IRCFREPTILES&AMPHIBIANS•21(4):108–115•DEC2014

Copulation and Oviposition in the Green-blotched Giant Anole,

Anolis smallwoodi (Squamata: Dactyloidae), from southeastern CubaYasel U. Alfonso1, Veronika Holanova2 and Kenneth L. Krysko1

1FloridaMuseumofNaturalHistory,DivisionofHerpetology,UniversityofFlorida,Gainesville,Florida32611,USA(YUA:anoles1983cuba@gmail.com,KLK:kenneyk@flmnh.ufl.edu)

2DepartmentofZoology,FacultyofScience,CharlesUniversityinPrague,CZ–12843Praha2,CzechRepublic(igu@centrum.cz)

Abstract.—Crown-giantanoleecomorphshaveevolvedindependentlyacrosstheGreaterAntilles(Cuba,Hispaniola,Jamaica,andPuertoRico),butonlyafewofthesespecieshavewelldocumentedlifehistories.Hereinwereportpre-viouslyundocumentedobservationsofcopulation,oviposition,andhatchlingsfortheGreen-blotchedGiantAnole(Anolis smallwoodi)ofsoutheasternCuba.Weobservedeightmatingevents(threewithA. s. smallwoodi and five with A. s. palardis)alongthecoastlinebetweenSantiagodeCubaandGuantánamoProvinceduringJune2010andSeptember2011.OnefemaleA. s. palardis matedwithtwomalesinthesameterritory.Femalesdigaholewiththeirsnouts,ovipositsingleeggs,andcoverthemwithsubstrate.Wemonitoredeighteggsincubatedinslightlymoistvermiculite.Theminimumincubationtimeswere54–69days(varyingwithtemperature).Juvenilesusuallyarecoloredlikeadults,brightgreenwithwhitespotsonthebody,palepostlabialbands,andsuprascapularstripes.

Key words:crown-giant,copulation,oviposition,hatchling

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IRCF REPTILES & AMPHIBIANS • VOL15, NO 4 • DEC 2008 189TABLE OF CONTENTS

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

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

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

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

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

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

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

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

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

H U S B A N D R Y

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

P R O F I L E

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

C O M M E N T A R Y

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

B O O K R E V I E W

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

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

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

Back Cover. Michael KernTotat et velleseque audant mo

estibus inveliquo velique rerchil erspienimus, quos accullabo. Ilibus

aut dolor apicto invere pe dolum fugiatis maionsequat eumque

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

IRC

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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

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Studies of Anolis lizardshaveindicatedanimportantrolefornaturalselectionactingonmorphology(Lososand

deQueiroz1997,CalsbeekandSmith2007)andphysiol-ogy (Calsbeek and Irschick2007,Calsbeek et al. 2008).Differencesinbodysizearethoughttoreflectdifferencesinmicrohabitatuse,andareimportanttraitsintheecomor-phologicaldiversificationofanoles(ButlerandLosos2002,Butleretal.2007).Environmentalfactorssuchastempera-ture,photoperiod,precipitation,relativehumidity,andfoodavailabilityare important in thecoordinationof seasonalreproductiveprocessesandalsomayhelpensurethatanoleswillhatchatthemostappropriatetimeofyeartogivethemtheirbestchanceatsurvival(Domínguezetal.2010). Anolislizardshavelowfecundity,withoneeggperovipo-sitionandseveralclutchesperseason(reviewedinHendersonandPowell2009).However,ofthe64currentlyrecognizedspeciesofCubananoles(Rodríguez-Schettinoetal.2013),thereproductivecycleisknownforonlyseven(A. allisoni, A. allogus, A. homolechis, A. bartschi, A. lucius, A. porcatus, A. sagrei;seeDomínguezetal.2010andreferencestherein).Regardingoviposition,someauthorshavedescribedintraspe-

cific(NovoRodríguez1985,EstradaandNovoRodríguez1986;Silvaetal.1982;Estrada1987,1993;Robinson2014)andinterspecific(Alfonsoetal.2012)communalsites. Crown-giantanoleecomorphshaveevolvedmultipletimesacrosstheGreaterAntilles(Cuba:Anolis baracoae, A. equestris, A. luteogularis, A. noblei, A. pigmaequestris, A. small-woodi;Hispaniola:A. baleatus, A. barahonae, A. ricordii; Jamaica:A. garmani;andPuertoRico:A. cuvieri, A. roosevelti; seeRodríguez-Schettino1999,HendersonandPowell2009,Losos2009).Anolis smallwoodiSchwartz1964(Fig.1A–C)hasbeenfoundinhabitingbothlowlandanduplandareas,semi-deciduous,evergreen,gallery,andrainforests,man-groves,andcoffeeplantations(Rodríguez-Schettino1999,HendersonandPowell2009).Thisspecieshaslargehomerangesandmultipleindividualsarerarelyobservedtogether(FläschendrägerandWijfels1996,2009);however,malesandfemalescanbeobservedtogetherduringthebreedingseason(VH,pers.obs.;Fig.1D).Colorationinnaturecanvaryindi-viduallyandgeographically(Rodríguez-Schettino1999),andoccasionaldeep-greenindividualshavebluespotsonthedor-sum(Fig.1E).

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Copulation time,oviposition,orhatchinghavebeenreported for someGreaterAntillean crown-giant species(summarizedinHendersonandPowell2009).Inthispaper,ouraimistodescribethesenaturalhistorydataforAnolis smallwoodi from semi-aridmicrohabitats in southeasternCuba.

Materials and MethodsStudy area and focal populations.—WeconductedfieldworkfromJune2010toSeptember2011at1000–1500halongthecoastlinebetweenSantiagodeCubaandGuantánamoProvincesinsoutheasternCuba.Dominantvegetationcon-sisted of xerophytic scrub,microphyllous evergreen for-

IRCFREPTILES&AMPHIBIANS•21(4):108–115•DEC2014ALFONSOETAL.

Fig. 1. Anolis smallwoodi smallwoodionthesoutheasterncoastofCuba.A:Adultfemaleperchingface-downonatreetrunk.B:AdultmalejumpingonabranchofDichrostachys cinerea.C:Adultfemalecapturedonatreetrunk,arrowindicatethebluenessofthenuchalcrest.D:pairedcoupleduringthebreed-ingseason.E:Adultfemale,notethestrongyellowcolorationonthehead,bluecolorationaroundthesuprascapularstripe,andbluespotsalternatingwiththebrightgreencolorationondorsum.PhotographsfromsoutheasternSantiagodeCubaprovince:Verraco(A,C–E)andSiboney-Jutici(B).PhotographsbyVeronikaHolanova(A,C–E)andYaselU.Alfonso(B).

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est,drymicrophylousforest,andcoastalhyperxeromorphicsemidesertscrub(CapoteandBerazaín1984,Reyes2006).Wesurveyed four localities formatingbehavior (copula-tion)inAnolis smallwoodi (A. s. smallwoodiinSantiagodeCubaProvince:Juraguá,19.936805ºN,75.670188ºW,36masl;Verraco,19.907291ºN,75.575775ºW,23masl;A. s. palardisinGuantánamoProvince:Baitiquirí,20.030244ºN,74.880822ºW,25masl;YacaboAbajo,20.052266ºN,74.699741ºW,11masl). Inmostinstances,weobservedcopulationwhileusingamonocularNikonFieldscopeatdistancesof13–37.4m.Wedescribematingbehaviorinvolvingthreephases:precopula-tion,copulation,andpostcopulation.Wemeasuredthedura-tionofeachstagewithadigitalchronometer(±0.1s).Meansarepresented±onestandarddeviation.

Ovipositioning behavior in captivity.—Captive individualsobservedduringovipositionbelongtotheprivatecollectionofVeronikaHolanovainPrague,CzechRepublic.Anoles weremaintainedinpairsinsideterraria(50x50x80cm)withapproximately5cmofmixedlignocelandsand(propor-tion2:1)asasubstrate.Brancheswereplacedverticallyandhorizontallyinsideterraria.Giantanolespreferwide,verti-terraria.Giantanolespreferwide,verti-Giantanolespreferwide,verti-calbranchesfordailyactivitiesandthin,horizontalbranchesforsleepingatnight(VH,pers.obs.).Liveplants(e.g., Ficus benjamina)fromtheleavesofwhichanoleslickwateralsowereplacedinterraria.Thebackwallofeachterrariumwascoveredwithcorkbark. Temperaturewasregulatedwith40-wattbulbsinceramicshades.Lightwasprovidedwithfluorescenttubes(2x25watt)above the terraria.Daytimetemperatures rangedbetween26–28°C(35°Cdirectlyunderthebulb)andnighttimetem-peraturesneverdroppedbelow22°C.Terrariaweremisteddailywithwater.Eachovipositedeggwasweighed(Ew)andmeasured(EL:egglength;EW:eggwidth).Afterhatching,neonatesweremeasured(TL:totallength,SVL:snout-vent-length,HL:headlength)andweighed(NW:neonatalweight).Measurementsweretakenwithadigitalcaliper(MitutoyoCD-20DC,Sakato,Japan;precision,±0.01mm)andmasswithaKern440-33Nelectronicbalance(±0.01mg).

ResultsCopulatory behavior.—Weobservedeightmatingevents,threeinvolvingA. s. smallwoodi and five A. s. palardis.DuringtheMay-to-Septemberbreedingseason,mostobservationswereofpairs(usuallyasinglemaleandfemaleonthesametree).Malescopulatedexclusivelywiththosefemaleslocatedwithintheirterritories(YUAandVH,pers.obs.).Onlyonefemale A. smallwoodi palardis matedwithmultiple(two)malesinthesameterritory(seedetailsbelow). Malesinitiatedtheprecopulatoryphasebycloselyfol-lowingreceptivefemalesthatusuallysubmittedtoamale’s

advanceswhenheclimbedontoherback.Atthatpoint,malesmadeasuddenlateralmovementandgraspedthefemale,effectivelyrenderingherimmobile(Fig.2A).Themale’sgriponthefemale’sdorsumdiffersbetweenthetwosubspecies;it is middorsal for A. s. smallwoodi(Fig.2B)andmoreante-rior for A. s. palardis(Fig.2C).Subsequentpreparationforpenetrationwasexpeditious,withmaletailandhindlimboverlappingthefemale’stail(Fig.2B–C).Precopulationtook7.11±3.78min(A. s. smallwoodi)and5.47±3.5min(A. s. palardis).Themale’sgraspofthefemalewasmaintainedthroughoutcopulation. Thecopulatoryphasebeganwhenfemalesrespondedtointromissionbymakinganabruptmovement.Matingwassimilarbetweensubspecies;neithermovedthebody,althoughfemalesperformedsomeslowlateralheadswings.Copulatory durationwas similar in both subspecies (A. s. smallwoodi:44.2±23.5min;A. s. palardis:40.6±20.1min).Thepostcopulatoryphasebeganwiththeseparationthatoccurswithoutanysuddenmovements.Femalesiniti-atedseparationwithelevationoftheirhindlimbstofacilitateextractionofthehemipenis.Afterseparation,lizardsinvari-ablymovedupwarduntiloneorbothwereconcealedamongthe leaves. WeobservedonecaseofmultiplecopulationofAnolis smallwoodi palardisinvolvingapair223cmhighonatreetrunk.Matingbehavioranddurationweresimilartoothersobservedforthissubspecies.However,afterseparation,theadultmale,seeminglyunawareofthepresenceofanotheradultmale,movedhigherintothecanopywhilethefemaleremainedinthesameinitialpositionwithoutanymalepro-tection.Anearbymale(ca.420cmawayand137cmabovetheground in abush)opportunistically entered the firstmale’sterritoryandsuccessfullycopulatedwiththefemalethat apparently remained receptive.Theperiodbetweenseparationafterthefirstcopulationandthebeginningofthesecondcopulationwas123min.Matingbehaviorforthesec-ondmatingwassimilartoothersobservedforthesubspecies,butdurationswererelativelyshortforbothprecopulation(2.5min)andcopulation(29.7min).Afterseparation,thefemaleclimbedupwardintothecanopyandthesecondmalejumpedfromthetrunktoanothertree.

Oviposition behavior in captivity.—Paired males and females (allAnolis smallwoodi smallwoodi)initiatedcourtshipbehaviorinMarch.Afterashort“chase,”copulationlasted10–30min.(Fig.3A–B).Femalesreadytolayeggsdescendedfromelevatedperchestothesubstrate,wheretheydugafew“test”holesappar-entlysearchingforasitewithoptimaltemperatureandhumid-ity.Whensatisfiedwithalocation,afemaledigsaholewithhersnout,laysaneggandrollsitinsubstratematerial(Fig.3C).Whentheeggiscovered,sheburiesitusinghersnoutandfore-limbsbeforeclimbingontoanelevatedperch.Theentireprocess

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takesabout20min.Malesusuallycopulatewithfemalesthesamedayshelaysanegg.Femalesalwayslayasingleegg(Fig.3C).Measurementsforeighteggswere2.59±0.45gand2.54±0.12cmby1.33±0.08cm.Well-fedfemalescanlayaneggeverysevendaysduringtheMarch–Septemberbreedingseason. Eggswere incubated in slightlymoistvermiculiteattemperaturesof27–29°C.Theminimumincubationtime

was 54–69 days (varyingwith temperature).Three eggswereinfertile,butmeasurementsforfourhatchedneonateswere2.6±0.11gand13.8±0.32cm(SVL=4.61±0.15cm,HL=1.54±0.03cm).Juvenilesusuallyarecoloredlikeadults,brightgreenwithwhitespotsonthebody,palepostla-bialbands,andsuprascapularstripes.Webeganfeedingneo-natestwodaysafterhatching.Theyacceptedawiderange

Fig. 2.MatingbehaviorinAnolis smallwoodiinsoutheasternCuba.Arrows represent the male graspposition.A–B:A. smallwoodi smallwoodiandC: A. smallwoodi palardis. PhotographsbyYaselU.Alfonso(A–B)andJosephBurgess(C).

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ofinsects(e.g.,crickets,grasshoppers,flies,andbeetlelarvae(Fig.4C–D).

DiscussionCopulatory behavior.—Reproductivebehaviorisnotwelldoc-umentedforGreaterAntilleancrown-giantanoles.Reviewsby Rodríguez-Schettino (1999),Henderson and Powell(2009),andLosos(2009)includedpertinentinformationforonlysomespecies(i.e.,Anolis baleatus, A. baracoae, A. baraho-nae, A. cuvieri, A. equestris, A. garmani, and A. pigmaequestris).ThematingbehaviorofthreeCubancrown-giantsforwhichwehavesomeinformation(A. baracoae, A. luteogularis, and A. noblei)issimilarinsomeaspectstothatofA. smallwoodi. Allhavebeenobservedinpairs(usuallyonemaleandonefemaleonthesametree)andprecopulationusuallyisiniti-atedwhenamalefollowsafemalefrombehindandendswiththemalegraspingandlargelyimmobilizingthefemale(YUA,pers.obs.). Ourobservations also suggest thatAnolis smallwoodi exhibitsbehaviorsimilartothatofotherGreaterAntilleancrown-giants (Losos,2009and references therein).Rios-LópezandPuente-Colón(2007)observedmultiplesleepingpairsofA. cuvieriandsuggestedthatthismightbeindica-tive of pair-bonding.Rodríguez-Schettino (1999) notedthatcourtshipandmatinginA. equestris and A. luteogularis occurredintalltrees,andthatonlyoneortwoindividualsofthelatteroccupyasingletree.Apparently,crown-giantshave

largehomerangesandtwoindividualsarerarelyseentogether(FläschendrägerandWijfels1996,2009). Thedurationofcopulationvariesconsiderablyamongsomespeciesofanoles(Jenssen1996).Itcanbelengthyinmanyspeciesand,inatleastsomespecies,durationseemstovaryconsistentlyamongmales(Crews1973,Tokarz1988)andappearstobegreaterlaterinthebreedingseason(Stamps1975,Tokarz1999).InJamaicanA. garmani,Trivers(1976)recordedanaveragedurationof25min.ForotherGreaterAntilleananoles,meandurationvariesfromafewsecondstomorethananhour(e.g.,A. sagrei,3.8min;A. bremeri,64.9min; A. valencienni,2min;A. websteri,±1sec;RegaladoandGarrido1993;HendersonandPowell2009;Losos2009andreferencestherein). Maleanoles(non-crown-giantspecies)usuallymatewithmultiplefemalesintheirterritories,andthosemaleswithlargerterritories(presumablysupportingmorefemales)suf-feredmorepaternity losstoextra-paircopulations(Losos2009andreferencestherein).OurobservationofmultiplecopulationsinonefemaleA. smallwoodi palardismightbeanexampleofcrypticmatechoice,whereinfemalesmatemul-tipletimesoveranextendedperiodandsubsequentlyselectaspecificmale’sspermtofertilizetheegg(Losos2009).FemaleA. garmanipositionthemselvesinexposedplaces,perhapsrenderingmatingmaleshighlyvisibletoothernearbymalesortosolicitmalessothatthefemalemaychooseamongthem(Trivers1976,HicksandTrivers1983).Extra-paircopu-

Fig. 3. A–B:PositionsduringmatinginAnolis smallwoodiincaptivity.C:Femaleburyingarecentlylaidegg(arrow)usingherforelimbs and snout. Note thebluecolorationofthisadultfemale.PhotographsbyVeronikaHolanova.

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lationshavebeenreportedatleastinonecrown-giantspe-cies(A. garmani;Trivers1976).HicksandTrivers(1983)observed49copulationeventsinA. garmani,includingbothresidentmalesandneighboringmales(i.e.,notwiththemainresidentmalewithintheterritory).

Ovipositioning behavior in captivity.—CourtshipbehaviorincaptivityhasbeenreportedforsomeGreaterAntilleancrown-giants(HendersonandPowell2009).Anolis baracoae canmate2–3timesperweek,andcourtshipinvolveshead-bobbing,dewlapextension,andpairedcouplesfacingdownonverticalbranches(HolanovaandHribal2004).Appler(1963)recordedthedurationofcopulationforA. cuvieri in captivity;therangeof20–90minismorevariablethanourobservationsofA. smallwoodi.Femaleanolesproduceonlyoneeggatatimewithanaverageof5–25daysbetweeneggs(AndrewsandRand1974,Andrews1985).Onlyasingleegg,usuallyfromalternatingovaries,isovulatedatatime,

butfemalescanretaineggsduringtimesofdroughtandthussometimescarryasmanyasthreeeggs,twoshelledandoneintheoviduct(Stamps1976). EggdepositionreportedhereinforAnolis smallwoodi is similartothatdescribedforotheranoles.Femalesusuallybeginbyselectingapotentialsite,probingthesubstratewiththesnout,diggingaholeintowhichaneggisdeposited,andfinallycoveringtheeggwiththesubstrate(Losos2009andreferencestherein).Forcrown-giants,Rodríguez-Schettino(1999)observedafemaleA. equestrisovipositinginaholenearthebaseofatree,andPanton(1928)describedafemaleA. garmanidescendingatreeatmidday,diggingasmallholewithhersnout,andlayinganegg. Eggandhatchlingsizes,incubationtemperatures,andminimumincubation timeshavebeenreported for somecrown-giants(Table1).IncubationtimeforseveralCaribbeanspeciescanrangefrom3.5–6weeksunderlaboratorycondi-tions(GreenbergandHake1990,Sangeretal.2008).Our

Fig. 4.A:NeonateAnolis smallwoodi smallwoodi.B:Holdinganeonateoutsidetheterrarium.C:AdultfeedingonlarvaeofZophobas morio(Tenebrionidae).D:AdultfeedingonlarvaeofPachnoda marginata(Scarabaeidae)insidetheterrarium.PhotographsbyVeronikaHolanova.

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resultsaregenerallysimilartothosepreviouslyreportedforCaribbeanspecies,althoughBech(1986)observedonecap-tive female A. equestris laying12eggsinoneyear(comparedtooneeggeverysevendaysduringbreedingseasoninA. smallwoodi).

AcknowledgementsWe thank Addael C. Casenave-Cambet (undergradu-ate student atUniversityofOriente) andTeresa Imbert(CATEDES/CITMA)fortheirassistanceinthefield.M.S.JavierTorres (UniversityofHavana) andananonymousreviewerprovidedhelpfulcommentsandsuggestionsonanearlierdraftofthismanuscript.

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Table 1.Meanovipositiondataofcrown-giantanolesintheGreaterAntilles.MIT=minimumincubationtime,IT=incubationtempera-ture,SVL=snout-ventlength.Anasteriskindicatesdataobtainedfromourstudy.NeonateSVLsaregivenbasedontotallength.RemainingdataarefromLosos(2009)andHendersonandPowell(2009),whichshouldbeconsultedfortheoriginalsourcesineachcase.

Eggs Hatchling Length Width Mass MIT IT SVLSpecies (mm) (mm) (g) (days) (ºC) (mm)

Anolis smallwoodi* 23–27 12–14 1.7–3.1 54–69 27–29 134–142

Anolis baracoae 23–25 6–7 — 74–81 26 130

Anolis cuvieri 21–23 11–12 — 95 20–25 41

Anolis equestris 23–25 12.8–18 — 60–92 18–30 80–140

Anolis garmani 12–15 8–11 — 60–75 24–26 27

Anolis pigmaequestris 23.5 13.3 — — — —

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