiansareimportantcom-ponentsofbothforestandwetlandecosystems.Amphibiansoftenoccurathighdensityandthereforeoccupyanimportantpositioninfoodwebswhilepotentiallydominatingenergytransferbetweenterrestrialandaquatichabitats(stebbinsandCohen1995,WelshandDroege2001,Gibbonsetal.2006).Finally,amphibiansaresensitivetoawidevarietyofnaturalandanthropogenicstressors(AlfordandRichards1999,booneetal.2007,DavidsonandKnapp2007),andtheworldwidedeclineofamphibianspe-ciesdiversityandabundanceiswell-documented(Wake1991,Green1997,lannoo2005). In2006and2007,theGlKnperformedpilotworksurveyingforamphibianswithinnetworkmanagementunits.theportionofthepilotworkreportedonherehadtwoprimaryobjectives:(1)togatherinforma-tiononspeciesdistributionandabundanceforparkinventoriesandasabaselineforfuturework,and(2)totesttheefficiencyandeffectivenessofdaytimesurveyandresearchmethods.

MethodsStudy Areas.—We performed pilot work at Apostle Islands nationallakeshore(ApIs)in2006,picturedRocksnationallakeshore(pIRo)in2007,andsleepingbearDunesnationallakeshore(slbe)inboth2006

and2007.ApostleIslandsnationallakeshoreislocatednearbayfield,Wisconsinandconsistsofanarchipelagoof21islandsandanarrow12-milesegmentalongthemainlandshoreoflakesuperior.theparkisprimarilyhemlock-hardwoodforest(butcontainselementsofsouthernborealfor-est),andhasawidediversityofcoastalfeatures.About190,000peoplevisittheparkannually.picturedRocksnationallakeshoreisheadquarteredinmunising,michiganandislocatedalongthesouth-centralshoreoflakesuperiorwithinatransitionzonebetweentheborealandeasterndeciduousforest.Wetlandsarecommonthroughoutthepark.About450,000peoplevisitannually.sleepingbearDunesnationallakeshoreisheadquartered

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Fig. 1.Daytimesurveypointsfor2006atApostleIslandsnationallakeshore,Wisconsin.Greenindicatesparkland.


inempire,michiganalongthenortheasternshoreoflakemichigan.theparkincludestwolargeislandsinlakemichiganaswellas65milesoflakemichiganshoreline,26inlandlakes,andfourstreams.About1.2millionpeoplevisittheparkannually(Routeandelias2007). Site Selection.—Daytimesurveysiteswerechosenbyseveralmeth-ods,dependingonthelogisticaldifficultiespresent.AtApIs,samplingofwetlandswasconstrainedbynumerouslogisticalfactors.theseincludedlakeconditionsandtheavailabilityofwatercraftandqualifiedpilots.theresource-managementstaffatApIscommunicatedthatonlya limitednumberofpermanentwetlandswerepresentontheislandgroup,andwesampledallofthesitesthatwereidentified(10sites;Fig.1). samplingofwetlandsatslbewaslimitedbyseveralfactors.thelarg-estsourceoflittoralhabitatswaslakemichigan.however,muchofthiswasunprotected,andthusexposedtotoomuchwaveactiontobesuitableasamphibianhabitat.thatactionalsocreatedsandybottomswithoutsub-mergedoremergentvegetationalongmuchofthecoastline.Additionally,manybodiesofwaterhadprivatein-holdingswithinthelakeshorebound-aries,andthuswerenotavailableforsampling.Waterbodiesknowntotheresourcemanagersthatwerenotaffectedbytheaboveconstraintswereselectedforsampling(sevensites;Fig.2).

siteselectionatpIRowasmoreprobabilistic.AllwetlandsfromthenhD(nationalhydrographyDataset)databasewithareasof0.02–2.0hawereassignedrandomnumbers,whichweusedtoselectthetop20of108sites.thenatural-resourcesstaffatpIRoindicatedwhichofthese20sitestheyconsideredreasonableforsampling(i.e.,relativeaccessibilityofhabitatswithinoneday’stime;Fig.3). Field Methods.—methodsfordaytimesurveysconsistedoffivecom-ponentsateachsite:callsurvey,samplingofphysicalandchemicalattri-butesofthewetland,visualencountersurvey(heyeretal.1994)anddip-netsweep(thomsetal.1997),andperimetersearch.Weperformedthesurveysinthatorderifwearrivedatthesiteinthemorning;buttoincreasedetectabilityinthecallsurveys,weperformedthoselastifwearrivedintheafternoon.twoobserversperformedonesubsampleofeachtypeofsurveyateachsite,oroneobserverperformedtwoofeachtypeofsurveyateachsiteiftwoobserverswereunavailable.Wherepossible,daytimesurveysitesweresampledonceeachduringeachofthreeseasons(Weirandmossman2005).the“earlyspring”seasonroughlycorrespondedtotheperiodfromearlyApril–earlymay,the“latespring”seasontomid-may–earlyJune,andthe“summer”seasontomid-June–earlyJuly. Callsurveysconsistedofstandinginanopenlocationontheperiph-eryofthesiteandlisteningforcallinganuransforaten-minuteperiod.GeneralmethodologyforcallsurveysfollowedWeirandmossman(2005).Werecordedthespeciescalling,themaximumcallingindexvalueforeachspecies(Weirandmossman2005),andthetimetofirstdetection(ttFD)foreachspecies. Forvisualencounteranddip-netsurveys,theobserverwalkedatran-sectthroughthewetland.transectswerelocatedarbitrarily(butfarenoughapartsothatobserversdidnotdisturbeachother)alongtheedgeofopenwaterandconsistedoftennodes,witheachnodebeingtwominutesinduration.Attheendofeachtwo-minutenode,theobserverperformedadip-netsweep(~1minlength).Werecordedthespeciesobserved,theapproximatenumberof individualsofeachspecies,andthettFDofeachspeciesduringeachtwo-minutenode.Wealsorecordedthespeciesobservedandtheapproximatenumberofeachspeciesforeachdip-netsweep.Anyensnaredanimalswereimmediatelyreleasedatthepointofcapture. perimetersurveysconsistedoftheobserverwalkingalongtheland-waterinterfaceofthesiteor,alternatively,alongtheedgeofthewetlandbasinifitwasclearlydefined.startingpointswerelocatedarbitrarily,butwerefarenoughapartsothatobserversdidnotdisturbeachotherwhilesearching.perimetersurveyswereterminatedafter20minutesor(rarely)whenthesitehadbeenthoroughlycircumnavigated,whichevercamefirst.Wescrutinizedtheland/waterinterfaceforadultamphibians,larvae,andeggmasses,andalsolookedunderlogsandotherpotentialcoverobjectsadjacenttothewetland.Werecordedthespeciesobserved,theapproximatenumberofindividualsofeachspecies,andthettFDforeachspecies. Analytical Methods.—toanalyzedaytimesurveydata,wefirstorga-nized species detectionsby year, park, site,method, and season.Wecombinedvisualencounteranddip-netsurveyresults fortheanalysis.Detectionsweredefinedasanobservationofaspeciesatagivensite,onagivenday,usingagivenmethod.Forexample,if20GreenFrogs(Lithobates clamitans)weredetectedbyadip-netsurveyatagivensiteonagivenday,itwasconsideredtobeonedetectionofthatspecies.IfGreenFrogswerealsodetectedduringthecallsurveyatthesamesiteonthesameday,itwasconsideredtobeaseparatedetection.thisapproachallowedustodeterminethespeciescompositionateachparkandsite,andtodeterminewhichspeciesweremostcommon(andmostcommonlydetected)amongsiteswithoutbiasingourresultstowardspeciesthatarelocallyabundantortowardaparticularsurveytechnique.Wecalculatedthepercentageofsurveysitesatwhicheachdetectedspecieswasfound(naïveoccupancy;mackenzieetal.2002). occasionally,weobservedspeciesatsitesoutsideoftheproscribedsur-veyperiods;thesespeciesdonotappearintheaboveanalyses.Wetherefore

AmphIbIAnsURVeYs

Fig. 2. Daytimesurveypointsfor2006and2007atsleepingbearDunesnationallakeshore,michigan.Greenindicatesparkland.

Fig. 3. Daytimesurveypointsfor2007atpicturedRocksnationallakeshore,michigan.DarkGreenindicatesparkland;lightgreenindicatesthenon-federalInlandbufferZone.

28 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 boWenAnDbeeVeR

table 1.Amphibianspeciesdetectionsbysite,method,andseasonatApostleIslandsnationallakeshore(Wisconsin)in2006.es=earlyspring,ls=latespring,s=summer.

Species Site Method Season

Anaxyrus americanus RockyIslandsouthswamp Dipnetsurvey ls michiganlagoon Dipnetsurvey s outerIslandlagoon perimetersurvey lsHyla versicolor littlesandbay Dipnetsurvey lsPseudacris crucifer littlesandbay Callsurvey es basswoodQuarry Callsurvey es,ls Dipnetsurvey es,s stocktonlagoonsouth Callsurvey es,ls Dipnetsurvey ls stocktonlagoonnorth Callsurvey ls RockyIslandsouthswamp Callsurvey ls sandRiver Callsurvey ls perimetersurvey ls michiganlagoon Callsurvey ls Dipnetsurvey ls,s outerIslandlagoon perimetersurvey ls outerIslandbeaverpond Callsurvey ls Dipnet ls perimetersurvey lsLithobates clamitans littlesandbay Callsurvey s Dipnetsurvey ls,s perimetersurvey s basswoodQuarry Callsurvey s Dipnetsurvey ls,s perimetersurvey s stocktonlagoonnorth Dipnetsurvey s sandRiver Dipnetsurvey ls,s perimetersurvey ls,s michiganlagoon Dipnetsurvey ls,s perimetersurvey s outerIslandbeaverpond Dipnetsurvey ls perimetersurvey ls outerlagoonnorth Dipnetsurvey s perimetersurvey sLithobates septentrionalis sandRiver Dipnetsurvey lsLithobates sylvatica basswoodQuarry Dipnetsurvey ls,s perimetersurvey ls RockyIslandsouthswamp Dipnetsurvey ls perimetersurvey es,ls michiganlagoon perimetersurvey lsAmbystoma maculatum basswoodQuarry Dipnetsurvey es,ls RockyIslandsouthswamp Dipnetsurvey es,ls michiganlagoon Dipnetsurvey ls,s stocktonlagoonsouth Dipnetsurvey s perimetersurvey s sandRiver Dipnetsurvey s outerIslandbeaverpond Dipnetsurvey ls perimetersurvey ls outerlagoonnorth perimetersurvey sNotophthalmus viridescens michiganlagoon Dipnetsurvey ls outerIslandbeaverpond Dipnetsurvey lsAmbystoma laterale RockyIslandsouthswamp perimetersurvey lsHemidactylium scutatum RockyIslandsouthswamp perimetersurvey ls


compiledaseparatelistofspeciesfoundoutsideofthesurveyperiodsateachsiteineachparkinordertoprovideamorecompleteaccountingofthespe-ciespresent.Weincludedanyobservedreptilianspeciesinthislist.Forallanalyses,commonandscientificnamesofspeciesfollowCrother(2008).

ResultsWewereunabletodeterminethenumberofdetectionsorthemost-com-monspecieswithregardtoseasonbecausenotallsiteswerevisitedinallthreeseasons,andsomesiteswerevisitedmultipletimesinoneseason.Furthermore,wewereunabletoquantitativelydeterminetheutilityofvari-ousmethodsbasedondetectionsbecausewewereunabletoperformallsurveysduringeveryvisitatallsites.Forthesesamereasons,naïveoccu-pancyestimatesandspeciesanddetectionspersiteshouldbeviewedwithcaution.however,quantityordiversityofdetectionsdifferedlittleamongmethodsorseasons,withthepossibleexceptionofsummerseasonsurveysresultinginlowerspeciesdiversity. Wedetectedatotalof10amphibianspeciesatApIs(Figs.4&5).outerIslandandmichiganIslandappearedtohavethemostdiverseher-petofaunas,withfiveamphibianandtworeptilianspeciesandsixamphib-ianandonereptilianspecies,respectively.Withregardtoindividualsam-plingsites,michiganlagoonandRockyIslandsouthswamphadthemostamphibianspeciesdetections(sixeach).basswoodQuarryproduced

thelargestnumberofamphibiandetections(13),followedbymichiganlagoon(11;table1).naïveoccupancyrangedfrom90%ofsites forthespringpeeper(Pseudacris crucifer)toonlyoneoftensitesforGraytreefrogs(Hyla versicolor),minkFrogs(Lithobates septentrionalis),blue-spottedsalamanders(Ambystoma laterale),andFour-toedsalamanders(Hemidactylium scutatum;table2).Wefoundpaintedturtles(Chrysemys picta)onouterIslandandstocktonIsland,andeasternGartersnakes(Thamnophis sirtalis)onouterIslandandmichiganIsland. Wedetectedatotalofnineamphibianspeciesandfourreptilianspe-ciesatslbeovertwoyears(tables3&4;Figs.6–8).In2006,ArallodgeandIndiantrailWesthadthemostamphibianspeciesdetections(fiveeach).Arallodgeproducedthelargestnumberofamphibiandetections(seven),followedbyIndiantrailWest(six;table3).naïveoccupancyrangedfrom85.7%ofsitesfortheGreenFrogtoonlyoneofsevensitesforGraytreefrogsandCentralnewts(Notophthalmus viridescens;table2).outsideofthesurveyperiods,weobservedGreenFrogsatArallodgeandnorthernleopardFrogs(Lithobates pipiens)atIndiantrailWest.Ifcombinedwithsurveydata,thisresultsinanaiveoccupancyof100%atdaytimesitesforGreenFrogsin2006.theonlyreptilianspeciesthatweobservedin2006weretheeasternGartersnakeatotterCreekandthenorthernRibbonsnake(Thamnophis sauritus)atArallodgeandIndiantraileast.

AmphIbIAnsURVeYs

Fig. 4. AnorthernleopardFrog(Lithobates pipiens)atlongIsland,ApostleIslandsnationallakeshore,Wisconsin.

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Fig. 5. stocktonIslandlagoonsouth,oneofthesamplingsitesatApostleIslandsnationallakeshore,Wisconsin.

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istable 2.numberofdaytimesamplingsites(inparentheses)atwhichamphibianspeciesweredetectedandpercentofsitesoccupiedbyamphibianspecies(naïveoccupancy)forApostleIslandsnationallakeshorein2006(ApIs,Wisconsin),picturedRocksnationallakeshorein2007(pIRo,michigan),andsleepingbearDunesnationallakeshorein2006and2007(slbe,michigan).

Anaxyrus Hyla Pseudacris Lithobates Lithobates Lithobates americanus versicolor crucifer clamitans pipiens septentrionalis

ApIs2006 (3)30.0 (1)10.0 (9)90.0 (7)70.0 (1)10.0slbe2006 (2)28.6 (1)14.3 (5)71.4 (6)85.7 slbe2007 (1)14.3 (5)71.4 (5)71.4 (6)85.7 (2)28.6pIRo2007 (3)42.9 (2)28.6 (5)71.4 (7)100.0 (1)14.3 (1)14.3 Lithobates Notophthalmus Plethodon Hemidactylium Ambystoma Ambystoma sylvatica viridescens cinereus scutatum laterale maculatum

ApIs2006 (3)30.0 (2)20.0 (1)10.0 (1)10.0 (7)70.0slbe2006 (2)28.6 (1)14.3 (2)28.6 slbe2007 (2)28.6 (1)14.3 (3)42.9 (1)14.3pIRo2007 (2)28.6 (1)14.3 (1)14.3


In2007,IndiantrailWesthadthemostamphibianspeciesdetec-tions(five),followedbyplatteRiver,Kelderhouse,andIndiantraileast(foureach).martinRoadproducedthelargestnumberofamphibiandetec-tions(14),followedbyIndiantrailWestandplatteRiver(12each;table4).however,detectionsatmartinRoadweredominatedbyGreenFrogsandspringpeepers.naïveoccupancyrangedfrom85.7%ofsitesfortheGreenFrogtoonlyoneofsevensitesforAmericantoads(Anaxyrus ameri-canus),spottedsalamanders(Ambystoma maculatum),andCentralnewts

(table2).WeobservedeasternGartersnakesatplatteRiver,Indiantraileast,andIndiantrailWestandnorthernRibbonsnakesatArallodge,Indiantraileast,andIndiantrailWest.Wealsoobservedpaintedturtles

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table 3.Amphibianspeciesdetectionsbysite,method,andseasonatsleepingbearDunesnationallakeshore(michigan)in2006.ls=latespring,s=summer.

Species Site Method Season

Anaxyrus americanus Arallodge Callsurvey ls Dipnetsurvey s IndiantrailWest Dipnetsurvey sHyla versicolor Arallodge Callsurvey ls Dipnetsurvey lsPseudacris crucifer platteRiver Callsurvey ls Arallodge Dipnetsurvey s IndiantrailWest Dipnetsurvey s Indiantraileast Dipnetsurvey s Kelderhouse Dipnetsurvey sLithobates clamitans otterCreek Callsurvey s Dipnetsurvey ls,s perimetersurvey ls,s platteRiver Dipnetsurvey ls,s perimetersurvey ls,s IndiantrailWest Callsurvey s perimetersurvey s Indiantraileast Callsurvey s perimetersurvey s Kelderhouse Callsurvey s Dipnetsurvey s perimetersurvey s martinRoad Callsurvey s perimetersurvey sLithobates sylvatica Arallodge Dipnetsurvey s Indiantraileast Dipnetsurvey sNotophthalmus IndiantrailWest Dipnetsurvey sviridescensPlethodon cinereus Arallodge perimetersurvey ls IndiantrailWest perimetersurvey s

Fig. 6. Indiantraileast,oneofthesamplingsitesatsleepingbearDunesnationallakeshore,michigan.

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Fig. 7. AnAmericantoad(Anaxyrus americanus)atsleepingbearDunesnationallakeshore,michigan.

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Fig. 8. AnabnormallypigmentedGreenFrog(Lithobates clamitans)larvafoundatKelderhouse,sleepingbearDunesnationallakeshore,michigan.

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atIndiantraileastandmartinRoadandsnappingturtles(Chelydra ser-pentina)atplatteRiver,IndiantrailWest,andmartinRoad. Wedetectedatotalofnineamphibianspeciesandonereptilianspe-ciesatpIRo(table5;Figs.9–11).northCountrytrailhadthemost

AmphIbIAnsURVeYs

table 4.Amphibianspeciesdetectionsbysite,method,andseasonatsleepingbearDunesnationallakeshore(michigan)in2007.es=earlyspring,ls=latespring,s=summer.

Species Site Method SeasonAnaxyrus americanus Kelderhouse Callsurvey lsHyla versicolor platteRiver Callsurvey ls perimetersurvey ls Arallodge perimetersurvey ls IndiantrailWest Callsurvey ls Dipnetsurvey ls perimetersurvey ls Indiantraileast Callsurvey es,ls Dipnetsurvey ls perimetersurvey ls martinRoad Callsurvey ls Dipnetsurvey es perimetersurvey lsPseudacris crucifer otterCreek Callsurvey es Arallodge Callsurvey ls Dipnetsurvey ls IndiantrailWest Dipnetsurvey ls Kelderhouse Callsurvey es,ls martinRoad Callsurvey es,ls Dipnetsurvey ls perimetersurvey lsLithobates clamitans otterCreek Dipnetsurvey es,ls,s perimetersurvey es,ls,s platteRiver Callsurvey s Dipnetsurvey es,ls,s perimetersurvey es,ls,s IndiantrailWest Callsurvey s Dipnetsurvey ls,s perimetersurvey ls,s Indiantraileast Callsurvey ls,s Dipnetsurvey ls,s Kelderhouse Callsurvey s Dipnetsurvey es,ls,s perimetersurvey ls,s martinRoad Callsurvey ls,s Dipnetsurvey es,ls,s perimetersurvey lsLithobates pipiens platteRiver perimetersurvey ls,s Indiantraileast Callsurvey esLithobates sylvatica platteRiver perimetersurvey ls Kelderhouse perimetersurvey esAmbystoma martinRoad perimetersurvey lsmaculatumNotophthalmus IndiantrailWest Dipnetsurvey lsviridescensPlethodon cinereus Arallodge perimetersurvey ls IndiantrailWest perimetersurvey es,ls Indiantraileast perimetersurvey ls

table 5.Amphibianspeciesdetectionsbysite,method,andseasonatpicturedRocksnationallakeshore(michigan)in2007.es=earlyspring,ls=latespring,s=summer.

Species Site Method SeasonAnaxyrus americanus ChapelRoadstream perimetersurvey ls ChapelRoadbeaverpond Callsurvey es northCountrytrail Callsurvey es Dipnetsurvey esHyla versicolor northCountrytrail Callsurvey es Dipnetsurvey ls ChapelRoadAlders Callsurvey esPseudacris crucifer 7-mileCreek Callsurvey es ChapelRoadstream Callsurvey es ChapelRoadbeaverpond Callsurvey es northCountrytrail Callsurvey es ChapelRoadAlders Callsurvey es Dipnetsurvey ls perimetersurvey esLithobates clamitans 7-mileCreek perimetersurvey s beaverbasinWest Callsurvey s perimetersurvey s beaverbasineast Dipnetsurvey s perimetersurvey s ChapelRoadstream Callsurvey s perimetersurvey es,ls ChapelRoadbeaverpond Callsurvey ls Dipnetsurvey es,ls perimetersurvey ls,s northCountrytrail Dipnetsurvey ls,s perimetersurvey ls ChapelRoadAlders Dipnetsurvey ls,sLithobates pipiens northCountrytrail Dipnetsurvey lsLithobates ChapelRoadbeaverpond Callsurvey sseptentrionalisLithobates sylvatica northCountrytrail Dipnetsurvey es perimetersurvey ls ChapelRoadAlders Callsurvey esNotophthalmus beaverbasinWest perimetersurvey sviridescens Plethodon cinereus beaverbasinWest perimetersurvey s

Fig. 9. AneasternGartersnake(Thamnophis sirtalis)atpicturedRocksnationallakeshore,michigan.

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The Eastern Collared Lizard (Crotaphytus collaris) was the subject of one of Dr. Fitch’s many contributions to better understanding the natural history of the Kansas herpetofauna (Fitch, H.S. 1956. An ecological study of the Collared Lizard (Crotaphytus collaris). University of Kansas Publications of the Museum of Natural History (8):213–274).

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proud sponsor of the IRCF and this centerfold

34 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 boWenAnDbeeVeR

amphibianspeciesdetections(six),followedbyChapelRoadAldersandChapelRoadbeaverpond(foureach).northCountrytrailproducedthelargestnumberofamphibiandetections(11),followedbyChapelRoadbeaverpond(eight;table5).naïveoccupancyrangedfrom100%ofsitesfortheGreenFrogtoonlyoneofsevensitesfornorthernleopardFrogs,minkFrogs,Centralnewts,andeasternRed-backedsalamanders(Plethodon cinereus;table2).outsideofthesurveyperiods,weobservedAmericantoadsatbeaverbasinWestandChapelRoadAlders.theonlyreptilianspeciesthatweobservedatpIRowastheeasternGartersnakeatnorthCountrytrail.

discussionWithregardtooverallnumberofspeciesdetected,daytimesurveyswereaneffectivewaytomonitoramphibians.morespeciesweredetectedusingthedifferentdaytimesurveymethodsthanbyusingnighttimecallsurveys(datanotshown).Inparticular,daytimesurveysresultedindetectionsofspeciesthatdonotcall,suchassalamanders.Daytimesurveysalsoallowedustodetermineifreproductionwasactuallyoccurringviadetectionsofeggmassesandlarvae,whereascallingmalesdonotnecessarilyequatewithsuccess-fulreproduction(heyeretal.1994).Atthistime,wecannotrecommendoneofthefourdaytimesamplingmethodsovertheothers.Dip-netsurveystendedtoproducemoredetections,butcallsurveysandperimetersurveysallowedustodetectspeciesthatwouldnothavebeennotedusingdip-netsurveysalone.ourdatasuggestthatchangingseasonsdoesnotchangetheeffectivenessofvisualencounteranddip-netsurveysfordetectingGreenFrogs(theonlyspecieswithenoughdetectionsforacomparison). slbewasthemostdiverseparkintermsofreptilianandamphib-ianspeciesdetected,followedbyApIsandpIRo.thisissensiblegiventhefactthatslbeisthesouthernmostparkthatwesampled.however,moreamphibianspeciesweredetectedatApIsthanateitheroftheotherparks.Anumberofspeciesatallthreeparksshouldhavebeenpresentbutwerenotdetected,suchasFowler’stoads(Anaxyrus fowleri)atslbe,Four-toedsalamandersatpIRo,andeasternRed-backedsalamandersatApIs(harding1997;Casper2001,2005;CasperandAnton2008).Insomecases,weknowthatthesespeciesarepresentbasedonpastsurveys.Regardless,wecannotstatewithconfidencethatanyspeciesisabsentwith-outmoresurveyingeffort(Kéry2002,mackenzie2005).notablespeciesdetectionsincludedtwonewrecordsforbasswoodIslandatApIs(GraytreefrogsandGreenFrogs;Casper2001).

Acknowledgementsthispilotworkwouldnothavebeenpossiblewithouttheadviceofandlogisticalsupportfromthenaturalresourcesstaffofthethreeparks.In

particular,weextendthankstos.YanchoandK.hydeatslbeandtoJ.belant,b.leutscher,andl.loopeatpIRo.s.mcmahon,e.ellis,t.VanZoeren,A.VanZoeren,andm.Cochranprovidedassistanceinthefield.Additionally,logisticalandprogrammaticsupportwasprovidedbytheU.s.nationalparkservice’sGreatlakesInventoryandmonitoringnetwork;inparticular,wearegratefultoW.Routeandt.KeniryforleadershipandtoU.Gafvert,whocreatedthemaps.portionsofthismanuscriptappearinanationalparkservicetechnicalreport.

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Casper,G.s.2001.Reptilesurveysoflong,michigan,andstocktonIslands,andlittlesandbay,intheApostleIslandsnationallakeshore,withnotesonamphibians.technicalReporttoApostleIslandsnationallakeshore,Wisconsin.

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Crother,b.I.(ed.).2008.Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico.herpetologicalCircular37.societyforthestudyofAmphibiansandReptiles,saltlakeCity,Utah.

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Gibbons,J.W.,C.t.Winne,D.e.scott,J.D.Willson,X.Glaudas,K.m.Andrews,b.D.todd,l.A.Fedewa,l.Wilkinson,R.n.tsaliagos,s.J.harper,J.l.Greene,t.D.tuberville,b.s.metts,m.e.Dorcas,J.p.nestor,C.A.Young,t.Akre,R.n.Reed,K.A.buhlmann,J.norman,D.A.Croshaw,C.hagen,andb.b.Rothermel.2006.Remarkableamphibianbiomassandabundanceinanisolatedwetland:Implicationsforwetlandconservation.Conservation Biology20:1457–1465.

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Fig. 10. AGreenFrog(Lithobates clamitans)atpicturedRocksnationallakeshore,michigan.

susa

n M

CM

aH

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Fig. 11. ChapelRoadstream,oneofthesamplingsitesatpicturedRocksnationallakeshore,michigan.

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oW

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macKenzie,D.I.2005.Whataretheissueswithpresence-absencedataforwildlifemanagers?Journal of Wildlife Management69:849–860.

macKenzie,D.I.,J.D.nichols,G.b.lachman,s.Droege,J.A.Royle,andC.A.langtimm.2002.estimatingsiteoccupancyrateswhendetectionprobabili-tiesarelessthanone.Ecology83:2248–2255.

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Route,b.2004.processandresultsofselectingandprioritizingVitalsignsfortheGreatlakesnetwork.Greatlakes Inventory andmonitoringnetwork.technicalReportGlKn/2004/05.nationalparkservice,Ashland,Wisconsin.

Route,b.andJ.elias.2007.Chapter1–backgroundinformation,pp.1–28.In:b.RouteandJ.elias(eds.),long-termecologicalmonitoringplan:Greatlakes

Inventoryandmonitoringnetwork.naturalResourceReportnps/GlKn/nRR/2007/001.nationalparkservice,FortCollins,Colorado.

semlitsch,R.D.2000.principlesformanagementofaquatic-breedingamphibians.Journal of Wildlife Management64:615–631.

stebbins,R.C.andn.W.Cohen.1995.A Natural History of Amphibians.princetonUniversitypress,princeton.

thoms,C.,C.C.Corkran,andD.h.olson.1997.basicamphibiansurveyforinventoryandmonitoringinlentichabitats,pp.35–46.In:D.h.olson,W.p.leonard,andR.b.bury(eds.),Sampling Amphibians in Lentic Habitats.societyfornorthwesternVertebratebiology,olympia,Washington.

Wake,D.b.1991.Decliningamphibianpopulations.Science253:860.

Weir,l.A. andm.J.mossman.2005.northAmericanAmphibianmonitoringprogram(nAAmp),pp.307–313.In:m.J.lannoo(ed.),Amphibian Declines: The Conservation of United States Species.UniversityofCaliforniapress,berkeley.

Welsh,h.h.ands.Droege.2001.AcaseforusingplethodontidsalamandersformonitoringbiodiversityandecosystemintegrityofnorthAmericanforests.Conservation Biology15:558–569.

male calling sites in Two species of australian Toadlets (anura: myobatrachidae: Uperoleia) at Two ponds in new south Wales

Francisl.lemckert1,2*,Georginalemckert2,Campbelllemckert2,andFrancesA.lemckert2

1schoolofenvironmentalandlifesciences,Universityofnewcastle,Callaghan,newsouthWales,Australia([email protected])2berowraheights,newsouthWales,Australia

Introduction“toadlets”ofthemyobatrachidgenusUperoleiaarecommonlyencoun-teredcallingaroundpondslocatedinsoutheasternAustralia.Usuallyonespeciesonlyispresentatanygivenpond,buttwospeciesoccasionallycanbecallingaroundthesamesite.thesmoothtoadlet(Uperoleialaevigata)andtheDuskytoadlet(U. fusca)callatthesametimeonthesamenights

aroundtwopondsonthecentralcoastofnewsouthWales,Australia,andobservationssuggestthatthemalescallinrelativelydiscretegroupsthatdiffertoatleastsomedegreeinlocation.thetwospeciesarecloselyrelated(tyleretal.1981),theadvertisementcallsofthetwospecies aresimilar(barkeretal.1995,Cogger2000),andtheypossesssimilarcallingseasonsandpreferredbreedingsites(barkeretal.1995,Cogger2000,Anstis2002).Undersuchcircumstancesthetwospeciesmaybeexpectedtoshowdif-ferencesintheircallsiteselection(littlejohnandmartin1969)thatwillassistindistinguishingthemalesofthetwospeciesbyfemalesmovingtothepondtobreed. Wecollecteddataonthecallingpositionsofthemalesofeachspeciestodetermineifthemaleswereselectingdifferentcallingareasortypesofcallingsites.Wecomparedlocationsofcallingmalesrelativetothepondsandalsomicrohabitatinformationassociatedwiththecallingpositionstodeterminewhatfeaturesthemalesofeachspeciesmightbeselectingforacallingsiteandhowmuchtheydiffer—ifatall.

Methodsthestudysiteconsistsoftwoadjacentponds(within5m)thatarelocatedapproximately15kmnorthwestofKulnuraonthecentralcoastofnsW,around120kmnorthofsydney(33°07’58.9”s,151°12’22.6”e).bothpondshavebeenpresentsinceatleastthelate1970sandareroughlycir-cularinshape.thesmallerpondisapproximately10mindiameterand0.3mdeep,andthelarger14mindiameterand0.9mdeep(depthsvarywithrainfall).AcallingmaleUperoleia fusca.


native vegetation in the surrounding area consists of dry openwoodland dominated by smooth-barked Apple (Angophora costata),Redbloodwood (Corymbia gummifera), andstringybarks (Eucalyptus oblonga),withasclerophyllousunderstory(ForestryCommission1989).Approximately25%ofthesurroundinglandhasbeenclearedforgrazingbylivestock. Wecollecteddataoncallingmalesonthenightof20February2006,locatingmalesofbothspeciesbytheircallsfrom2000–2300h.locationsofmalesweremarkedwithabambooskewercolor-codedforeachspe-cies.Wereturnedtothepondduringdaylighthoursandobtainedadigitalphotographfromaheightofonemeterofeachcallingsite,withtheskewerinplace.Werecordedthefollowingattributesofeachcallingsite,eitheratthetimeoftakingthephotographorfromthephotograph:(1)Distancefromtheedgeofthewatertothecallingsite(inmeters),(2)percentagebareground(rockorsoil)withina10-cmradiusofthecallsite(asopposedtobeingcoveredwithleaflitterorvegetation),and(3)percentageshadingof

thecallingsitewithina10-cmradiusofthecallsite.thismeasureprovidesanindicationofthelevelofcoveraffordedtothecallingmale. eachcategoryofdatawasinspectedvisuallytodetermineifitwasnor-mallydistributed.thiswasthecasefordistancefromthepond.percentageofbaregroundandpercentageofcoverwerearcsinetransformedtomeettestassumptions.Weusedt-testsinmicrosoftexcel2007tocomparethemeasurementsmadeforthetwospeciesusingp<0.05astheacceptedlevelofsignificance.

ResultsWecollecteddatafor19callingmalesofU. fuscaand16callingmalesofU. laevigata(table1).ComparisonsofthedatafromthecallsitesofthemalesindicatedthatthedistanceofthecallingsitesfromtheedgeofapondwassignificantlygreaterforU. laevigatathanforU. fusca(t=7.39;df=33;p<0.001).thepercentageofbaregroundwasgreateraroundthecallingsitesofU. laevigatathanatthoseofU. fusca(t=4.14;p<0.001).

lemCKeRtetAl

AcallingmaleUperoleia laevigata.

table 1.mean(±onestandarddeviation)andrangeofhabitatvariablesatsyntopiccallingsitesformalesofUperoleia fuscaandU. laevigata.

Species Uperoleia fusca (n = 19) Uperoleia laevigata (n = 16)Variable Mean (range) Mean (range)

Shading 18±24.7%(0–90%) 9±22.7%(0–100%)

Bare Ground 9±16.1%(0–70%) 39±26.4%(0–84%)

distance to Pond 2.1±2.9m(0.0–8.9m) 10.9±4.1m(1.9–18.2m)


themeanpercentageofcoverabovethecallingsiteswasgreaterforU. fuscamalesthanU. laevigata, butthedifferencewasnotstatisticallysignificant(t=1.62;p>0.10),althoughthisresultmighthavebeeninfluencedbyonemaleU. laevigatathatcalledfromunder100%cover.AnumberofU. laevigatamaleswereobservedcallingonpatchesofbaresand,whereasU. fusca malesalmostalwayschoselocationswheretheyrestedonleaflitterorvegetation.Uperoleiafuscamalesgenerallycalledfromundersomeformofvegetation,butalsocalledfrompositionspartiallyconcealedbyrocksordeadfall.

Discussionthemalesofthetwospeciesdoappeartohaveslightlydifferentpreferencesincallingsitelocation.maleU. fuscacallsignificantlyclosertotheedgeofthepondthandomaleU. laevigata.theyalsoprefersitesaffordedprotec-tionbysomecoverabovethecallingsite,whereasmaleU. laevigataoftencallfromexposedpositions. thepreferredcallingdistancefromthepondmayprovideasimplemeansofseparatingthetwospeciesatacommoncallingsite.bothspeciescallconsistentlyonthesamenightsoftheyear,oftenincombinedcho-rusesofmorethan40malesandsometimesover150males(F.l.lemckert,unpubl.data).Atthetimeofmaximumcallingactivity,asyntopicchorusisverycomplexandnoisy,andaspatialseparationofthemalesofthetwospecieswouldlikelybeofconsiderableassistancetofemalesattemptingtolocateconspecificmales. Uperoleiafuscamalesweremorelikelytocallfrompositionsthatwereatleastpartiallyobscuredbyvegetation.Wenotedthatthedensestveg-etationcoverwasclosetothepondandthemostopenareaswerefartherfromthepond.hence,therelativelocationofthehabitatsthatprovidethepreferredcallingsitesforthemalesmaybeenoughtoallowfortheobservedseparationofthetwospecies. theseparationofsitesmightalsobearesponsetocallingcompetition,aspredictedbylittlejohnandmartin(1969).theypredictedthatmales,inthepresenceofacousticcompetition,wouldchangetheircallingsitesorpatternstoreducethiscompetitionandavoidheterospecificmatings.Determiningthecharacteristicsofchosencallingsitesatpondswheremalesofonlyoneofthetwospeciescallwouldindicateifthemaleshavedifferentpreferredcallinglocationsintheabsenceoftheotherspecies.Ifnochangeinbehaviorisevident,thendatacouldbecollectedonthespatialstructureofsuitablegroundcovertotestifthedifferenceinstructuralelementsinflu-encesthechoiceofcallingsiteorwhethertheselectionofacallingsiteissimplyafunctionofdistancefromtheedgeofthepond.

AcknowledgementsWethankmikemahonyandRachaelpeakfortheirassistanceinformulat-ingthisstudyandmurraylittlejohnforcommentsonadraftofthismanu-script.thisworkwascarriedoutwithanappropriateanimalresearchlicenseobtainedfromtheDepartmentofenvironmentandClimateChangeandanAnimalWelfarelicensefromtheAnimalCareandethicsCommittee.

Literature CitedAnstis,m.2002.Tadpoles of South-eastern Australia: A Guide with Keys.Reed-new

holland,sydney.

barker,J.,G.C.Grigg,andm.J.tyler.1995.A Field Guide to Australian Frogs.surreybeatty&sons,sydney.

Cogger,h.G.2000.Reptiles and Amphibians of Australia. 6th ed.Reed-newholland,sydney.

ForestryCommission.1989.Forest Types in New South Wales.Researchnoteno.17.ForestryCommissionofnewsouthWales(nowForestsnsW),sydney.

halliday,t.andm.tejedo.Intrasexualselectionandalternativematingbehaviour,pp.419–468.In:h.heatwoleandb.K.sullivan(eds.),Amphibian Biology. Volume 2. Social Behaviour.surreybeattyandsonsptyltd,sydney.

littlejohn,m.J.andA.A.martin.1969.Acousticinteractionsbetweentwospeciesofleptodactylidfrogs.Animal Behaviour 17:785–791.

tyler,m.J.,m.Davies,andA.A.martin.1981.AustralianfrogsoftheleptodactylidgenusUperoleia Gray.Australian Journal of Zoology Supplementary Series79:1–64.

AUstRAlIAntoADlets

Viewofthestudyponds(left).malesofUperoleia fuscagenerallycallinthevegetationaroundtheedgesoftheponds,whereasmaleU. laevigatatendtocallonornexttotheroadonthelefthandsideoftheimage.ontherightisthelargerofthetwoadjacentpondsshowingtheareasofvegetationonthebanksaswellasareasofbarebankfromwhichthemalescall.

AmplexusinUperoleia fusca.

38 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 CAmposAnDmAGnUsson

browntreeClimbers(Uranoscodon superciliosuslinnaeus1758),knownlocallyastamacoré,occurthroughouttheAmazonbasin,andoften

arefoundalongtheedgesofwatercoursesperchedonbranchesoftreeandvines(Fig.1;Vittetal.2008).theselizardsalsoarecalled“Divinglizards”becausetheydiveintothewatertoescapedanger.theyaremodelsit-and-waitpredators.thediethasbeenstudiedinbrazilbyhoowlandetal.(1990),Vittetal.(1991),andGasnieretal.(1994)andinsurinambyhoogmoed(1973).thespeciesspendsmostofitstimeontree-trunks,butfeedsmainlyonpreythatoccurexclusivelyormuchmoreabundantlyontheground(Gasnieretal.1994).themainpreyitemsareorthopterans,cockroaches,earthworms,smallfrogs,andlepidopteranlarvae.thehigherconsumptionoflarvaewascoincidentwithorsoonafterovipositioninJuly–november(Gasnieretal.1997). here,wereportillegalhuntingofU. superciliosus inthebrazilianAmazon. on13August2008,nearCururulakenearthesolimõesRiver(3°34’30.4”s,60o40’03.3”W;datum:WGs84;elev.80m),stateofAmazonas,alocalresidentcapturedandkilled18individualjuvenilesandadultU. superciliosus(Fig.2).lizardsontreebranchesandvinesinfloodedforest(igapó)aroundCururulakewerecapturedbyhand.Abuyerhadordered30individuals.eachanimalwasevisceratedandstoredinalcohol.thefollowingday,thebuyerwenttothehunter’shouseandpaidR$1.00(aboutUs$0.60)perlizard.hesaidthatthelizardsweretobeusedinanUmbanda(anAfro-brazilianreligionthatblendsAfricanreligionswithCatholicism)ritualaspartofatreatmentformalesexualinadequacy. thespeciesalso iscommonlyfoundinthemarket inthecityofbelém,stateofthepará,andsuchritualsmightbepartofthepopularcultureinmuchofAmazonia.Allwildlifehasbeenprotectedinbrazilsince1967,exceptthattakenforsubsistence—andtradeforaphrodisiacsisunli-kelytobeconsideredsubsistence.Althoughthespecieshasanenormousrange,lizardsarecapturedduringthereproductiveperiod,andcollectioncouldaffectlocalpopulations,especiallyinareaswheremuchoftheriparianvegetationhasbeencleared.

Literature CitedGasnier,t.R.,W.e.magnusson,andA.p.lima.1994.Foragingactivityanddietoffour

Fig. 1. brown tree Climbers (Uranoscodon superciliosus) occur throughoutAmazonia,oftenalongwatercoursesperchedonbranchesoftreesandvines.

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J. V

itt

Fig. 2.Uranoscodon superciliosus killedbyamarkethunteratCururulake,brazilianAmazon,August2008.

ziLC

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Hunting Tamacoré, Uranoscodon superciliosusZilcaCampos1andWilliammagnusson2

1embrapapantanal,Cp109Corumbá,msbrazil79320-900,brazil([email protected])2InpA,Coordenaçãodepesquisasemecologia,Cp478,69069-020manausAm,brazil([email protected])

IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 39AllIGAtoRsnAppeR

sympatriclizardspeciesinatropicalrainforest.Journal of Herpetology28:187–192.

Gasnier,t.R.,W.e.magnusson,andA.Waichman.1997.GrowthcurveshapeandgrowthvariationofthetropicallizardUranoscodon superciliosus(sauria:tropiduridae).Ecotropica3:101–107.

hoogmoed,m.s.1973.notesontheherpetofaunaofsurinam.IV.thelizardsandamphisbaeniansofsurinam.Biogeographica4:1–419.

howland,J.m.,l.J.Vitt,andp.t.lopez.1990.lifeattheedge:theecologyandlifehistoryofthetropidurineiguanidlizardUranoscodon superciliosum.Canadian Journal of Zoology68:1366–1373.

Vitt,l.,W.e.magnusson,t.C.Avila-pires,andA.p.lima.2008.Guide to the Lizards of Reserva Adolpho Ducke, Cntral Amazonia.Guia de Lagartos da Reserva Adolpho Ducke, Amazônia Central.ÁttemaDesigneditorial,manaus,brazil.

alligator snapper stuck in the mud: evidence of aestivation

stevenG.George

U.s.ArmyCorpsofengineersengineerResearchandDevelopmentCenter,environmentallaboratory

Vicksburg,mississippi39180([email protected])

theAlligatorsnappingturtle(Macrochelys temminckii)isthelargestfreshwaterturtleinnorthAmerica,capableofreachingaweightof

113kgwithacarapacelengthof80.0cm(ernstetal.1994).theseturtlesareconfinedtoriversystemsofthelowermississippibasinandriversthatdrainthenorthernGulfofmexico(ernstetal.1994,trauthetal.2004).theAlligatorsnappingturtleishighlyaquatic,andonlythefemaleleavesthewatertonest(ernstetal.1994,pritchard1989).littleisknownofthebehaviorofthisspeciesundernaturalconditions,especiallyduringlowwaterwhenindividualsmightbecomestranded. on3september1993,whileworkingonpantherCreek (YazooCounty,mississippi)duringlow-waterconditions,IobservedanAlligatorsnappingturtlethatappearedtobeaestivatingwhileburieddeeplyinmud.Afterencounteringtheturtle,mycoworkerandIreturnedthefollow-ingdaytotakepicturesandcollectdata.thesedimentdepthoftheburiedturtlewas35.6cm,andtheturtlehadmoved17.8cmsincefirstobserved16hearlier.thesedimentsurroundingtheturtlewassoftmudthatcon-tainedseepagefromaspring.thetemperatureoftheseepagewas22ºC.themaximumcarapacelengthoftheturtlewas66cm.theindividualwasamaleandweighedjustover45.3kg. on24october2003,whilesamplingfishes intheQuiverRiverupstreamfromhwy3insunflowerCounty,mississippi,acoworkerandIobservedasecondAlligatorsnapperstrandedinmud.onlytheupperhalfoftheturtle’sbodywasexposed.Apparentlytheturtlehadbeenstrandedduringlow-waterconditionsandwaswaitingforariseintherivertoiniti-atemovement.ernstetal.(http://nlbif.eti.uva.nl/bis/turtles.php)reportedturtlesaestivatingindryingriverbedstopreventdesiccation.AlthoughothercolleaguesalsohaveobservedAlligatorsnappingturtlesstrandedoraestivatingduringlowwaterconditions(billlancaster,retiredturtletrap-per,sunflowerCounty,mississippi;brentharrel,UsFWs,pers.comm.),thisaccountappearstobethefirstpublisheddocumentationofMacrochelys temminckiistrandedoraestivatinginnaturalhabitat.

AcknowledgmentsWilliamt.slackreviewedthismanuscriptandbradleylewisandJayCollinsprovidedfieldassistance.WorkconductedinthemississippiDeltawassup-portedbytheVicksburgDistrict,U.s.ArmyCorpsofengineers.permissionwasgrantedbytheChiefofengineerstopublishthisinformation.

Literature Citedernst,C.h.,J.e.lovich,andR.W.barbour.1994.Turtles of the United States and

Canada.smithsonianInstitutionpress,Washington,D.C.

ernst,C.h.,R.G.m.AltenburgandR.W.barbour.turtlesoftheWorldonlineDatasource(http://nlbif.eti.uva.nl/bis/turtles.php).

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

trauths.e.,h.W.Robison, andm.V.plummer2004.The Amphibians and Reptiles of Arkansas.theUniversityofArkansaspress,Fayetteville.

excavatinganAlligatorsnappingturtlefromthemudduringlow-waterconditionsinpantherCreek,YazooCounty,mississippi.

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40 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 GeoRGe

Timber rattlesnake (Crotalus horridus) swims the mississippi river

stevenG.George

U.s.ArmyCorpsofengineersengineerResearchandDevelopmentCenter,environmentallaboratory

Vicksburg,mississippi39180([email protected])

thetimberRattlesnake (Crotalus horridus) iswidelybut sporadi-callydistributedthroughoutmuchofthesoutheasternUnitedstates

(trauthetal.2004,ConantandCollins1991).numerouslifehistorystudieshavebeenconductedonthespecies,addressingtopicsthatincludereproduction,migration,andmovement(e.g.,palmeretal.1995,trauthetal.2004);however,reportsofswimmingbehaviorarefew.Walker(1963)observedanindividualswimmingacrossalakeinJacksonparish,louisiana.Viosca(1944)commentedthatthemississippiRiveranditsfloodplainpro-videapotentialbarrierforeast-westmovementforvariousamphibiansandreptiles;however,themississippiRiveritselfmightprovetobeanexcellentcorridorfordispersal.

on19July2002,whileworkingonthemississippiRiverjustsouthofVicksburg,mississippiatRiverKilometer685.6(Rm426),ourfieldcrewandIobservedatimberRattlesnakeswimmingacrosstheriverpresumablyfrommadisonparish,louisianatoWarrenCounty,mississippi.thestreamwidthwheretheswimmingrattlesnakewasobservedwas750m,andthewatertemperaturewas29.5ºC.thesnakewasoverhalfwayacrosstheriverandapproximately300mfromthenearestshore.Itlookedtobeinexcellentcon-ditionandappearedquitecapableofcompletingitsjourneyacrosstheriver. Interestingly,Dr.Davidbiedenharn(UsACeCoastalandhydraulicslaboratory,eRDC;pers.comm.)alsoobservedatimberRattlesnakeswimmingthemississippiRiverintheVicksburgareaduringthesummer,althoughtheeventtookplaceseveralyearsago.Inaddition,whilework-ingontheAlabamaRiverduringlatesummer2004,IobservedtimberRattlesnakesontwoseparateoccasionsswimmingtherivernearCamden,Alabama.theseobservationsprovidesupportforthecontentionthatlargeriversarenotamajorbarriertotimberRattlesnakedispersal.

AcknowledgmentsspecialthankstoWilliamlancasterandbradleylewisforfieldassistanceandphotographs,andtoWilliamt.slackforreviewingthemanuscript.WorkconductedonthemississippiRiverwassupportedbythemississippiValleyDivision,U.s.ArmyCorpsofengineers.permissiontopublishwasgrantedbytheChiefofengineers.

Literature CitedConant,R.andJ.t.Collins.1991.A Field Guide to Reptiles and Amphibians of

Eastern and Central North America.3rded.houghtonmifflinCo.,boston.

palmer,W.m.andA.l.braswell.1995.Reptiles of North Carolina.UniversityofnorthCarolinapress,Chapelhill.

trauths.e.,h.W.Robison,andm.V.plummer.2004.The Amphibians and AtimberRattlesnakeswimmingandfloatingwiththecurrentsofthemississippiRiverapproximately300mfromland.

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the author and a timber Rattlesnake that had been swimming across themississippiRiverfrommadisonparish,louisianatoWarrenCounty,mississippi.

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AtimberRattlesnakeswimmingtheAlabamaRiver,CamdenCounty,Alabamain2004.

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a Defensive Display by a smooth earth snake (Virginia valeriae)

Andrewm.Durso1andKevinp.Durso2

1odumschoolofecology,UniversityofGeorgia,Athens,Georgia30602([email protected])2CollegeofnaturalResources,northCarolinastateUniversity,Raleigh,northCarolina27607([email protected])

thesmoothearthsnake(Virginia valeriae)isasmall(maximumsize393mm)andubiquitousnatricinesnakeofthesoutheasternUnited

states.Whereasmanylargenatricinesnakeswilldisplaydefensivelywhenunabletoretreat,gaping,vibratingthetail,andstrikingatwould-bepreda-tors,thesebehaviorsarelesscommonlyobservedinsmallsnakes,possiblyduetotheirnon-threateningandinconsequentialeffectonlargepredators.bothVirginia valeriae andRed-belliedsnakes(Storeria occipitomaculata)areknowninsteadtoexhibit“lip-curling,”anunusualandelaborateexhibitionofthemaxillaryteeth,whichishypothesizedtoconstituteanagonisticdis-playtargetedatsmallerpredatorsandadeliveryagentforDuvernoy’sglandsecretions(doAmaral1999,Conant1951). At1100hon24July2008,weobservedadefensivedisplayinanadultV. valeriaewhilephotographingtheanimalinsuburbandeciduouswoodlandsnearthesitewhereitwascapturedatnighttwodaysearlier.thesnakehadbeenmovingacrossthepavedwalkwayofaprivateresidenceinWakeCounty,northCarolinaat0015hafteralightrainandwasheldforlaterphotographyduetopoorlightconditionsduringtheinterveningtwodays.Immediatelyuponbeingremovedfromitstemporarycontainerandplacedunderlightrestraint,thesnakefacedusandgapedinastrikingposi-tion,butdidnotstrike.thisbehaviorcontinuedforapproximately30sec,afterwhichthesnakechangeditsbehaviorandinsteadattemptedtoescapeforapproximately2min.theinitialescapebehaviorwasthenfollowedbyapproximately2minofgaping,slightneck-flaring,andactualstrikingbefore

thesnakeresortedentirelytoescapeandburrowingbehaviors,whichitcon-tinuedaswephotographeditforalmost20mindespitebeingremovedfromhospitableburrowinghabitatandplacedonafirm,opensurface. thisaccountofdefensivebehaviorinV. valeriaeisstrikinglysimilartothatoftodd(2008)foranindividualfrombarnwellCounty,southCarolina.Inbothinstances,thebehaviormighthavebeenanatypicaldis-playbyahighlyagitatedindividualoritcouldbeatraitthatismorecom-monthanpreviouslythought.otherdefensivebehaviorsdocumentedforV. valeriaeincludewrithingviolently,sprayingmusk,voidingfeces,feign-ingdeath(ernstandernst2003),andloop-knottingthebodytopreventingestion(Yeatman1983).

Literature CitedConant,R.1951.The Reptiles of Ohio.2nded.andrevisionaryaddenda,University

ofnotreDamepress,notreDame,Indiana.

doAmaral,J.p.s.1999.lip-curlinginRedbellysnakes(Storeria occipitomacu-lata):Functionalmorphologyandecologicalsignificance.Journal of Zoology: Proceedings of the Zoological Society of London248:289–293.

ernst, C.h. and e.m. ernst. 2003. Snakes of the United States and Canada.smithsonianInstitutionpress,Washington,D.C.

todd, b.D. 2008. Virginia valeriae (smooth earth snake). threat display.Herpetological Review39:475.

Yeatman,h.C.1983.Virginia v. valeriae(easternsmoothearthsnake).Defense.Herpetological Review14:22.

Asmoothearthsnake(Virginia valeriae)exhibitingaprolongeddefensivedisplay(seetext).

VIRGINIA VALERIAE

Reptiles of Arkansas.theUniversityofArkansaspress,Fayetteville.

Viosca,p.,Jr.1944.Distributionofcertaincoldbloodedanimalsinlouisianainrelationshiptothegeologyandphysiographyofthestate.Proceedings of the

Louisiana Academy of Sciences8:47–62.

Walker J.m. 1963. Amphibians and reptiles of Jackson parish, louisiana.Proceedings of the Louisiana Academy of Sciences26:91–101.

42 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 DelIsle

sharing the resource: six species of rattlesnakes in

Joshua Tree national park, californiaharoldF.Delisle

Joshuatreenationalpark,twentyninepalms,California92277([email protected])

photographsbytheauthorexceptwherenoted.

thecompetitiveexclusionprinciplewouldseemtoapplytosixspeciesofrattlesnakesinthegenusCrotalus,allofwhichfeedmostlyonsmallmammals.InJoshuatreenationalpark,California,however,sixspeciesoccurinanareaofjust400,000ha.Apatternnotedinecologyisthatdiversityatonelevelbegetsdiversityatotherlevels.Almost70yearsoflocalitydatacombinedwithpresentfieldresearchwasusedasevidenceforthehypothesisthattheserattlesnakespeciesmostlyavoidcompetitiveexclusionbymicrohabitatdifferenceswithinthegreatdiversityofecologicalcommunitiesinthepark.theselong-termrecordsmightalsoindicatethatthedynamicsofthedesertecosystemcouldbechanging,possiblytheresultofclimatechangeand/orlocalurbanization.

Introduction

ecologicaltheorypredictsthatspeciesinecologicalcommunitiescancoexistonlyiftheydifferintheirresponsestolimitingresources.this

competitiveexclusionprinciplewouldseemtoapplytothesixspeciesofrattlesnakesinthesinglegenusCrotalus inthearea(400,000ha)ofJoshuatreenationalpark(Jtnp).Adultrattlesnakesfeedalmostexclusivelyonsmallmammals(mackessy1988,beavers1976),withtheexceptionofsidewinders(C. cerastes),forwhichmammalscompriseabout50%ofthediet(Funk1965).InJtnp,theubiquitousmerriam’sKangarooRat(Dipodomys merriami)probablymakesupthebulkofthedietofallsixspe-cies,asitdoeselsewhereinthedesert(Reynoldsandscott1982),whereasside-blotchedlizards(Uta stansburiana)almostcertainlyservethesameroleforhatchlingrattlesnakes.howdoesthislimitedresource supportsomanycloselyrelatedspecieswithoutviolatingecologicaltheory?

Materials and MethodsJoshuatreenationalparkisanidealplacetostudyrattlesnakeecology.Itsprotectedstatushaspreservednaturalhabitatsinfairlygoodconditionfor70years.thefirstherpetologicalsurveyofwhatwasthenanationalmonu-mentwasconductedbyRobertC.stebbinsfrom1945–1955.R.b.loomisandhiscolleaguescontinuedtoinventorythereptilesduringthe1960s.Ibeganperiodicherpetologicalvisitstotheparkinthe1970s,andcon-ductedseason-long(march–november)surveysin1999–2005.Inaddition,observationrecordsmadebyrangersandothershavebeenaccumulatingforalmost50years.theresult(table1)isthatwehaveaccumulatedsome400localitydatareportsonWesternDiamondbackRattlesnakes(C. atrox;18),southernpacificRattlesnakes(C. helleri;60),RedDiamondRattlesnakes(C. ruber; 15),mojaveRattlesnakes (C. scutulatus; 21), southwesternspeckledRattlesnakes(C. pyrrhus;155),andsidewinders(C. cerastes;133). All localitydatawereplottedon amapof theparkusingesRIArcView.thesespeciesmapswerethencomparedwithmapsshowingvegetationpatterns(leary1977).extensivefieldworkwasconductedinareasofsympatrytoidentifypossibledifferencesinmicrohabitatswithinleary’shabitat(vegetationassociation)types.thesedatawereusedtotestthehypothesisthattherattlesnakesintheparkavoidcompetitiveexclusionprimarilybypartitioningmicrohabitats. multivariateanalysesofhabitatswerenotemployed,butworkelse-where(Reinert1984a,1984b)indicatedthathabitatseparationisprobably

thechiefmethodforsnakesthatpreyonsmallmammalstoavoidnichecompetition.because“habitat”isaverybroadcategorythatcanincludea

theubiquitousmerriam’sKangarooRat(Dipodomys merriami)istakenbylargerrattlesnakesofallspecies,whereashatchlingsfeedlargelyonthephenomenallyabundantside-blotchedlizards(Uta stansburiana).

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varietyofdissimilarentities,approachingtheconceptfromasnake’sper-spective,ratherthanourown,isimportant.everyindividualhasto:(1)Gettoaplace(orbebornthere),(2)beabletoexploitallessentialresources,and(3)avoiddeleteriousconditions,whichmightbeepisodicbutcanbesevere.thesefactorsaredistributedonseveralscales,from<1mtomanykm2.

locationofJoshuatreenationalparkandlocalecoregions(thesmallsquareindi-catestheIndianCovearea).

theecologicalcommunitiesinJoshuatreenationalpark.

theIndianCovearea,withlocalityrecordsforCrotalus cerastes (green),C. helleri (darkblue),C. pyrrhus (magenta),C. ruber (lightblue),andC. scutulatus (yellow).

WesternDiamondbackRattlesnakes(Crotalus atrox)areconfinedmostlytothesonoranDesertecoregion.

sidewinders(Crotalus cerastes)appeartobelimitedtoareaswithsandysubstrates.

southernpacificRattlesnakes (Crotalus helleri) inhabit rockyareaswithin thepinyon-Junipercommunity.

ResultsCrotalus atrox reaches the northern-most point of its distribution inCaliforniawithinJtnp. It ismostlyconfined to thesonoranDesertecoregionsection.Crotalus cerastes seemsstrictlylimitedtoareasofsandsubstrate.Crotalus scutulatus reachesthesouthern-mostextentofitsrangeinCalifornia in thenorthwesternpartofJtnp.Itappears limited inoccurrencetoJoshuatreeflats.Crotalus helleriinhabitsrockyareaswithinthepinyon-Junipercommunity.AlllocalityrecordsarewithinareaswithCaliforniaJunipers(Juniperus californicus).Crotalus ruber occursintwoapparentlydisjunctpopulationsinJtnp.one,inthesouthwesterncornerofthepark,isprobablytheeasternterminusofthemainpopulationofthedesertfoothills(e.g.,WhitewaterCanyon).thesecondhaslongbeencon-

sideredanisolatedpopulationwithintheWonderlandofRocks,whereitisassociatedwithlargeoutcroppingsofmonzograniteinveryruggedterrain.Crotalus pyrrhus isthemostabundantspeciesofrattlesnakeinJtnp.Intheeasternhalfofthepark,itistheonlyspeciesoccurringinrockycanyons,hills,andridges.Inthewesternhalfoftheparkitseemstohaveanarrowerniche,andappearstobelimitedtorockycanyons.Itdoesoccurtoeleva-tionsashighas1,525mabovesealevel. most specieswereclearlyassociatedwithparticularhabitat types.however,thelocalitydatarevealedoneareaoftheparkinwhichfivespeciesofCrotalus(cerastes, ruber, helleri, pyrrhus, scutulatus)appeartobesympatric.thisareacentersaroundIndianCove.Groundreconnaissanceofthelocali-tiesofsympatryrevealedthatthespeciesarenotsyntopic.

44 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 DelIsle

southwesternspeckledRattlesnakes(Crotalus pyrrhus)occurinrockycanyons,hills,andridgesintheeasternhalfofthepark,butappeartobelimitedtorockycanyonsinthewesternhalf.

table 1.basichabitattypesandlocalityrecordsbyspecies.

Habitat types Species of Crotalus

C. atrox C. cerastes C. helleri C. pyrrhus C. ruber C. scutulatusRockycanyonsW 20 19 45 10 14Rockycanyonse 80 31 RockyridgesW(pinyon-juniper) 81 4 90Rockyridgese 1 JoshuaflatsW 11 86Creosoteflatse 8 sandywashes/flatsW 10 sandywashes/flatse 90

RedDiamondRattlesnakes(Crotalus ruber)occurinthesouthwesterncorneroftheparkandanisolatedpopulationisknownfromtheWonderlandofRocks.

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DiscussionResourcespotentiallysubjecttointerspecificcompetitionarefood,place,andtime.localityrecordsandfieldstudiesseemtosupportthehypothesisthatnichesaredistinctatthemicrohabitatlevel.FortherattlesnakesofJtnp,shelterisprobablyinmorethanadequatesupply,withholesandcrevicesessentiallyeverywhere.nocturnal-diurnalbehaviorisdictatedby

seasonaltemperaturesandappearstobethesameforallspeciesatthesamealtitude.Veryfewquantitativestudiesofthedietsofrattlesnakeshavebeenconducted.thefewexistingdietarydataforthesespeciesweredeterminedelsewhereintheirrange.thattheyareessentiallythesameinJtnpmayormaynotbeavalidassumption,asdietaryshiftsareknowntooccurinwide-rangingspecies. therattlesnakedistributionintheparkmaybechangingalongwithecosystemchanges.thelastverifiedsightingintheparkofC. scutulatus wasin1991(KeysRanch).thetwoflatlandcorridorsthatleadfromthenorth(mainrangeofthemojaveRattlesnake)arenowlargelyblockedbyurban-ization.Iftheparkwasformerlya“sink”formovementsofthisspecies,thecorridorsarenowmostlyblocked,andthespeciesmayhavebecomeextirpatedfromthepark,althoughitremainsabundant25kmtothenorth,ontheothersideoftheurbandevelopment. Inthelasttwodecadesexoticgrasses(mostlyCheatGrasses,Bromus spp.)haveinvadedmuchoftheJuniperandJoshuatreewoodlands.theseexoticshaveprovidedfuelforrecurringwildfiresinplantcommunitiesnotadaptedtofire.Anintensivesurveyafterthemay1999fireshowedthatmostreptilessurvivedthefire,butasubsequentsurveyduringthefollowingspringindicatedaneartotalabsenceofallspeciesintheburnedarea.

AcknowledgementsIthankhenrymcCutcheon,ChiefofResourcemanagement,Jtnp,forhisencouragementandthenecessarypermitstodoresearchinthepark,andespeciallytoddhoggan,sanbernardinoCountymuseum,forhishelpwithfieldwork.theleeFamilyFoundationandtheJoshuatreenationalparkAssociationprovidedfunding.

Literature Citedbeavers,R.A.1976.FoodhabitsoftheWesternDiamondbackRattlesnake,Crotalus

atrox (Viperidae) intexas.Southwestern Naturalist20:503–515.

Delisle,h.F.2003.Amphibiansand reptilesof Joshuatreenationalpark:Annotatedchecklist.Unpubl.Reportforthenationalparkservice,Joshuatreenationalpark,California.

Funk,R.s.1965.FoodofCrotalus cerastes laterorepens inYumaCounty,Arizona.Herpetologica21:15–17.

Geest,J.C.1971.studiesofapopulationofsidewinderRattlesnakes,Crotalus ceras-tes hallowell,fromthesuggestedareaofintergradation.Unpubl.Reportforthenationalparkservice,Joshuatreenationalpark,California.

leary,p.J.1977.Investigation of the Vegetation Communities of Joshua Tree National Monument.technicalReport2.nationalparkserviceandtheUniversityofnevada,lasVegas.

loomis,R.b. andR.C.stephens.1962.Recordsof snakes fromJoshuatreenationalmonument,California.Bulletin of the Southern California Academy of Sciences61:29–36.

loomis,R.b.andR.C.stephens.1967.AdditionalnotesonsnakestakeninandnearJoshuatreenationalmonument,California.Bulletin of the Southern California Academy of Sciences66:1–21.

mackessy,s.p.1988.Venomontogenyinthepacificrattlesnakes,Crotalus viridis helleri andC. v. oreganus.Copeia1988:92–101.

miller,A.h.andR.Cstebbins.1964.The Lives of Desert Animals.UniversityofCaliforniapress,berkeley.

Rainey,D.1962.AmphibiansandreptilesofCovingtonFlat,Joshuatreenationalmonument.Unpubl.Report for thenationalparkservice, Joshuatreenationalpark,California.

Rainey,D.1965.thedistributionofamphibians,reptiles,andmammalsofJoshuatreenationalmonument.Unpubl.Reportforthenationalparkservice,Joshuatreenationalpark,California.

Reinert,h.K.1984a.habitatseparationbetweensympatricsnakepopulations.Ecology65:678–486.

Reinert,h.K.1984b.habitatvariationwithinsympatricsnakepopulations.Ecology65:1673–1682.

Reynolds,R.p. andn.J. scott, Jr. 1982.Useof amammalian resourceby aChihuahuan snake community, pp. 99–118. In: n.J. scott, Jr. (ed.),Herpetological Communities.UnitedstatesDepartmentoftheInteriorFishandWildlifeserviceResearchReport13,Washington,D.C.

mojaveRattlesnakes(Crotalus scutulatus)reachthesouthern-mostextentoftheirrangeinCaliforniainthenorthwesternpartofJtnp.thespeciesappearstobelimitedinoccurrencetoJoshuatreeflats.

theIndianCovearea,withthetopographicmap(top)andphotographshowingthesameview.bluedotsindicatehabitatofCrotalus helleri,greendotsthatofC. cerastes,andmagentadotsthatofC. pyrrhus.

46 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 smIthAnDlemos-espInAl

observations of amplexus and oviposition in Ollotis [Bufo] occidentalis in the

río salado, puebla, mexicoGeoffreyR.smith1andJulioA.lemos-espinal2

1Departmentofbiology,DenisonUniversity,Granville,oh43023,UsA([email protected])2laboratoriodeecología,Unidaddebiología,tecnologíayprototipos(Fes-Iztacala,UnAm),Av.delosbarrioss/n,

losReyesIztacala,tlalnepantla,estadodeméxico,méxico54090

theRíosaladorunsthroughelValledeZapotitlánsalinasinsouth-easternpuebla,mexico.poolsalongtheRíosaladoareseasonal,form-

ingduringthedryseasonasthewaterlevelintheRíosaladofalls.DuringamonthlysurveyofasectionoftheRíosaladofortadpoles(Woolrich-peñaetal.,unpubl.data),weobservedanamplectantpairofOllotis [Bufo] occi-dentalisandtheireggmass.theonlypreviouslypublishedreportsonrepro-ductioninO. occidentalisareanobservationbyDuellman(1961)oftad-polescollectedon“may3inaquietsectionofafaststreamnearbarrancaseca”inmichoacán,mexico,andanobservationofanamplectantpairin

elValledeZapotitlánsalinason27october1998thatproducedaneggmasswith>10,000eggs(oliver-lopezetal.2000). Weobservedtheamplectantpairon24november2007at0745h.Indicationsthatithadrainedthepreviousnightwereabundant.theeggmasswasdepositedinashallowpoolalongthemainchanneloftheriver.thedepthofthewaterwas2cm.theconductivityofthewaterwas1,966mscm-1,thesalinitywas1.2ppt,thedissolvedoxygencontentwas6.49mgl-1,andthetemperaturewas17.0°C(measuredusingaYsImodel85handheldDo/conductivitymeter).Asurveyofseveralotherpoolsalonga

AmplectantpairofOllotis [Bufo]occidentalisandeggmassinapoolalongtheRíosaladoinpuebla,mexicoon24november2007.notetheshallownessofthewater.

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2-kmsectionoftheriverrevealednoothereggmasses,amplectanttoads,oranyotheradulttoads,althoughweencounteredseveraltadpolesofvaryingsizesthroughoutthissectionoftheriver. ourobservationsuggests that, intheRíosalado, thesetoads laytheireggsinrelativelyshallowpools.DuringtheperiodwhentadpolesarepresentintheRíosalado(november–February),poolsremainrela-tivelyshallow;however,tadpolesaremorefrequentlyfoundindeeperandlargerpools,andarefoundonlyinpoolsalongthemainchanneloftheRíosalado(Woolrich-piñaetal.,unpubl.data).Giventherelativelyshallownatureofthepoolinwhichweobservedtheamplectantpair,choicesofovipositionsitesbytheadultsmightbelimitedbeyondlayingeggsinthemainchanneloftheriver,andtadpoleslikelymoveamongpoolsuntiltheriverdriestherebyisolatingmanyofthepools.however,furtherstudythatmoresystematicallyexaminestheovipositionsiteselectionofthesetoadswouldbeinformativeandmightprovidevaluableinformationtoguideanyassessmentofconservationormanagementplansoftheRíosalado,whichispotentiallyaffectedbyhuman-alterations,includingtheuseofwaterfromtheriverfortheproductionofsaltin“salineras.”

Acknowledgementsthisresearchwassupportedby funds fromtheDirecciónGeneraldeAsuntos depersonalAcadémico throughprojectpApIIt-In221707,“Factoresquedeterminanladistribucióndelosanfibiosenlaspozasaso-ciadasalRíosalado,puebla,méxico”;andbytheFacultaddeestudiossuperioresIztacalathroughtheprogramadeApoyoalosprofesoresdeCarrera(pApCA)2007–2008fortheproject“CaracterizacióndelaspozasasociadasalRíosalado(puebla)ysuinfluenciaenladistribucióndelosanfibios:aspectosecológicosygeográficos.”theresearchwasapprovedbytheDenisonUniversityInstitutionalAnimalCareandUseCommittee(protocol07-004).

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

University of Kansas Publications, Museum of Natural History15:1–148.

oliver-lopez,l.,A.Ramirez-bautista,andJ.A.lemos-espinal.2000.Bufo occiden-talis.Fecundity.Herpetological Review31:39–40.

InChina,thefirstleopardGeckowasreportedfromhainanIslandin1908as G. “lichtenfelderi”(ZhaoandAdler1993).thisinsularformwaspreviouslythoughttobeaVietnamesespeciesbutisnowseparatedasG. hainanensis(Grismeretal.2002,blairetal.2009).Goniurosauruslich-tenfelderiisconfinedtograniticsubstratesinnorthwesternVietnamand

leopardGeckosofthegenusGoniurosaurus(squamata:eublepharidae)typically inhabitcavesandcliffs in forestedareas in theRyu-Kyu

ArchipelagoofJapanandsoutheasternAsia(blairetal.2009).tentotwelvespecieshavebeendescribed,mostrecentlyG.catbaensis(Ziegleretal.2008)andG.huuliensis(orlovetal.2008),bothfromnorthernVietnam.

Discovery of Goniurosaurus Geckos (squamata: eublepharidae)

in northwestern Guangdong, chinaZu-shengYi1,Zhen-Changli2,Wei-liangWen3,mianhou4,Wen-hualu5,andJameslazell5

1Collegeoflifesciences,GuangzhouUniversity,panyu,Guangdong510006,China2Collegeoflifesciences,southChinanormalUniversity,Guangzhou,Guangdong510063,China

3ForestrybureauofQingyuanprefecture,Guangdong511500,China4CenterforChineseendemicherp-breedingandConservationResearch,shenyangnormalUniversity,liaoning110034,China

5theConservationAgency,Jamestown,RhodeIsland02835,UsA

photographsbymianhou,exceptwhereindicatedotherwise.

table 1.ComparisonofGoniurosaurusindet.withspeciesoftheG.luiigroup(G.araneus,G.bawanglingensis,G.catbaensis,G. huuliensis,andG.luii)onscalecountsandmorphometriccharactersinmm(fromourdata,Grismeretal.1999,orlovetal.2008).sl=supralabials;Il=infralabials;pm=post-mentals;po=preorbitals;eF=eyelidfringescales;pVt=paravertebraltuberclesbetweenlimbinsertions;bs=scalesaroundmidbody;Gst=granularscalessurroundingdorsaltubercles;lt4=basalanddistalsubdigitallamellaeunderfourthtoe(hindlimb);pp=precloacalpores,sVl=snoutventlength;hl=headlengthfromsnouttoposteriorofexternalear;hW=headwidthatthewidestareaofhead.

SL IL PM PO EF PVT BS GST LT4 PP SVL HL HWG.indet.(male) 8 7 2 16 50–51 34 106 9–10 17–21 12 80 23 14G.indet.(female) 8 7–8 4 14–16 51–55 33 107 10-11 17–21 — 86 23 15G. araneus 8–11 8–10 4–6 13–18 61–69 32–38 129–149 10–14 23–24 18–23 111–130 30–33 19–25G.bawanglingensis 8–10 7–11 2–3 12–18 56–67 32–36 104–133 9–13 18–22 37–46 104 — —G.catbaensis 8–9 6–8 2–3 — 52–56 33–34 112–127 8–11 22–24 16–21 84–113 23–32 16–23G. huuliensis 10–11 9–11 2–3 — 41–44 34–36 118–129 12–13 18–20 25–28 108–117 29–33 18–23G.luii 9–12 8–11 2–6 14–17 52–63 33–35 119–148 9–14 20–24 21–29 106–120 30–33 19–23

48 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 YIetAl.

offshoreislandsintheGulfoftongkin(orlovetal.2008).today,threespeciesareknowninChina:G. hainanensisandG. bawanglingensisarecon-finedtohainanIsland,whereasthepresenceofG.luiionhainanIsland(Grismeretal.1999)isdiscredited(l.l.Grismer,pers.comm.,2march2010);itdoesoccurinGuangxiprovince(Grismeretal.1999)andnorth-ernVietnam(Vuetal.2006).AnotherGoniurosaurushasbeenfoundinGuangdongprovince,andhereinwereportthisnewrecord.thispopula-tionofGoniurosaurus(currentlyidentifiedas“G.indet.”)probablyrepre-sentsanewspeciesintheG.luiigroup,whichalsoincludesG.araneus,G.bawanglingensis,G.catbaensis,and G.huuliensis(table1),allofwhichhavefivepairsofblacktransversedorsalbandsbeginningjustbehindthehead(the“nuchalloop”ofGrismer1988)andextendingtothebaseofthetail. thefirstindividualofGoniurosaurusindet.,collectedon16July2007inakarstcaveapproximately160–170kmnWGuangzhou,Guangdongprovince,atapproximately250minelevation,wasanadultmale,80mmsVl(southChinanormalUniversity[sCnU]26115).Detailsofthelocality,includingGpscoordinates,arewithheldbecauseofthedirecon-sequencesofover-collecting—whichhasledtotheextirpationofentirepopulationsofthisgenus(e.g.,stuartetal.2006). AfterseveralfailedattemptstocollectmoreanimalsinJune2008,wereturnedtothesiteon9February2010andsecuredtwoadditionalspeci-mens:anadultfemale,86mmsVl(sCnU26116)andajuvenile,48mmsVl(sCnU26117).Allthreespecimenshaveregeneratedtails.thecavesysteminwhichthesegeckoswerefoundissmall,withtwochambersandthreeopenings.thelowestopeningislevelwiththevalleyfloorandabout10mfromtheedgeofacultivatedpaddy;onecannotenterthecavesystemthroughthisopening.eachofthetwoupperopeningshasachamberthatis

connectedwiththeother(wecouldheareachother);theyare~30mapartinupslopedistance.themiddleentranceis~5mverticallyabovethevalleyfloorandtheupperentranceis~15mabovethelowestopening.Asmallstreamcoursesthroughthelowerreachesofbothcavechambersandflowsoutthelowestopening.stalactitesandstalagmitesaredevelopedinbothchambers,whicharewellusedbyhumans.Wesawpotsandpans,feathersofchickensandotherbirds,ashes,andcharcoal.Weobservedcavecricketsandspiders.theslopeiscutoveranddominatedbyshrubvegetation,withonlyafewscattered,maturetreesabout50cmintrunkdiameter.theslopeandcaveentrancesfacewest.sCnU26115wasfoundonthecavewalldur-

theotherthreespeciesofGoniurosaurusthathavebeenrecordedfromChina:G. hainanensis(top)andG. bawanglingensis(middle)areconfinedtohainanIsland,whereasG.luii(bottom)occursinGuangxiprovinceandinnorthernVietnam.

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Adult female (top) and immature of Goniurosaurus indet. from Guangdongprovince,China.


ingsummeratnight;sCnU26116andsCnU26117werecaughtamongrubbleonthecavefloorinwinterduringdaytime(wereturnedlateratnightbutfailedtofindmoregeckos).Althoughthesamplesizeisverysmall,thesedatamightbesuggestiveofseasonalbehavior. the two adults are among the dullest in color of all knownGoniurosaurus.Inlife,ourfemale(sCnU26116)hadagray-browndorsalgroundcolorwithdullyellowtints;thefivetransversedorsalbandsaresootytonearblack,bold,andinpairs;eachpairhasapalegray-brownbandwithadullyellowtintatcenter;theselightcenterbandsareimmaculate.thedorsalzonesbetweenthebandsandthetopoftheheadwerespottedandmarbledwithnear-black.theventerwaspale,immaculate,lavendertogray-brown.theiriswasbrick-red.themale(sCnU26115)isadrabversionofthefemalewhenpreservedandwassimilartoherinlife;itsstom-achcontainedfragmentsofacavecricket’sfemurandtibia.sexualsizedimorphisminGoniurosaurustypicallyisfemale-biased(femalesarelarger),probablybecausemalesdonotengageincombatbehavior,whichwouldfavorlargemalesize(KratochvílandFrynta2002). ImmatureGoniurosaurustypicallyhavebrightercolorationthanadults(Grismeretal.1994,1999).ourjuvenile(sCnU26117)wasstronglycontrastingredandyellow,and,exceptforitssmallersize,resemblesthemostcolorfuladultsofG.bawanglingensisandG. luii,asfiguredbyblairetal.(2009)andasdescribedbyGrismeretal.(1999,2002)andVuetal.(2006).thedorsalgroundcolorwaslightred;theventerwaspaleyel-low;thetransversedorsalbandswerebrightyellowedgedbythenear-blackpaireddorsalbands.theiriswasbrightred.Darkspottingontheheadandinthedorsalgroundcolorwassparse,indicatingthatincreasedspottingisafunctionofage.thetwoadultsdiffermostnotablyfrommembersoftheG. luiigroupindullercolorationandpattern,smallersize,andfewerprecloacalpores(table1).becausethetaxonomicstatusofGoniurosaurusindet.mightrequireageneticassessment,ourspecimenshavebeenpre-servedinethanoltofacilitateDnAextractionforsuchcomparisons.

Literature Citedblair,C.,n.l.orlov,h.-t.shi,andR.W.murphy.2009.Ataxonomicre-evalu-

ationofGoniurosaurus hainanensis (squamata:eublepharidae)fromhainanIsland,China.Russian Journal of Herpetology16:35–40.

Grismer, l.l. 1988. phylogeny, taxonomy, classification, and biogeographyof eublepharid geckos, pp. 369–469. In: R. estes and J. pregill (eds.),Phylogenetic Relationships of the Lizard Families.stanfordUniversitypress,stanford,California.

Grismer,l.l.,h.ota,ands.tanaka.1994.phylogeny,classification,andbio-geographyof Goniurosaurus kuroiwae(squamata:eublepharidae)fromtheRyu-KyuArchipelago,Japan,withdescriptionofanewsubspecies.Zoological Science11:319–335.

Grismer,l.l.,b.e.Viets,andl.J.boyle.1999.twonewcontinentalspeciesofGoniurosaurus(squamata:eublepharidae)withaphylogenyandevolutionaryclassificationofthegenus.Journal of Herpetology33:382–393.

Grismer,l.l.,h.shi,n.l.orlov,andn.b.Ananjeva.2002.AnewspeciesofGoniurosaurus (squamata: eublepharidae) from hainan Island, China.Journal of Herpetology36:217–224.

Kratochvíl,l.s.andD.Frynta.2002.bodysize,malecombatandtheevolutionofsexualdimorphismineublepharidgeckos(squamata:eublepharidae).Biological Journal of the Linnean Society76:303–314.

orlov,n.l.,s.A.Ryabov,n.t.tao,n.Q.truong,andh.t.Cuc.2008.AnewspeciesofGoniurosaurus(sauria:Gekkota:eublepharidae)fromnorthVietnam.Russian Journal of Herpetology15:229–244.

stuart,b.l.,A.G.J.Rhodin,l.l.Grismer,andt.hansell.2006.scientificdescrip-tioncanimperilspecies.Science312:1137.

Vu,n.t.,Q.t.nguyen,l.l.Grismer,andt.Ziegler.2006.FirstrecordoftheChineseleopardGecko,Goniurosaurusluii(Reptilia:eublepharidae)fromVietnam.Current Herpetology 25:93–95.

Ziegler,t.,Q.t.nguyen,A.schmitz,R.stenke,andh.Rösler.2008.AnewspeciesofGoniurosaurusfromCatbaIsland,haiphong,northernVietnam(squamata:eublepharidae).Zootaxa1771:16–30.

Zhao,e.andK.Adler.1993.Herpetology of China.Contributionstoherpetology10.societyforthestudyofAmphibiansandReptiles,Ithaca,newYork,UsA.

SNAKES uSING StuMPHOLES

theforestfloor.After~30secofobservation,thesnakewasdisturbedandmadeitswaytoastumpapproximately70cmfromitsoriginallocation.thesnakeretreatedintoatunnelassociatedwiththestump.bothobser-vationsoccurredinokaloosaCounty,Florida.toourknowledge,theyrepresentthefirstaccountsofthesespeciesusingthistypeofrefuge.on8April2005inUpsonCounty,Georgia,weobservedalargeadulteasternKingsnake(Lampropeltis getula getula)coiledandpartiallyvisibleatabout1400hwithintheleaflitterofalongleafpinestump. on 16 June 2009 at 0730 h, we observed an adult easternDiamondbackRattlesnake(Crotalus adamanteus)withinacavityundertherootsystemofalargedownedsandliveoaktree.on23July2009at1950h,weobservedanadultCottonmouth(Agkistrodon piscivorus)withinacavityassociatedwiththebaseofadownedtreeonthebankofasmallclear-waterstream.bothobservationsoccurredinokaloosaCounty,Florida.

stumpholesareacommonhabitatfeatureoffire-maintainedlongleafpineecosystemsinthesoutheasternUnitedstates.thesestumpholesandasso-

ciatedsubterraneantunnelsthatformasstumpsdecayorareconsumedbyfirehavebeenidentifiedasimportantrefugiafornumerousvertebratespecies(means2005;steenetal.,inpress).treesdownedbywind,suchasthoseseenaftermajorstormevents(Greshametal.1991),maycreatesubterraneandepressionsorcavitiesneartheirbasesasrootsareexposed.theserepresentpotentialmicrohabitatsforanimalsthatdonotconstructtheirownburrows.hereinwereportonfiveobservationsofsnakesusingrefugiaassociatedwithstumpholetunnelsorrootcavitiesofdownedtreesinlongleafpineforests. on26may2009at1300h,weobservedanadultpigmyRattlesnake(Sistrurus miliarius)alongsideaburntstump;therattlesnakeretreatedintoatunnelwithinthestumpholeshortlyafterbeingobserved.on13July2009at0640h,weobservedanadultCoralsnake(Micrurus fulvius)on

snakes using stumpholes and Windfall Tree-associated subterranean structures

in longleaf pine forestsDavidA.steen1,2,AmandaD.steen1,scottpokswinski1,seanp.Graham2,andloral.smith1

1JosephW.JonesecologicalResearchCenter,newton,Georgia39870,UsA([email protected])2Departmentofbiologicalsciences,AuburnUniversity,Auburn,Alabama36849,UsA

50 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 steenetAl.

theseobservationsindicatethatmanyspeciesofsoutheasternsnakes maybeinfluencedbyforestmanagementstrategiesthataffectthepres-enceof stumpholes and cavities associatedwithwindfall treeswithinlongleafpineforests.theserefugiamaybeespeciallyimportantinareasoflowGophertortoise(Gopherus polyphemus)burrowdensities,suchasinokaloosaCounty,Florida,wherefouroftheseobservationsoccurred,astortoiseburrowsaredocumentedsheltersforallaforementionedsnake

species(Jacksonandmilstrey1989).Forestmanagementpracticesinclud-ingtheharvestingofstumpsandfiresuppressionhavereducedstumpholehabitatinmanysoutheasternforests(means2006).Inaddition,harvestingtreesdownedbywindmayreduceavailablesubterraneanshelterforforest-associatedwildlifespecies,althoughlarge-scaleexperimentalmanipulationsofcoarsewoodydebrisinthesoutheasternCoastalplaindidnotdocu-mentcompellingtrendsthatsuggestthatamphibiansandreptilesgenerallyrespondedonapopulationlevel(owensetal.2008).

Literature CitedGresham,C.A.,t.m.Williams,andD.J.lipscomb.1991.hurricanehugowind

damagetosoutheasternU.s.coastalforesttreespecies.Biotropica23:420–426.

Jackson,D.R.ande.G.milstrey.1989.thefaunaofGophertortoiseburrows,pp.86–98.In:J.e.Diemer,D.R.Jackson,J.l.landers,J.n.layne,andD.A.Wood(eds.),Gopher Tortoise Relocation Symposium Proceedings.FloridaGameandFreshWaterFishCommissionnongameWildlifeprogramtechnicalReportno.5,tallahassee.

means,D.b.2005.thevalueofdeadtreebasesandstumpholesashabitatforwildlife,pp.74–78.In:W.e.meshakaandK.babbitt(eds.),Status and Conservation of Florida Amphibians and Reptiles.UniversitypressofFlorida,Gainesville.

means,D.b.2006.Vertebratefaunaldiversityinlongleafpinesavannas,pp.155–213.In:s.Jose,e.Jokela,andD.miller,(eds.),Longleaf Pine Ecosystems: Ecology, Management, and Restoration.springer,newYork.

owens,A.K.,K.R.moseley,t.s.mcCay,s.b.Castleberry,J.C.Kilgo,andW.m.Ford.2008.Amphibianandreptilecommunityresponsetocoarsewoodydebrismanipulationsinuplandloblollypine(Pinus taeda)forests.Forest Ecology and Management256:2078–2083.

steen,D.A.,J.m.linehan,andl.l.smith.multiscalehabitatselectionandrefugeuseofCommonKingsnakes,Lampropeltis getula,insouthwesternGeorgia.Copeia:Inpress.

thesunriseilluminatestheunderstoryofalongleafpineforestinokaloosaCounty,Florida.Althoughherpetologiststypicallyarenotactiveatdawn,aviansamplingobliga-tionsinthespringof2009enabledmetotakeadvantageofthemorninglight—andtofindtheCoralsnakeandeasternDiamondbackRattlesnakedescribedinthetext.

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theeasternDiamondbackRattlesnake(Crotalus adamanteus)usessubterraneanrefugesassociatedwithdownedtrees.however,onemayhavetofruitlesslyexaminescoresofsuchstructuresbeforeasnakeisencountered.theobservationofthisindi-vidualisdescribedinthetext.

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bythetimeDr.Fitch’sstudyoftheRingnecksnake(Diadophis punctatus)waspublishedin1975,hehadalreadyestablishedhimselfasthe“fatherofsnakeecol-ogy”(Fitch,h.s.1975.AdemographicstudyoftheRing-necksnake(Diadophis punctatus)inKansas.University of Kansas Museum of Natural History Miscellaneous Publication(62):1–53).thiswasthesinglemostfrequentlyencounteredspeciesofsnakeinhis50-yearstudyofreptilesontheUniversityofKansas(nowFitch)naturalhistoryReservation(Fitch,h.s.1999.A Kansas Snake Community: Composition and Changes Over 50 Years.KriegerpublishingCo.,malabar,Florida).

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manyofthesametechniquesusedtostudysnakesappliedaswelltoanotherreptilianspeciesnativetonortheasternKansas.theslenderGlasslizard(Ophisaurus attenuatus),althoughverysnake-likeinmanyways,isquitelizard-likeinitsinsectivorousdiet(Fitch,h.s.1989.AfieldstudyoftheslenderGlasslizard,Ophisaurus attenuatus,innortheasternKansas.Occasional Papers of the Museum of Natural History, The University of Kansas(125):1–50).

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52 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 ReeDetAl.

I n v a s I v e s p e c I e s

Is the northern african python (Python sebae) established in southern florida?

Robertn.Reed1*,Kennethl.Krysko2,RayW.snow3,andGordonh.Rodda1

1U.s.Geologicalsurvey,FortCollinsscienceCenter,2150CentreAve,bldgC,FortCollins,Colorado805262Floridamuseumofnaturalhistory,Divisionofherpetology,UniversityofFlorida,Gainesville,Florida32611

3nationalparkservice,evergladesnationalpark,40001stateRoad9336,homestead,Florida33034*Correspondingauthor:[email protected],970-226-9464

morethan45speciesofnonindigenousamphibiansandreptilesarecurrentlyestablishedinFlorida,largelyasaconsequenceofinterna-

tionaltradeinliveanimalsaspets(meshakaetal.2004,Kraus2009,K.Krysko,unpubl.data).Amongthese,theburmesepython(Python molu-rusbivittatusKuhl1820)hasexpandeditsrangethroughseveralthousandsquarekilometersofthesouthernevergladesandadjacentareas(snowetal.2007).theboaConstrictor(Boa constrictorlinnaeus1758)alsoisestablishedinaforestedcountyparkinsoutheasternmiami,miami-DadeCounty(snowetal.2007),butdelineatingthegeographicdistributionofthispopulationisdifficultbecauseboasfoundnearbycouldbeeitherdispersersfromtheknownpopulationorrecentlyreleasedorescapedani-malsfromcaptivity.Free-rangingindividualsofseveraladditionalspeciesoflargeconstrictors(e.g.,GreenAnaconda,Eunectes murinuslinnaeus1758;YellowAnaconda,E. notaeusCope1862;Reticulatedpython,Broghammerus reticulatus schneider1801; andWhite-lippedpython,Leiopython albertisiipetersandDoria1878)havebeenfoundinvariouspartsofFlorida,butevidenceofreproductionforthesespeciesispresentlylacking.herein,weprovideevidencesuggestingthepossibilityofarepro-ducingpopulationofathirdspeciesofgiantconstrictorinFlorida,thenorthernAfricanpython(Python sebaeGmelin1788;Fig.1). In2002,alarge(ca.4.9mtotallength)Python sebaewasfoundattheintersectionofsW26thstreetandsW147thAvenue,miami,miami-DadeCounty(25.74298on,-80.43221oW;datumWGs84;Fig.2A),andrecovered,butnotretained,bythemiami-DadeVenomResponseUnit(A.Cruz,pers.comm.;Internetbroadcastingsystems2005).Asnakeofthissizewouldalmostcertainlybefemale,asthisspeciesexhibitsfemale-biasedsexualsizedimorphism,andmalesarenotknowntoattainsuchsizesinitsnativerange(ReedandRodda2009).

on11october2005,anadult(ca.3mtotallength)Python sebae(photographicvoucherUF153699)wasfoundinaplantnurseryjustnorthofthesameintersection(Fig.2b).thesnakehadenteredanoutdoorenclo-sureforraisingfowl,andhadconsumedadomesticturkey(A.Cruz,pers.comm.;Internetbroadcastingsystems2005). Innovember2008,anexperiencedsnakecollectorobservedalarge(3.5–4.0mtotallength)adultpythonalongacanalbank(25.72739on,-80.46548oW),3.7kmsouthwestoftheintersectioncitedabove(Fig.2C;A.Flanagan,pers.comm.2009).heattemptedtocapturethesnake,butitescapedfromhisgraspafterabriefstruggle.thisobserverhascaptured100P. molurusinsouthernFlorida,andwasconfidentofhisidentificationofthissnakeasP. sebae,includingtheobservationthattheindividualboreanaberrantmiddorsaldarkstripe.

Fig. 1. northernAfricanpython(Python sebae)capturedbyAnthonyFlanaganon19December2009.

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Fig. 2.specimensandobservationsofthenorthernAfricanpython(Python sebae)inmiami,miami-DadeCounty,Florida.localities(seetext)include:A=2002,b=11october2005,C=november2008,D=30may2009,e=5August2009,F=16August2009,G=19December2009.


on30may2009,anapproximately2.75m-longPython sebaewasstruckbyamotorvehicleatthesameintersectioncitedabove(Fig.2D),andrecoveredbythemiami-DadeVenomResponseUnit(A.Cruz,pers.comm.).thissnakesubsequentlydiedfromitsinjuries,anddissectionrevealedthatitwasanadultfemalewith37“undevelopedova”(unknownwhetherthesewereovarianfolliclesoroviductaleggs,astheywerediscardedpriortopreservation).thisspecimen,alongwithphotographicimages,wastransferredtotheFloridamuseumofnaturalhistory(UF155725)forpreservationanddocumentation. on5August2009,aneonate(59cmsVl)Python sebae(UF155500)wasfounddeadonUs41,0.09kmwestof160thAvenue(25.76089on,-80.45596oW;Fig.2e),whichis3.0kmnorthwestoftheintersectioncitedabove.on16August2009,anotherjuvenileP. sebae(110g,UF155726)wascollectedat15527sW18thstreet,miami(25.74984on,-80.44637oW; Fig. 2F), and recovered by the miami-Dade VenomResponseUnit(l.Woods,pers.comm.).Dissectionrevealeda72-gboat-tailedGrackle(Quiscalus major)inthestomachofthepython.seetable1forbodylengthsanddispositionofspecimens. mostrecently,on19December2009,anadultmaleP. sebae(UF157193,249cmsVl,281cmtotallength,10.374kg)wascollectedfromaMelaleucaslashpile(Fig.3)justwestof157thAvenuesouthofUs41(25.75245˚n,-80.45079W;specimennotyetaccessionedintoFlmnh).thesnakewasbaskingontopofdeadlogsandappearedreproductivelycompetent;itstesteswereswollenandmildlyturgid,spermductswerecon-voluted,andmicroscopicexaminationoffluidexpressedfromtheductsrevealedmultiplespermatozoa. theabovespecimensandobservationsarepresentlyconfinedtoasmall(~10km2)areaofmiami,just1.6kmeastofevergladesnationalpark(Fig.2).AllsevenpythonswerefoundsoutheastoftheintersectionofUs41andsR997,anareaoftenreferredtoasbirdDrivebasin.takenasawhole,thesesevenobservationsoveraseven-yearperiod,includingmultipleadults,agravidfemale,andyoung-of-yearhatchlingssuggestthepossibilityofareproducingpopulationofPython sebae.Recoveryofadultsoverseveralyearsfollowedbyrecentlyhatchedjuvenilesarguesagainstthenotionthatalloftheseindividualsweretheresultofasinglereleaseofmul-tipleindividuals,leavingmultiplereleasesorareproducingpopulationasthemostlikelyexplanations.theseobservationsdonotrepresentdefinitiveevidenceofareproducingpopulation,andsomewouldarguethatonlythediscoveryofafemalebroodingeggswouldqualifyasdefinitiveevidence.bythetimesuchdiscoveriesaretypicallymade,ofcourse,aspeciesisoftenwellestablished.Indeed,theobservationsofPython sebaeinmiamihighlightthedifficultyofdeclaringwhenapopulationofextremelycrypticreptileshasbecomeestablished. threeofthefiveadultpythonswerefromasmallareajustsouthoftreeIslandpark,whichissouthofUs41andboundedontheeastandwestbysW146thstreetandsW149thstreet,respectively.thetwohatchlingswerefoundfarthertothewestandnorth,andcouldconceivablyrepresentdispersingindividuals.habitatsinthisareaincludehigh-densitysingle-fam-ilyhousingdevelopments,undevelopedbuthighlydisturbedhabitatswitha

preponderanceofinvasiveMelaleucatrees,smallman-madecanalsandlakes,agriculturalareas(primarilytothesouthwest),andseasonallyfloodedwet-lands.landownershipintheareaiscomplex,withvariousparcelsbelongingtohomeowners,housingdevelopers,plantnurseryandagriculturalinter-ests,miami-DadeCounty,southFloridaWatermanagementDistrict,miccosukeetribeofIndians,stateofFlorida,andothers. theareadescribedaboveiswithinoradjacenttotheeasternedgeoftheknowndistributionofinvasiveburmesepythons(Python molurus bivittatus;snowetal.2007).DistinguishingbetweenP. molurusandP. sebaecanbechal-lenging,andoftenhingesoncolorpattern;althoughmeristiccharacters(e.g.,scalecounts)aresufficienttodistinguishmanyindividuals,moderateoverlapexistsamongthesespeciesinmanyofthosecharacters.Python molurus andP. sebae areknowntohybridizeincaptivity(branchanderasmus1984andrefer-encestherein),whichcouldfurthercomplicatetheidentificationofapythonrecoveredfromthisarea.Fertilityandfitnessofhybridsisunknown,asarethepotentialimplicationsofaddingP. sebaegenestotheexistingpopulationofP. molurus.thepotentialdifficultyofaccuratelyidentifyingfree-rangingindividualsofdifferentspeciesofexoticpythonsisexemplifiedbytwoP. sebaerecoveredfromsouthwesternFlorida(eastofsarasota)in2006and2009.bothofthesespecimenswereoriginallyreportedasadifferentspecies(oneidentifiedasP. molurusandoneasaReticulatedpython,Broghammerus[for-merlyPython] reticulatus).AsthepopulationofburmesepythonscontinuestoexpandwithinFlorida,accurateidentificationoflargesnakeswillbecrucialtoidentifyingincipientpopulationsofotherspeciesbeforetheirpopulationsbecometoowidespreadforeffectiveeradicationprograms. effortsarecurrentlyunderwaytodevelopeducationandoutreachmaterialsthatwillallowdiscriminationamongtheseandothergiantcon-strictorspeciesbycitizensandresourcemanagerslackingherpetological

InVAsIVespeCIes

Fig. 3.scottGoetz(U.s.Geologicalsurvey)precariouslymakeshiswaythroughalargepileofdeadMelaleucalogswhilesearchingforpythonsinbirdDrivebasin.thepythonpicturedinFig.1hadbeencapturedfromthispile.

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table 1.specimeninformationforsevenP. sebaerecoveredfromthewesternboundariesofgreatermiami,Flfrom2002to2009.Whenavailable,speci-menswereaccessionedintotheFloridamuseumofnaturalhistoryinGainesville,Fl.

Date Specimen No. Specimen Type Approx. Total Length

19December2009 157193 Wholebody 2.81m16August2009 155726 Wholebody 0.71m05August2009 155500 Wholebody 0.67m30may2009 155725 Wholebody 2.75mnovember2008 n/A Crediblesighting ~3.7m11october2005 153699 mediaaccount 3.05m2002 n/A Crediblesighting 4.9m


expertise.stakeholdersfromvariousfederal,state,tribal,andlocalagen-ciesalsoplantopartnerwithnon-governmentalcooperatorstoconductintensivesurveysforPython sebaeinthebirdDrivebasinareain2010(D.Giardina,FloridaFishandWildlifeConservationCommission,pers.comm.2009).suchsurveyswillaimtodelineatethesizeandgeographicextentoftheincipientpopulationandattempteradicationofremainingindividuals.

AcknowledgmentsWethankA.Cruz,m.easter,A.Flanagan,C.Gillette,andl.Woodsforprovidinginformationand/orspecimens,m.Granatoskyforassistingwithspecimenpreparation,andC.Doveforidentificationofavianpreyremains.K.hart,W.meshaka,m.Rochford,andananonymousreviewerprovidedhelpfulcommentsonearlierversionsofthismanuscript.

Literature Citedbranch,W.R.andh.erasmus.1984.Captivebreedingofpythons insouth

Africa,includingdetailsofaninterspecifichybrid(Python sebae natalensis

xPython molurus bivittatus).Journal of the Herpetological Association of Africa 1984:1–10.

Internetbroadcastingsystems.2005.Rare10-footpythonattacksfarmer’spoultry.<http://www.clickorlando.com/news/5087908/detail.html> (accessed31August2009).

Kraus,F.2009.Alien Reptiles and Amphibians: A Scientific Compendium and Analysis.Invadingnature—springerseriesinInvasionecology,volume4.springer,newYork.

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

Reed,R.n.andG.h.Rodda.2009.Giantconstrictors:biologicalandmanage-mentprofilesandanestablishmentriskassessmentforninelargespeciesofpythons,anacondas,andtheboaConstrictor.U.s.GeologicalsurveyopenFileReport2009-1202.FortCollins,Colorado.

snow,R.W.,K.l.Krysko,K.m.enge,l.oberhofer,A.Warren-bradley,andl.Wilkins.2007.IntroducedpopulationsofBoa constrictor(boidae)andPython molurus bivittatus(pythonidae)insouthernFlorida,pp.416–438.In:R.W.hendersonandR.powell(eds.),Biology of the Boas and Pythons.eaglemountainpublishinglC,eaglemountain,Utah.

ReeDetAl.

IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 55ConseRVAtIonReseARChRepoRts

Conserving Mexican Amphibianstraditionally, biodiversity conservation gapanalyseshavebeen focusedongovernmentalprotected areas (pAs). however, an increas-ingnumberofsocialinitiativesinconservation(sICs) are promoting a new perspective foranalysis.sICsincludealloftheeffortsthatsoci-etyimplementstoconservebiodiversity,suchaslandprotection,fromprivatereservestocom-munityzoningplans,someofwhichhavegener-atedcommunity-protectedareas.thisisthefirstattempttoanalyzethestatusofconservationinlatinAmericainwhichsomeofthesesocialini-tiativesareincluded.theanalyseswerefocusedonamphibiansbecausetheyareoneofthemostthreatenedgroupsworldwide.OchOa-OchOa et al.(2009.PLoS ONE4:1–9)usedanichemodelapproachtomapthepotentialandrealgeographicaldistribution(extractingthetrans-formedareas)ofendemicmexicanamphibians.Allspecieshavesufferedsomedegreeof loss,but36specieshavelostmorethan50%oftheirpotentialdistribution.For50micro-endemicspecies,theauthorscouldnotmodeltheirpoten-tialdistributionrangeduetothesmallnumberof recordsper species; therefore, theanalyseswereperformedusingtheserecordsdirectly.theauthorsthenevaluatedtheefficiencyoftheexist-ingsetofgovernmentalpAsandestablishedthecontributionofsICs(privateandcommunity)forlandprotectionforamphibianconservation.theyfoundthatmostofthespecieshavesomeportionoftheirpotentialecologicalnichedistri-butionprotected,but20%arenotprotectedatallwithingovernmentalpAs.73%ofendemicand 26% of micro-endemic amphibians arerepresentedwithinsICs.however,30micro-

endemicspeciesarenotrepresentedineithergovernmentalpAsorsICs.thisstudyshowshowtheroleoflandconservationthroughsICsisbecominganincreasinglycrucialelementforanimportantnumberofspeciesnotprotectedbygovernmentalpAs.

Artificial Refuges Facilitate Reptilian Recovery

ecosystem restoration requires that habitatrequirementsof all speciesbe considered. In1998,amassivequantityoftailingsbrokeoutoftheholdingpondoftheAznalcollarmineinsouthwesternspainandpollutedtheGuadiamarRivervalley.Aftertheaccident,asoilandveg-etation restoration program began, and theGuadiamarGreenCorridorwascreatedtocon-necttwolargenaturalareas,anationalparkandthesierramorena.themine-tailingspillpol-lutedalargearea,givingrisetoaninterestingcasestudyinvolvingsoilandvegetationrestorationandrecoveryoftheterrestrialreptiliancommu-

nityinthecontaminatedarea.the5-yearstudyby Marquez-FerrandO et al. (Restoration Ecology17:660–667)evaluatedwhethertherep-tiliancommunityinthepollutedarearemaineddramaticallyimpoverishedbecauseoftheelimi-nationofallnaturalrefuges.theareamanagedwithartificialrefugesexhibitedabetterandfasterrecoveryofthereptiliancommunitythanthecontrolareawithnorefuges.

Power Boating and Northern Map turtles

Recreational power boating is growing inpopularityinnorthAmerica.thisactivityisknowntohavelethalandsub-lethaleffectsonaquaticwildlife,andfreshwaterturtlesmaybeparticularlysensitivetothisactivity.Bulté et al. (2009.Aquatic Conservation: Marine and Freshwater Ecosystems, early view publishedonline26october2009)reportedonpatternsof traumatic injuries inflicted by powerboatpropellerstoCommonmapturtles(Graptemys geographica)fromtwositesdifferinginboattraf-ficintensityinontario,Canada.therelativevulnerabilityofturtleswasassessedinlightofseasonalpatternsinboattraffic,asafunctionof sex- and age-specific movement patterns,habitatuse,andbaskingbehaviorobtainedbyradio-telemetry.theauthorsconductedpopu-lationviabilityanalyses(pVA)toevaluatethepotentialdemographicconsequencesofmortal-ityinducedbypowerboats.theprevalenceofpropellerinjurieswastwotoninetimeshigherinadultfemalesthaninadultmalesandjuve-nilefemales.patternsofmovement,habitatuse,andaquaticbaskingindicatedthatadultfemales

aremoreexposedtocollisionswithboats.pVAshowed thatboat-inducedmortality inadultfemalescouldleadtorapidpopulationextinc-tioniftheriskofmortalitywhenhitbyaboatisgreaterthan10%.theresultsofthisstudyshowedthatrecreationalpowerboatingisaseri-ousthreattonorthernmapturtles,evenunder

horseshoe Whip snakes (Hemorrhois hippocrepis)werefoundsignificantlymorefrequentlyinplotswithartificialrefugesthanincontrolplotswithoutrefuges.

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thecriticallyendangeredAxolotl(Ambystoma mexicanum)isoneofmanymexicanamphibianswithdecliningdistributions.

c o n s e r v a T I o n r e s e a r c H r e p o r T s

AdultfemaleCommonmapturtles(Graptemys geo-graphica)aremorevulnerabletocollisionswithboatsthanareadultmalesandjuveniles(illustrated).

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moderateboattraffic,whichspeakstotheneedtoadoptmeasuresrestrictingboattrafficinareasimportanttoturtles.

the trade in Vertebrates Promotes Extinctions and Introductions

the process of taxonomic homogenizationoccurs through twomechanisms, extinctionsandintroductions,andleadstoareductionofglobalbiodiversity.rOMagOsa et al.(2009.Conservation Biology23:1001–1007)usedavail-ableU.s.tradedataasaproxyforglobaltradeinlivevertebratestoassessthecontributionoftradetotheprocessoftaxonomichomogeniza-tion.DataincludedallavailableU.s.importa-tion and exportation records, estimation of

extinctionrisk,andreportsofestablishmentout-sidethenativerangeforspecieswithinsixver-tebrategroups.basedonmonteCarlosampling(aclassofcomputationalalgorithmsthatrelyonrepeatedrandomsamplingtocomputetheirresults), thenumberof species traded,estab-lishedoutsideofthenativerange,andthreat-enedwithextinctionwasnotrandomlydistrib-utedamongvertebratefamilies.twenty-eightpercentofvertebratefamiliesthatweretradedpreferentiallywerealsoestablishedorthreatened

withextinction,anunusuallyhighpercentagecomparedwiththe7%offamiliesthatwerenottradedpreferentiallybutthatbecameestablishedorthreatenedwithextinction.theimportanceoftradeinhomogenizationofvertebratessug-geststhatadditionaleffortsshouldbemadetopreventintroductionsandextinctionsthroughthismedium.

Road-kill HotspotsRoadswithwetlandsonbothsidesareconsid-eredhotspotsbecauseofthehighmortalityratesofamphibiansandreptiles that livenear theroads.langen et al.(2009.Journal of Wildlife Management 73:104–114)studieda219-milehighwaynetworkinnewYork,wheretheydis-

coveredbothamphibianandreptilianroadmor-talitytobespatiallyclustered,andcausewaysweremorelikelytohaveroad-killsthanpointswithoneadjacentwetlandorwithnowetlandpresent.Roadcrossingoccurswhentheanimalsmigrateduringseasonalmigrationstoandfromhibernationorbreedingsites.theauthorsstatedthatplannerscouldidentifyvalidpredictorsofhotspotswhendesigningorrestoringroadstoavoidasmuchharmaspossibletoamphibianandreptilianpopulations.

Australian Snakes Vulnerable to Climate Change

thebroad-headedsnake(Hoplocephalus bun-garoides),Australia’smost endangered snake,will findareasofhigherelevationmost suit-ableforsurvivingclimatechange.PenMan et al.(Diversity and Distributions2010:109–118)analyzedthesydneybasinbioregionandpre-dictedthedistributionofthisspeciesunderlowandhighclimatechangescenariosfor2030and2070.populationswillbelostunderbothcli-mate-warmingscenarios.thespecieshashighlyspecializedhabitat requirementsandexhibitsdelayed maturation and a low reproductiverate.thesefactorsreducethespecies’abilitytorecoverfrompopulationreductions.populationshavedeclineddramaticallythroughouttherangein the 200 years since european settlementbecauseofanthropogenicreasons,particularlyurbanization.theauthorsconcludedthatmanyareasthatarecurrentlyoccupiedwillbecometoohotanddryforthisspecies,andonlycoolerareasathigherelevationswillremainsuitable.

ConseRVAtIonReseARChRepoRts

northernleopardFrogs (Lithobates pipiens)wereamongninespeciesofanuranroadkillsfoundmorefrequentlyoncausewaysthanonroadswithoneadja-centwetlandornowetland.

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Ifthebroad-headedsnake(Hoplocephalus bungaroi-des),Australia’smostendangeredsnake,willsurviveclimatechange, itwillmost likelyberestrictedtocoolerareasathigherelevations.

speciesinthefamilyIguanidaeweretradedsignificantlymorethanexpectedbasedonrandomizationprocedures—despitethefactthatsomespecies,suchasthissanestebanIslandChuckwalla(Sauromalus varius),rarelyappearinthetradeforpets,skins,ormeat.

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turtles and Wetlandsseasonaldifferencesinwetlandusebyspottedturtles (Clemmys guttata) and blanding’sturtles(Emydoidea blandingii)arecomplexandrequireadynamic landscape to sustain theserareturtles.Beaundry et al.(2009.Journal of Herpetology43:636–645)examinedthesefactorsinsouthernmaine,wherewetlandsareabun-dantanddiverse.thestudyconcludedthatthecharacteristicofthewetlandsusedbytheturtles

variesbetweenthespecies.Inthespring,spottedturtlesusedwetlandswith abundantWoodFrogeggmassesandshowedanegativeassocia-tionwithforestedswamps.Inthesummerandfall,theyusedwetlandswithhighsunexposure.blanding’sturtlesusedwetlandswithindecidu-ousforestsandwithahighcoverofsphagnuminthespring.Duringlatesummerandfall,theseturtleswere associatedwithdeep-waterwet-lands.seasonally,blanding’sturtleswereactiveearlierthanspottedturtles.

Horned Lizards and Climatethepredictionthatvariationinspeciesmor-phologyisrelatedtoenvironmentalfeatureshaslongbeenofinteresttoecologistsandevolution-arybiologists.luxBacher and KnOuFt(2009.

Journal of Evolutionary Biology22:1669–1678)usedmorphological,environmental,andphylo-geneticdatacompiledfromstudiesofhornedlizards(Phrynosoma)toexaminemorphologi-calandclimaticvariationacrossthegeographicrangesofthesespeciesinanevolutionarycon-text.Inthefirststudy,patternsofenvironmentalnicheandmorphologicaltraitvariationamonghornedlizardswereexaminedinaphyloge-neticcontext.theresultsindicatedthatcloselyrelated species of Phrynosoma occupy moresimilar environmental niches than distantlyrelatedspecies,whichwassuggestiveofnicheconservatism.Inthesecondstudy,morphologi-calcharactersof14hornedlizardspeciesweremeasured.theresultssuggestednodiscernablepatternintheevolutionofbodysizeorformamonghornedlizards.thelackofaphyloge-neticsignalinbodysizemayseemsurprising,butphylogeneticsignalsareresponsivetosamplesizes,andthisanalysiswaslimitedto14species.thefinalstudylookedatassociationsbetweenenvironmentalnichesandmorphologicaltraits.Althoughthestudycouldnotdetermineifmor-phologicalcharacterswererespondingdirectly

toclimateorto,forexample,vegetativecom-munitiesthatareaffectedbyclimate,theresultsshowed that morphological traits (excludingbodysize)arecorrelatedwithclimaticfeaturesamonghornedlizards.speciesofPhrynosomawithlonglimbsoccurindrylowlandhabitatswithrelativelyhightemperaturesandspecieswithlargeheadsoccurinwetenvironmentswith

hightemperatureseasonality(i.e.,wideannualfluctuationsintemperature).thispatternisalsosupportedbyempiricaldata.

Bird and Crocodilian Lungs are Similar

When it breathes, the American Alligator(Alligator mississippiensis),asemi-aquaticecto-thermwithoutairsacs,possessesthesameuni-directionalairflowinthelungsasbirds.Duringinhalationandexhalation,airflowintheavianlungmovesgasesinthesamedirectionthroughsmalltubescalledtheparabronchi.Crocodilianlungsaredistinctlydifferentthanthoseofbirds,butthetopographyoftheintrapulmonarybron-chiissimilarinboth.FarMers and sanders(2010.Science327:338–340)testedthehypoth-esisthatairflowinalligatorlungsisunidirec-tionalandconcludedthatitwasextremelybird-like.themechanismsforunidirectionalairflowinalligatorlungsremainunknown,butcroco-dilians andbirdsbothbelong to the crown-groupArchosauria,whichincludescrocodiles,dinosaurs,andpterosaursofthetriassicperiod(251–199millionyearsago), suggesting thattheunidirectionalairflowinalligatorsandbirdscouldhaveevolvedbeforethesegroupsdiverged.

nAtURAlhIstoRYReseARChRepoRts

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

morphologicaltraits(excludingbodysize)arecorre-latedwithclimaticfeaturesamonghornedlizards.Forexample,short-limbedmountainshort-hornedlizards(Phrynosomahernandesi)occurathigheleva-tions and cooler temperatures across the westernUnited states and into sonora, Chihuahua, andDurango,mexico.

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UnidirectionalairflowthroughthelungsofAmericanAlligators(Alligator mississippiensis)isessentiallysimi-lartothatinbirds,suggestingthatairflowmecha-nismsevolvedbeforecrocodiliansandbirdsdiverged.

58 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 neWsbRIeFs

Amphibian Reserve Launched in Colombia

An exciting new partnership between theIUCn/ssC Amphibian specialist Group,IUCnnetherlands,Dendrobatidaenederland,Conservation International-Colombia, andFundaciónproAveshasledtothelaunchofanewamphibian reserve toprotect spectacularpoisonfrogsinColombia.Colombiahasmorethreatenedamphibiansthananyothercountry,largelyrestrictedtohighlyfragmentedsubtropi-calandmontaneforestswhichareunprotectedandatthreatofagriculturalexpansion.WithinColombia,thehotspotforthreatenedamphibiansistheCentralCordillera,whichalsohasthegreat-estconcentrationofcoffeeproductioninsouthAmericaandisalmostdenudedofnaturalforests.

After extensive searches of the CentralCordillera for amphibians,herpetologist andFundaciónproAvespresident,AlonsoQuevedo,withecologistoscarGallego,discoveredoneofthe largestsurvivingforest fragments,amere200acresofforestonitseasternflank.notonlydidAlonsodiscoverthatthe200acrescontainedmanythreatenedamphibians,butthatitheldmanypreviouslyundescribedspeciesincludingtwospectacularpoisonfrogs,recentlynamedas the swainson’s poison Frog (Ranitomeya doriswainsonae)andlittleGoldenpoisonFrog(Ranitomeya tolimense). sadly, these last 200 acres were in theprocessofbeingclearedforavocadoandcof-feeplantationsandthatwouldalmostcertainlysealthefateofcountlessamphibiansandotheruniquebiodiversity.“Inanurgentbidtosavethisuniqueislandofamphibiandiversity,”saidAlonso Quevedo, “I negotiated with differ-entlandownersofthe200-acreforesttostopclearingforestandsellthelandtothenationalconservationnGo,FundaciónproAves.theownersagreed,soIimmediatelyapproachedtheIUCn/ssCAmphibianspecialistGroupandIUCnnetherlandsforemergencysupport.” the twogroupspromptlyprovided thesupport necessary to purchase the land and

resources for the long-term protection andmanagementof thearea. InearlyDecember,thenewlynamed“RanitaDoradaAmphibianReserve,”namedafterthelittleGoldenpoisonFrog,waslaunched.thisactioncomesatacru-cialmomentwhenconservationmeasuresareurgentlyneededforthishighlysensitivegroup.

Robin Moore AmphibianspecialistGroup

Species, Magazine of the Species Survival Commission(IUCn)50:15

tracking the Illegal Capture of Marine turtles

Criticallyendangeredmarineturtlesfromsouth-easternAsiaarebeingtargetedbyforeignvesselsoriginatinginhainan,China,and(toalesserdegree)Vietnam.theseboats leaveportwiththeexpresspurposeofcatchingmarineturtles,apracticewhichisillegalbothintheirhomecoun-triesandinthewatersofcountriesinwhichtheyfish.thelasthalfdecadehasseensuchanote-worthyincreasethatthereisaneedtodeterminetheseverityofthispractice.howmuchpoachinggoesunrecordedorundetected?howseverearetheimpactstoturtlepopulations?Whatdrivesthistrade,andhowcanitbecurtailed? significant and urgent inroads need tobe made into curbing this illegal trade andfurtherresearchisneededtofullyunderstandthe market forces at play in illegal wildlifetrade inorder todesigneconomicdeterrentsto it. In June 2009 members of the IUCnmarineturtle specialistGroup coorganizedaworkshopalongwiththeterengganustateGovernment,Universitymalaysiaterengganu,themarineResearchFoundation,ConservationInternational (philippines), and the IUCnmarineturtlespecialistGroup.thepurposeof

theworkshopwastodocumenttheapprehen-sionsofforeignvesselsandfishermeninvolvedintheillegaldirectcapture,topresentinforma-tiononthedecliningtrendofmarineturtlesintheregionandtheeffortsundertakentoarrestthedecline,andtodemonstratethatdirectcap-tureofadultandsub-adultmarineturtleswillrapidlycauseacollapseofturtlepopulationsinthesoutheastAsianRegion. solutionstothedirectcaptureofseaturtleswerediscussed,andthereareplanstoaddresstheproblem through apartnershipwithChineseauthoritiesandscientists,analyzingmarketforces,raisingawarenesswithinthefishery,trainingofcustomsstaffinturtlemeatandpartsidentifica-tion,trackingconsignmentsthroughDnAevalu-ation,translationandprovisionofconservationmaterialsintomandarin,andcontinueddialogueandinformationexchangesessions.

Nicolas J. Pilcher, Chan Eng Heng, and Kevin Hiew marineturtlespecialistGroup

The State of the World’s Sea Turtles (sWot)reportisproducedbythemarineturtlespecialistGroup in partnership with ConservationInternationalandDukeUniversity.thefourthvolume,releasedin2009,featuresthefirst-evermapofglobalFlatbackturtle(Natator depressus) nestingdata,geneticstocks,andinwaterdistribu-tion.otherhighlightsincludearticlesaboutwhyleatherbackturtlepopulationsvaryglobally,howretailsaleshelpcommunitiesandseaturtlesinbrazil,andhowfishermenareaidingconser-vationeffortsinCanada.thefullreportcanbedownloadedatwww.seaturtlestatus.org.

Roderic Mast and Nicolas J. Pilcher Co-Chairs,marineturtlespecialistGroup

Species, Magazine of the Species Survival Commission(IUCn)50:24–25

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AnewamphibianreserveinColombiaishometomanythreatenedspecies,includingthepreviouslyundescribedlittleGoldenpoisonFrog(Ranitomeya tolimense).

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Criticallyendangeredmarineturtles,suchasthisAustralianFlatbackturtle(Natator depressus),arebeingtargetedbyforeignvesselsoriginatinginhainan,China,andVietnam.

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IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 59neWsbRIeFs

dog Hunts Invasive turtlesIf the turtles inblackburnlakedon’t smellright,Anguswillsniffthemout.thepure-bredlabradorRetrieverhasbeentrainedspecificallytotrackandhuntdowntheillegallyintroducedRed-earedslider(Trachemys scripta elegans).theinvasivepesthasbeenseeninblackburnlakesanctuary,elsternwickparklake,andRuffeylakeparkinWhitehorse,Australia. AturtlewascapturedinblackburnlakeinJanuaryofthisyear,butauthoritiesbelieveuptothreeturtlesandpossiblyanestremain.GaryJackson,Angus’trainer,saidthefive-year-oldwastheonlydogintheworldtrainedspecificallytocatchRed-earedsliders.“Wehavetrainedhimtoignorenativeturtlesandeggs,”mr.Jacksonsaid.“helovesfindingthesliderturtle’snests;whenhesniffsonehestartsdiggingforitandunlesswepullhimoff,hewoulddigallthewaythroughtothenestandprobablyeattheeggs.” Angus is on loan from theQueenslandGovernmentforthenextweekandwillbehunt-ingfortheelusiveturtleinblackburn. AgricultureministerJoehelpersaidtheWorld Conservation Union listed the Red-earedsliderasoneof theworld’s100most-invasivespecies.Femaleslayupto70eggsand,if unchecked, the species can destroy nativehabitats and food supply. Red-eared slidershaveadistinctiveredstripebehindeacheyeandadome-shapedshell.mr.helperurgedthepub-lictoreportanysightingstotheDepartmentofprimaryIndustries.

James Dowling Whitehorse(Australia)Leader

27october2009

African dwarf Frogs Implicated in Salmonella Outbreak

AmphibiansandreptilesarecarriersofSalmonella. thistypeofbacteriaisahealthhazardmostfre-quentlycontractedbytouchingrawfoods(e.g.,meats,eggs),butalsocanbeacquiredbytouch-inganimalsthatcarrySalmonella.thoroughlywashing hands after any potential contact isimportantifinfectionsaretobeavoided.AfricanDwarfFrogs(Hymenochirus boettgeri)havebeenimplicatedin85casesofpatientswithSalmonella

typhimurininfectionsin2009.thiswasthefirstreportedmultistateoutbreakofSalmonellainfec-tionsassociatedwithamphibians.theinvestiga-tionconductedbytheCenterforDiseaseControlandpreventionfoundtheinfectionin31states,extendingfrommassachusettstoCalifornia,andtheagerangeofpatientswas3weeksto54years.the multistate investigation revealed that allpatientswereexposedtofrogs,whichweretracedbacktothesamebreederinCalifornia.AfricanDwarfFrogsareaquaticanimalssoldasorna-mentalaquariumpets.themostlikelysourceoftransmissioninthisoutbreakwascontactwithwaterfromthefrogs’aquaria.

National Center for Zoonotic, Vector-borne, and Enteric Diseases(ZVeD)

7January2010

Florida Wildlife (Native and Non-native) Affected by Protracted Cold Spell

As January daytime temperatures hoveredinthe50sanddroppedbelow30atnightforseveraldaysinarow,Florida’scrops,aswellasnativeandnon-nativefloraandfaunafeltthechill.Wildlifeofficialsreportedatleast70deadcrocodiles,morethan60manateecarcasses,andperhapsthebiggestfishkillinmodernFloridahistory.Floridasea-turtlerehabilitatorsorganizedamassiverescueofmorethan2,000turtlessuf-feringcold-stunsyndrome.thecoldwaterpara-lyzesthem,makingthemunabletoswimorfeed. “Whatwewitnessedwasamajorecologicaldisturbanceeventequaltoafireorahurricane,”saidFrankmazzotti, awildlife ecologist andassociateprofessorattheUniversityofFlorida.“AlotofthingshavehappenedthatnobodyhasseenbeforeinFlorida.” thecoldtemperaturesalsoaffectedmanyof themarqueenames in the state’s invasivespecieszoo.“Anecdotally,wemighthavelostmaybe half of the pythons out there to thecold,”saidscotthardin,theFloridaFishandWildlife Conservation Commission’s exoticspeciescoordinator.“Iguanasdefinitely.Fromacollectionofobservationsfrompeople,morethan50percent fatalityonGreen Iguanas.”

Catatoniciguanaswerereportedfallingfromtreesandlitteringwalkwaysandcanalsinastateofsuspendedanimation. thecoldsnapplayedintoahighlypoliti-cizeddebateoverhowtopreventnon-nativespe-ciesfromcolonizingtheUnitedstates.ReptiledealersandhobbyistsstronglyopposeaproposalbytheU.s.FishandWildlifeservicetobantheimportofandinterstatetradeinburmesepythonsandseveralotherlargesnakes.theysaysouthernFlorida’scoldsnapshowsthatthesespeciesdon’tthreatentospreadnorth,assomeclaim,andafederalcrackdownisunnecessary. “pythons are tropical animals,” saidAndrewWyatt,presidentoftheUnitedstatesAssociationofReptileKeepers.“Whentem-peratures fall below a certain level, they areunabletosurvive.Itreinforcestheideathatthepythonscan’texistmorethanashortperiodoftimenorthoflakeokeechobee.eventhepythons in the everglades are dying duringthecoldsnap.”Wyattsaidthatscientistsaredownplayingtheeffectofcoldweatheronthepythonsbecausethatwouldunderminetheirabilitytowingrantstostudyaproblemthathasreceivedinternationalpublicity. no one knows how many burmesepythonsliveintheeverglades,wheretheywerereleasedasunwantedpetsorwheretheyfoundrefugeafterhurricanesdestroyedtheirbreedingfacilities—butwhat’scertainisthatalotfeweraretheretodaythanamonthago. GregGraziani,apoliceofficerwhoownsa reptile breeding facility, is one of severallicensedpythonhunterswhostalksthesnakesin the everglades. In cold weather, Grazianisaid,pythonsgointoacatatonicstate,andiftheydon’tmakeit toasafeplacetorideouttheweather,freezetodeath.“We’refindingthesmallerpythonsarehandlingitbetterthanthelargeones—thesmalleronescangetintodiffer-entcracksandcrevicestomaintainthetempera-turestheyneed.”

AdaptedfromarticlesintheFort Lauderdale Sun-Sentinel,The Miami Herald,

andThe Orlando Sentinel

AfricanDwarfFrogs(Hymenochirus boettgeri)havebeenimplicatedinamultistateoutbreakofSalmonellainfections.

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GreenIguanas(Iguana iguana)wereliterallyfallingfromthetreesinsouthernFlorida.Unabletoreviveafterthesustainedcoldtemperatures,theymadetheircontributiontotheworkloadofthestate’s“clean-up”crewofvulturesandotherscavengers.

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AlabradorRetrieverhasbeentrainedtotrackille-gallyintroducedRed-earedsliders(Trachemys scripta elegans)inWhitehorse,Australia.

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o b I T u a r y

theconservationcommunitysustainedagreatlosson14February2009.Johnthorbjarnarsondiedtragically innewDelhi,Indiaof

advancedfalciparummalarialikelycontractedinUgandawhilework-ingtosavetheanimalsheloved.JohnservedasseniorConservationofficeroftheWildlifeConservationsocietysince1993.healsowasanAdjunctAssistantprofessoratColumbiaUniversity. Johnt.,ashewasbetterknowntofriendsandcolleagues,wasprimarilyknownandrespectedasoneoftheworld’spremiercrocodilianbiologists;hisresearchencompassedreproductiveecol-ogy,diet,feedingbehavior,movementpatterns,habitatuse,socialbehavior,andpopulationdynamics.however,researchforhimwasnotpurelyanacademicendeavor.heutilizedhisfindingstohelpdevelopconservationprogramsforendangeredspecies.hisapproach,basedonthesustainableuseofreptilesthroughcommu-nity-basedprograms,wasbothinnovativeandeffective. “Whatarealtragedythisis,”saysGrahameWebb,chairoftheIUCnCrocodilespecialistGroup.“Atapersonallevel,Jtwastrulyadmiredandafriendofsomany.Ataprofessionallevel,youjustcannotreplacepeoplelikeJt...Idonotbelievetheworldwilleveragainallowindividualstolearnsomuchaboutcrocodilesinthefieldandthencraftthatknowledgeoveryearssoitbecomeseffectiveatthefrontline...hewasoneofthemostcarefulofscientists,whothroughWCswasdoingmoreoncrocodileconservationaroundtheworldthananyotherindividualhasdoneinthepast—orwilldointhefuture.” Johnalsowill bemournedbyhis colleagues in chelonianconservation. he was a major contributor to the book Turtle ConservationbymichaelKlemens,buthewillberememberedbestforspearheadingtheorganizationofaworkshopinChinatobegindevelopmentofastrategicplanforsavingRafetus.thisworkshop—andtheresultingChinasoftshellturtleguide—ultimatelyledtothediscoveryoftheChangshaZoofemaleandinvolvementoftheturtlesurvivalAlliance(tsA).Johnalsoco-authoredaseminal2003paperinChelonian Conservation and BiologythatprovidedathoughtfulanalysisrankingAsiancountriesinorderoftheirimpor-tanceforturtleconservation.thatpaperessentiallybecametheorganizingprinciplebehindthetsA’sAsianfieldprogram. AndersRhodin,chairoftheIUCntortoiseandFreshwaterturtlespecialistGroup,eloquentlyreflectedonJohn’spassing:“IechothesentimentsofothersasIstoptoreflectonthededicationtoconservation,crocodiles,andturtlesthatJohnhad,andhowhewasalwaystravelingtotheendsoftheearthandintowildernessregionstopursuehispassionandwork.hisenergyanddevotionwillindeedbesorelymissed…JohnlovedhisworkandI’msurehewouldnothavechangedwhathedid.” Johnisperhapsbestdescribedinthewordsofhisnephew,writerAndrimagnason:“hewasascientist,specializedincold-bloodedanimalsbuthimselffullofwarmthwithastronghumantouch—hecouldactasapeacekeeperbetweenpeopleandthe

creaturestheyfearedthemost,hecouldeliminateprejudiceandcreateunderstandingforthegracefulbutunpopularcreaturesinthecrocodilefamily.hecouldgetpeopletounderstandthatacroco-dileisahealthysigninanecosystem—notsomekindofapesttobeexterminated.bydestroyingthehabitatofthecrocodile,thewetlands,swamps,andrivers,peoplewouldeventuallyharmtheirownexistence.hewasrealisticandunderstoodthatpeopleneededasourceofliving—andbypromotingsustainablehunting,thelongtermbenefitofaspeciescouldbesecured.” John’spassingleavesatremendousvoidinthelivesofthosewithwhomheworkedandintheprojectshespearheaded.hewasapassionateadvocateforconservationandissimplyirreplaceable.hislegacywillbetheremarkablenumberofyoungandaspiringbiologiststhathetrainedandmentored.hisworktrulyspannedtheglobewithprojectsinsouthAmerica(mostnotablybrazil),Asia(includingpioneeringworkinburma,China,andCambodia),Africa,andtheCaribbean(especiallyCuba).Aspecialsessionpay-ingtributetothelifeofJohnt.isbeingplannedfortheupcomingtsAconferenceinorlando.

John Thorbjarnarson (1957–2010)1

1AdaptedfromobituariesandremembrancesbyRickhudson,Chuckshaffer,andAndrimagnason.

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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

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

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.

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theresponsetoourcall-for-papersforthisspecialtributetoDr.henrys.Fitch,whichwillextendintoportionsofthenexttwoissuesofReptiles & Amphibians,wasoverwhelming.ItspeakseloquentlytotheinfluenceDr.

Fitchhadontheherpetologicalcommunityandtothehighesteeminwhichhewasheldbyhisstudents,colleagues,acquaintanceswhofrequentlycalleduponhimforadvice,andthosewhomerelyreadhismanyseminalpublications. RobertW.henderson,whostudiedunderDr.Fitch,andGadperry,whosteppedupfromhisusualroleasassociateeditor,servedasco-editorsforthistribute.theyreviewedorfacilitatedthereviewofallcontributions,solicitedcontentandimages,andprovidedvaluableinsightsthroughouttheprocess. WethankAliceFitchechelleforhersupportandhelpinprovidingphotographs.Dwightplatt,oneofDr.Fitch’searlieststudents,sharedcommentshepresentedatthe36thAnnualmeetingoftheKansasherpetologicalsocietyinnovember2009.Joeandsuzannel.Collins,CenterfornorthAmericanherpetology,publicizedthecall-for-papersanddonatedanumberofimages.scottA.schaefer(Copeia),Roberthansen(Herpetological Review),andtheauthorsgavepermissiontoreprintcontentoriginallypublishedinthosejournals.mostly,however,weweregratifiedbytheresponsetoourtributetoDr.Fitchandhislegacy—andtakethisopportunitytothankallofthecontributors,includingthosewhoseworkwillappearinlaterissues.

The Editors of Reptiles & Amphibians

Editors’ Remarks

64 IRCFReptIles&AmphIbIAns•Vol17,no1•mAR2010 FoCUsonConseRVAtIon

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

rescue Darwin’s frogs

Whileonhisfamousvoyagearoundtheworld,CharlesDarwincollectedthefirstspecimensofwhatlaterbecameknownasDarwin`sFrog(Rhinoderma darwinii)(seethearticleinReptiles & Amphibians16(4):246–255). theselittlefrogshaveauniquereproductivebiology,inwhichtadpolesdevelop

inthevocalsacofthemaleuntilmetamorphosis.thegenuscontainsasecondspecies(R. rufum), inwhichthemaleskeepthetadpolesforashorterperiodbeforereleasingthemintosmallstreams. Rhinoderma rufum hasnotbeenseenformorethantwodecadesandmightbeextinct.thehabitatofR. darwinii, thecoolValdivianrainforestsofeasternpatagonia,isvanishingquicklyasaconsequenceofdeforestationanddamconstruction.thechytridfungus,whichisknowntocauseextinctioninmanyamphibianspecies,hasbeenfoundinwildpopulations.theoutlookforthesespectacularfrogsisdire. In2002,europeanzoos,privatekeepers,scientists,andconservationorganizationsandagenciesinChileandeuropeinitiatedacollaborativeconserva-tionprojectforRhinoderma.theproject,ledbytheGermanherpetologicalmagazineReptilia,themuseumAlexanderKoeniginbonn,andtheZoologicalsocietyfortheprotectionofspeciesandpopulations(ZGAp),instigatedeffortstoprotectthesefrogs:(1)surveystosearchforR. rufum,(2)a6-yearfieldstudyofselectedpopulationsofR. darwinii,(3)in-situandex-situhusbandryandbreedingofR. darwinii,(4)statussurveysofwildpopulationsofR. darwinii andscreeningforthepresenceofthechytridfungus,(5)constructionofalabandbreedingstationbyZooleipzigattheUniversityofConcepcióninChile,and(6)supportforotherprojectsregardingRhinoderma.

tosupportthisdifficultandexpensivework,pleasecontact:heikoWerning,RedaktionReptIlIA,seestr.101,13353berlin,Germany(e-mail:[email protected]).

Although Dr. Fitch is probably best known for his five decades of work on the herpetofauna of northeastern Kansas, he spent considerable time in the neotropics, where one focus of his work was the exploitation of black Iguanas (Ctenosaura similis; illustrated) and Green Iguanas (Iguana iguana) in Central America (see also the photograph on p. 14).

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Documents

IRCF Reptiles & AmphibiAns · IRCF ReptIles & AmphIbIAns • Vol 17, no 1 • mAR 2010 In thIs Issue male calling sites of the Australian toadlet (Uperoleia laevigata) are less densely