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Araneae Biodiversity in the Amazonian Foothills: An analysis of Araneae Biodiversity in the Amazonian Foothills: An analysis of

family and guild presence across habitat types family and guild presence across habitat types

Rae Lewark SIT Study Abroad

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Recommended Citation Recommended Citation Lewark, Rae, "Araneae Biodiversity in the Amazonian Foothills: An analysis of family and guild presence across habitat types" (2019). Independent Study Project (ISP) Collection. 3265. https://digitalcollections.sit.edu/isp_collection/3265

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AraneaeBiodiversityintheAmazonianFoothillsAnanalysisoffamilyandguildpresenceacrosshabitattypes

Lewark,RaeAcademicDirector:Xavier,SilvaPhD

ProjectAdvisor:GermaniaEstevez,PhDFieldAdvisor:AlexBentleyHawaiiPacificUniversityEnvironmentalStudies

SouthAmerica,Ecuador,PastazaProvince,EcomingaRioAnzuSumakKawsayinSitu

SubmittedinPartialFulfillmentoftherequirementsforEcuador:ComparativeEcologyandConservation,SITStudyAbroad,Fall2019

AbstractTheactivityoftheorderAraneaewithinecosystemsisoftenconsideredanecologicalmystery.Consideredtop-leveltrophicpredators,spidersactasregulatorsofinsectpopulationsandhaveinfluenceuponthefoundationofthetrophicweb.IntheEcuadorianAmazonianfoothills,itisimportanttoaddressthelackofknowledgeofthebiodiversityofspidersintheregion.Conductingbiodiversitystudiesacrosslandusegradientscanbegintocharacterizeanthropogenicimpactsonneotropicalspiderspecies.ThisstudyquantifiesthearaneaebiodiversityinthetransitionbetweentheCloudandAmazonforestsinEcuador.Populationsinprimaryforest,secondaryforest,cultivatedfieldsandcavesystemswereanalyzedtoidentifythefamiliesandguildspresentintheecosystems.Thebiodiversityofspiderfamiliesandguildtypewerefoundtodiminishwithincreasinglanduseimpactandthedecreaseofvegetationcomplexity.ResumenLaactividaddelordenAraneaeentrelosecosistemasfrecuentementeseconsideraunmisterio.Consideradepredadoresimportantesarañassonreguladoresdelaspoblacionesdeinsectosytieneninfluenciasobrelabasedelaredtrófica.PorlasestribacionesdelaAmazonia,esimportanteaenfocarenlafaltadeinformacióndelabiodiversidaddearañasenelregión.Larealizacióndemásestudiosentrelosgradientesdelusodelatierrapuedeempezaracaracterizarlosimpactoshumanosenlasarañasdelosalotrópicos.EsteestudiocuantificólabiodiversidaddearaneaeenlatransicióndelBosqueNubladoylaAmazoniandeEcuador.Laspoblacionesdebosqueprimario,bosquesecundario,camposcultivados,ylossistemasdecuevasfueronanalizadosaidentificarlasfamiliasylosgremiosquesepresentenenlosecosistemas.Labiodiversidaddelasfamiliasylosgremiosdearañasseencontraronadisminuirconelcrecimientodelusodetierrayconeldecrecimientodelacomplejidaddelavegetación.AcknowledgementsTheauthorwouldliketoextendgratitudetoGermaniaEstevezforadvisementandguidancethroughoutthestudy,toAlexBentleyforadvisementinthefieldandAnaMariaOrtez,XavierSilvaandDianaSerranoforaidthroughoutthedevelopmentandcompletionofthestudy.TothestaffofSumakKawsayinSituandthefellowAnzuteamresearchers,ReneeHeller,TeoCarr,andZaneLibkefortheircompanionship,theirresilienceandtheirconstantabilitytoinspireeveninthehardestconditions.ToalltherepresentativesofSITstudyabroad.IntroductionSpidersrepresentamegadiverseorderwithinthearthropodphylum,with48,262currentlydescribedspecies(TheWorldSpiderCatalog20.5).Spidersposeataxonomicchallenge,asmorphologicalvariationwithinspeciesisabundantandcanbeduetogeneticdeviationorgeographicaldifference(Coddington&Levi1991).Thecurrentnumberofdescribedspeciesisspeculatedtoonlyconstitute35%ofthetotalspiderspeciespresentonearth.Incomparisontoothernon-invertebrategroups,spidersandotherarachnidsrepresentbothalackofresearchandanareafornewdiscovery(Pinzon2010,Brooks2004.)Whileresearchintotheecologicalimportanceofspidersisincreasing,onlyanaverageof605newspeciesaredescribedeachyear.Atthisrateofdescription,itwouldtakemorethan150yearstodevelopasufficientdatabaseontheplanet’sspiderbiodiversity(Platnick2013).Intheneotropics,wherebiodiversityisnotablyhigherthantemperateregions,researchintospiderbiodiversityisneededand

presentsafrontierforthediscoveryanddescriptionofnewspecies.SpidersoftropicalregionssuchasinEcuador,presentanunder-exploredworldofecologicalandphysiologicalpotential(Basset2012.)Spidervenomcontainsneurotoxinsthatarebecomingimportantinneurobiologicalinnovation,studiesintopropertiesofspiderwebforfiberresearch,andalsointhedevelopmentoflessenvironmentallydamaginginsecticides(Coddington&Levi1991).Spidersmaybenefittheeffectivenessofpestcontrolinagroforestry.Increaseddiversityofspiderpopulationshasbeenfoundtobenefitthemitigationofpestimpactoncrops.Arangeofspeciesfromdifferentguildscreatesaneffectivenetworktotargetpestsacrosstheirlifecycles.Thispromoteslandmanagementwithlesschemicalimpactandfocusesonharnessingthenaturalecosystemtopromotecropproductivity(Marc&Conrad1997).Inordertoquantifytheimpactoflanduse,spiderpopulationsmustbethoroughlysurveyed.Themosteffectivetechniqueforspidercollectingis“nocturnalhandcollecting.”Thisisduetohigherprobabilityofencountersinthenighthourswhenmanyspeciesaremorelikelytobehunting.ThisstudyfocusedonhandcollectingasthemainsamplingmethodasadvisedbyRego(2009)whenothermethodsarenotavailable.Thismethodwasthefocusofthestudyinordertoencounterarangeofspidersacrossguildsandvegetationtypes,aswellastoobserveeachindividual’secologicalinteractionsandbehaviorwhileintheirnaturalenvironments.Nightcollectingsessionswereavitalcomponentofthestudyasspidersareoftenobservedinhigherquantityinhoursofnolightintheneo-tropics,withsomespeciesexclusivelyrecordedasnocturnallyactive(Green1999).Thisisoftenduetospider’smainpredatorsbeingactiveduringdaylighthours,andinordertohuntspidersmustputthemselvesatrisk.Buildingaweborambushhuntingdemandshighlevelsofenergyandrequiresindividualstoleavelocationswheretheyaresafefrompredators(Venner2005).Huntinginlowornolighthoursdecreasestheriskofpredationandalsoincreasestheindividual’spotentialforcatchingunsuspectinginsectswhomaynotbeabletodetectthespider’spresenceaswellatnight(Rypstra1986).Spidersareconsideredtopleveltrophicpredatorswithintheirhabitats.Spidersfilltheecologicalregulatorrolewithintheirecosystems,withthecapacitytodictatethecompositionandbalanceofinsectpopulations(Martin1982).Throughouttheplanet’secosystems,predatorsaredistinguishedaskeystonecomponentsinoverallhealth.Thedisproportionatelackofresearchintospider’simpactonecosystemfunctionalitydrawsastarkcomparisontopredatorsonlargertrophiclevels.Whilesmallinphysicalsize,spidershavethecapacitytoactasimportantcontrolagentsinthebalanceofinsectpopulations(Silva1992).Spiderfamiliespresentadiverserangeofguilds.Guildsareclassifiedasgroupsofspecieswhocompeteforthesameresourcesinanecosystem.Identifyingtheguildspresentinanecosystemcanleadtoabetterunderstandingoftheecologicalimpactspidercommunitiesarehavingonthetrophicweb.Classifyingguildscanalsodevelopanunderstandoftheimpactoflanduseinahabitat.ThedescriptionsofspiderguildsbyCardosoetal(2011)characterizedspiderguildsby8definitions:sensingwebweavers,sheetwebweavers,spacewebweavers,orbwebweavers,specialists,ambushhunters,groundhuntersandotherhunters.

Thestratosphericrangeofaspidercanhelpidentifythetaxonomyaswellastheguilditbelongsto(Yanoviak2013).Spidersthatrelyonspecificvegetationstructureforwebbuildingorforhuntingwillbemorevulnerabletochangesinvegetation.Spiderswhorelyoncamouflageforhuntingmayalsobenegativelyimpactedbychangesinvegetationtype(Thery2002).Spiderswillutilizevegetationstructuretobuildwebs,forage,oruseambushhunting.Webbuildingspiderswillelectlocationsofhigherinsecttraffic,inlocationswheretheycanattractspecificindividuals,orlocationswheretheycanutilizesurprisetoentrapprey(Venner2005).Thisstudycollectedspecimensacrossfourdistincthabitats.Withineachhabitata200mtransectorientedaroundabodyofwateranda200mtransectinadryregionweresampled.Thefourecosystemssampledwereprimaryforest,secondaryforest,cultivatedfields,andsubterraneancaves.Thefirstthreeecosystemtypeswereselectedinordertocomparethebiodiversitygradientacrosstheregionwithincreasinganthropogenicimpact.Thecavetransectswereselectedinordertoobservetheimpactsoftheecologicalpressuresofsubterraneanecosystemsonspiderdiversity(Mammola&Isaia2017).Theprimaryforesttransectswereconsideredregionsunalteredbyanthropogenicactivity,withlittletonodisturbance.Thefirsttransectselectedwassituatedinavalleyandcutthroughthepathofastream.Thesecondtransectselectedwassituatedalongtherideofthesamevalley,andcutthroughtheforest.Bothtransectshadlittletonohumanimpactexceptforcollectingreasons.Thestructureofvegetationhasasignificantimpactonspiders.Astheyrelyheavilyoninfrastructureforhunting,areaswithmorediverseflorawilloftenprovidethefoundationfordiversespiderpopulations(Schüpbach2013).Thesecondaryforesttransectsweredefinedasareaswereprimaryforesthadbeencleared,buthadthenbeenabandonedandsuccessionalgrowthwaspermitted.Thefirsttransectselectedwasorientedclosetoariveralongastreamtributary.Thesecondtransectcutthroughtheforest,andcrossedthedryportionofthefirsttransectsstream.Secondaryforestgrowthisdenserwhilelessdiversethanprimaryforests,andiffoundadjacenttoprimaryforestswilloftenpresentidealconditionsforspiderdiversity(Floren&Deeleman-Reinhold2005).Thecultivatedtransectswerecharacterizedasareaswhereprimaryforesthadbeenremovedtoopenthelandforagriculturalactivity.Thetransectsselectedweredominatedbygrassesandsomesuccessionalforestflora.Cultivatedhabitatsoftenexhibitpatchworklikeregions,withthealtercationsoflandcausingfragmentationofhabitatsforthespeciespresent.Spidersareadaptivepredators,oftenexhibitingresiliencewhenimpactedbyhabitatdisturbance.Oftenabletoadapt,spidersthriveinareaswithhighinsectpopulations,suchasagriculturalsites,orhabitatsaroundbodiesofwater.Theycanadapttheirhuntingmethods,whetherweborientedorambushhunting,toalteredecosystems(Schüpbach2003).Ithasbeenobservedhoweverthatoveralldiversityofspecieswilldecreaseinareasofdisturbanceimpactedbyagricultureorotheranthropogeniclanduse.Thisusuallyresultsinafewspeciesthrivinganddominatingtheecosystem(Cardosoetal2011).

Thecaveswerecharacterizedassubterraneangeologicalformations.Subterraneanhabitatscreateextremeenvironments,exposinginhabitantstohighlevelsofecologicalpressure.Duetolowresourceavailabilityaswellaslowlightlevels,specieslivingincaveecosystemswilloftenevolveawayfromtheirterrestrialcounterparts(Mammola&Isaia2017).ThisstudyaimedtoevaluatethebiodiversityofspidersinthetransitionalhabitatsbetweentheEcuadorianCloudForestandAmazonForest.Bydevelopinganunderstandingofthespeciesintheregionandbysamplingvarioushabitatstypes,theimpactofhumanlanduseonspiderdiversitywasalsoevaluatedacrossincreasinglevelsofaltercation.MethodsIntheFieldThestudywasconductedinNovember,2019inthetransitionbetweenthedryandthewetseasoninEcuador.ThestudysitewasintheRioAnzuregionofthePastazaProvince,intheEcomingaandSumakKawsaybiologicalreserves.Collectionsessionswereperformedfor16daysbetweenthe12thandthe29thofNovember.Thefourhabitatssampledwereprimaryforest,secondaryforest,cultivatedfields,andsubterraneancaves(cavesnotmapped).Foreachtypeofhabitat,excludingthecavesystems,two200mtransectswereused,onethatwasorientedaroundabodyofwater(wettransect)andonethatwasnotimpactedbywater(drytransect)(figure1).Thecavetransectswereevaluatedinsearchhoursinsteadofdistancesurveyed.

Figure1.Mapofthe6maintransects,TCM=cultivatedwettransect,TCS=cultivateddrytransect,TSMIandTSM2=secondarywettransects,TSS=secondarydrytransect,TPM=primarywettransect,TPS=primarydrytransect.

Surveyingsessionsutilizedhandcollectingmethods,withemphasisonidentifyingindividualsaswellasobservingbehaviorandenvironmentalinteractionofeachmorpho-species.Whenanewindividualwasencountereditwasphotographed,theheightabovegroundwhereitwasfoundwasmeasured,andthetypeofvegetationorstratawasclassified.Vegetationorstrataclassificationwasbrokenupinto9categories:gravel/sand,surfaceofwater,soilbank/wall,leaflitter,rootsystems,fallentrees/branches,grass,low-levelfoliageandhigh-levelfoliage.Thepresenceofawebanditstypewasnoted,aswellasbehaviorwhendisturbed,eggsacktype,orothernotablefeaturesthatcouldbeusedinidentification.AsfoundbyAvezedoetalin2014,nocturnalhandcollectingisthemosteffectivemethodforcollectingspiderspecimensaboveothermethodssuchastreebeatingorpitfalltraps.Whileacombinationofthethreemethodsisideal,nocturnalhandcollectingisthemostsuccessfulespeciallywhenotheroptionsarenotaccessible.Thisisduetohigheractivityofspiderspeciesatnightcomparedtotheday(Green1999).Thisstudyaimedtomaptheactivityofdifferentspidergroupsthroughoutdifferentlightlevels,anddevelopedcollectingschedulesinnight,dayandtransitionallightsessions.Thisstudyutilizedhandcollectingastheprimarysamplingtechniquebecausepitfalltrapsandtreebeatingeitherkillsindividualsordisruptstheirnaturalactivity.Thisinhibitsobservationtoobserveoftheirecologicalinteractionandbehaviorintheirnaturalenvironment.Fifteencollectinghourswerespentineachtypeofhabitat,withevenamountsoftimespentcollectingintransitionalhours(sunriseandsunset),daylighthours,andnighthours.Thecavesystemsweresampledforonlyanequivalentofthreesamplingsessions,andnotsubjecttolightlevelspecificsampling.Atotalof45searchandcollectionhourswereperformedinthemainthreehabitattypes,and3hoursand45minutesinthecavetransects.Eachtransectwaswalkedslowly,withsweepingvisualsearchesdonefromsidetosideofthevegetationorwaterway.Leaflitter,soilbanks,fallentrees,rootsystemsandtheundersidesofleaveswerethoroughlyexaminedateachsearchinterval.Nighttimesessionsutilizedbothaheadlampandspottingflashlightinvisualsearches.Samplesweretakenatthreedifferentlevelsoflight.Daylightsamplesweretakenbetweenthehoursof8:00and17:00.Nightsamplesweretakenbetween19:00and0:00.Transitionallightsamplesweredividedevenlybetweenmorningandeveningtransitionhours,withmorningsessionstakingplacewithinthehoursof5:30and7:45,andeveningsessionstakingplacewithin17:15and18:45.Fifteenhoursofcollectingineachlightlevelwereperformedacrossthethreemainhabitats,withtransitionalhoursbrokeninto7hoursand30minutesforthemorningtransitionsand7hoursand30minutesfortheeveningtransitions.TheapplicationiNaturalistwasusedtoorganizedataduringcollectingsessions.Eachindividualwasphotographedinthefieldwithamacro10mmMomentLensattachedtoaniPhoneXR.Ifpossible,photosweretakenofeachindividualfromabovetocapturethelegandbodyoverallshape,alongthedorsalplanetocaptureshapeofthecarapaceandtheabdomen,andfromthefronttocapturetheocularpattern.Whenanewmorpho-specieswasfounditwascollectedtobepreservedandidentifiedoutsideofthefield.

PreservationandIdentificationSampleswerepreservedina70%alcoholsolution,andphotographedwiththesamelensusedinthefield.Amicroscopewasusedforidentificationwhenmacrophotoswerenotsufficientforidentification.Thefamilyofeachindividualwasidentifiedaswellasthegenuswhenpossible.IdentificationwasbasedonthedescriptionsofJocqué(2007),Wegner(2011),Duperre(2013,2015,2016),Brescovit([nodate]),andtheWorldSpiderCatalog20.5(2019).GuildswereassignedtoeachfamilybasedonthedescriptionsofCardosoetal(2011).StatisticalAnalysisThediversitywithineachhabitataswellasthecompositediversityoftheregionwascalculatedusingtheentropyoftheShannonDiversityIndex,andtheinverseSimpsonDiversityIndex.TheShannonDiversityIndexwhichaccountsfortheincreaseofrarespecies,aswellastheabundanceandevennessinthecommunityandfunctionsonanordinalscale.Theentropywasappliedinordertoavoidunbalancedinfluenceofrareorcommongroupsdisproportionately(Jost2006).TheGini-SimpsonDiversityIndexdescribestheoveralldiversityofacommunity,accountingforrichness,evennessanddivergence(Solow1993).Theinverseoftheformulawasappliedinordertoaccountfordominanceofgroupswithinthesample(Jost2006).TheShannonIndexiscalculatedasfollows,where(p1)isthequantityofspecies“1”inrespecttothetotalnumberofspecies.

= 𝑒𝑥𝑝 &'(𝑝))|ln 𝑝)|.TheGini-SimpsonIndexiscalculatedasfollows,wherenisthetotalnumberofindividualsinaspecificspecies,andNisnumberofindividualsofthecumulativespecies:

= 1

0 1∑(𝑛𝑁)

45

ThesamplecompletenessofeachhabitattypeaswellastheregionasawholewascalculatedusingthesoftwareiNext(Chaoetal2016).Thisisanimportantmeasuretotakewhenstudyingarachnidpopulationsbecausetheyoftenpresenthighnumbersofsingleordoublemorpho-speciesencounters(singletonsanddoubletons)(Rego2009).Aspeciesaccumulationcurvewasmappedinordertocomparethenumberofspeciesfoundinaregioninrespecttotheeffortusedtofindthem(Chaoetal2016).TheJaccardIndexwasusedtomeasurethepercentsimilarityofmorpho-speciesbetweenthethreemainhabitatssampled(primary,secondaryandcultivated.)Theformulaoperatesbydividingthenumberofsharedspeciesbythetotalnumberofspecies,wherenequalsnumberofsharedspeciesandNequalstotalnumberofspeciesbetweenthehabitats.

%𝑠𝑖𝑚𝑖𝑙𝑎𝑟𝑖𝑡𝑦 =𝑛𝑁

Thesamplescollectedfromthecavetransectswerenotstatisticallycomparabletotheotherhabitatsasthesamplingeffortwasdisproportionate.Theresultsfromdatasampledfromthecaveswasincludedinthediscussionportionofthestudy.Results&DiscussionAtotalof1036spiderswerecollectedacrossthefourhabitattypes.Thetotalnumberofindividualscouldbebrokenupinto150morpho-speciesand20families(appendixA).The20familiescouldbeclassifiedinto8guilds(table1).Ofthetotalmorpho-species4couldnotbeidentifiedandfallintotheunknownclassification.

Table1.SpiderFamiliesfoundacrossallhabitatsdesignatedintoguilds.Guild FamilyAmbushHunter Deinopidae

Thomisidae

GroundHunter CorinnidaeHeteropodidaeLycosidae

OrbWeaver AraneidaeSymphytognathidaeTetranathidae

OtherHunter AnyphaenidaeCtenidaeMiturgidaeSalticidaeScytodidae

SensingWeb Theraphosidae

SheetWeb AgelenidaePisauridae

SpaceWeb PholcidaeTheridiidae

Specialist DysderidaeGnaphosidae

Ofthe150morpho-species,onefromeachhabitatwasfoundtodominanttheirrespectivehabitattypes.Themostdominantmorpho-speciesfoundintheprimaryhabitatwasPisauridaegen.sp.1(15%),Thenextmostdominantmorpho-specieswere

Pholcidaegen.sp1.andTetranathidaegen.sp.1makingup12%and11%ofthetotalpopulation,respectively.Themostcommonmorpho-speciesinthesecondaryhabitatwasPholcidaegensp2whichmadeup17%ofthetotalmorpho-speciesfoundinthehabitat.Thenextmostdominatemorpho-specieswereTheridiidaegen.sp.1andPholcidaegen.sp.2whichmadeup17%and13%ofthetotalmorpho-speciesfound,respectively.Themostcommonmorpho-speciesfoundinthecultivatedhabitatwasTetranathidaleucagesp.whichrepresented40%ofthetotalpopulation.ThefollowingdominantspecieswereTheridiidaegen.sp.1,andPisauridaegen.sp.2,makingup11%and8%oftheremainingpopulation,respectively.

Figure2.Thethreemostcommonmorpho-speciesfoundineachhabitattype.Fromlefttoright:themostabundantmorpho-speciesintheprimarytransects,secondarytransects,andthecultivatedtransect.Themostcommonmorpho-speciesinthesecondaryhabitatwasPholcidaegensp2whichmadeup17%ofthetotalmorpho-speciesfoundinthehabitat.Thenextmostdominatemorpho-specieswereTheridiidaegen.sp.1andPholcidaegen.sp.2whichmadeup17%and13%ofthetotalmorpho-speciesfound,respectively.Themostcommonmorpho-speciesfoundinthecultivatedhabitatwasTetranathidaleucagesp.whichrepresented40%ofthetotalpopulation.ThefollowingdominantspecieswereTheridiidaegen.sp.1,andPisauridaegen.sp.2,makingup11%and8%oftheremainingpopulation,respectively.Ofthethreemainhabitats,thecultivatedhabitatspresentedthehighestdominancebyonemorpho-specieswitha40%representationofTetranathidaleucagesp.Thismorpho-speciesisnotablyadaptedtocultivatedenvironments,andareproficienthuntersingrassdominatedecosystems(Hall2019).Similartospiderbiodiversityin

Pisauridae Pholcidae Tetranathidae

othercultivatedareas,themostalteredenvironmenthadthehighestdominanceofasinglemorpho-species(Cardosoetal2011).ComparisonofDiversityTheShannonEntropycalculationsshowedhigherdiversityandevennessofthepopulationfromthecultivatedtotheprimaryhabitatgradient(table2).Anincreaseof61.31%wasfoundbetweenthecultivatedandsecondaryhabitats,andanincreaseof29.02%betweenthesecondaryandprimaryhabitats.Theincreaseof72.54%betweenthecultivatedtoprimaryhabitatswasthemostsignificantchangeinbiodiversity(appendixD).TheTransformedGini-Simpsoncalculationsshowedasignificantincreaseofbiodiversitywhenhabitattypeswerelessimpactedbyanthropogenicaltercation(table2).Anincreaseof67.23%wasfoundbetweenthecultivatedandthesecondaryhabitat,andanincreaseof33.31%betweenthesecondaryandprimaryhabitat.Theincreaseof78.15%betweenthecultivatedandprimaryhabitatswasthemostnotableinchangeinbiodiversityacrossthelandaltercationgradient(appendixD).Table2.Diversityvalues,foreachhabitattyperespectivelyandforallthreecombinedtorepresenttheregionalbiodiversity.Habitat ShannonEntropy Gini-SimpsonTransformedPrimary 3.86 25.58Secondary 2.74 17.06Cultivated 1.06 5.59Regional 2.88 51.91Thepercentsimilaritiesbetweenhabitatsshowedtheprimaryandcultivatedhabitatshadtheleastsimilaritysharingonly10%morpho-speciesbetweenthem.Secondaryandcultivatedshared31%ofthetotalmorpho-species,showingthegreatestsimilarity.Theprimaryandsecondaryhabitatshowedaslightlylowersimilaritywith28%ofthetotalmorpho-speciesshared(table3).

Table3.Jaccard’sIndicesbetweenthe3mainhabitatssampled.HabitatComparison Shared

Morpho-speciesTotalMorpho-species

JaccardIndices

PrimarytoSecondary 30 106 28%

SecondarytoCultivated 34 108 31%

CultivatedtoPrimary 15 143 10%

Habitatswithsimilarlanduseimpactssharedmoremorpho-species.Boththesecondaryandprimarytransectsshowedmoresimilarity,asitcanbeassumedthemorpho-speciesinthosehabitatsrelyonmoreunalteredvegetationstructuretosurvive.Morpho-speciesinthecultivatedregionsmaybeabletocrossovertoboththesecondaryandprimaryhabitats,butmanyoftheprimaryandsecondarymorpho-

speciesmaybetoospecializedtotheirvegetationinfrastructuretosurviveinhighlyimpactedregions(Gollan2010).SpeciesAccumulationandSpeciesCoverageTheindividualsfoundacrossthethreemainhabitatsrepresentedfrom60%to75%ofthetotalexpecteddiversityoftheregion.Thisleavesanunidentifiedmarginfrom35%to40%ofmorpho-speciesnotaccountedforinthesamplingperiod(figure3).

Figure3.SpeciesAccumulationCurveofthethreemainhabitats:cultivated,primaryandsecondary.Theinterpolatedlinerepresentsdatacollectedandtheextrapolatedlinerepresentstheprojectionofspeciesandindividualsnotfoundinthehabitatsduringthestudy(Chaoetal2016).Thesamplingeffortsofthestudysuccessfullyaccountedforaround60%to75%ofthetotalspeciesinthehabitatssurveyed.Thisleavesanunidentifiedmarginfrom35%to40%ofthemorpho-speciesnotfoundinthesamplingperiod(figure4).

Figure4.SampleCoverageofthesurveysacrossthethreemainhabitats:cultivated,primaryandsecondary.Theinterpolatedlinerepresentsdatacollectedandtheextrapolatedlinerepresentstheprojectionofdiversitythatremainsun-surveyedinthehabitats(Chaoetal2016).

Thisresultreflectsthefindingsofsimilarstudieswherearachnidbiodiversitystudiesofthisgeneralsizeareusuallyabletoaccountforaround70%ofthetotalregionaldiversity(Azevedoetal2014).Duetohighvariabilityamongstpopulations,itiscommontoencounterahighnumberofsingletonsanddoubletonswhilesurveyingarachnids(Rego2009).FamilyCompositionTheregionalfamilycompositionshowedthefamilyPholcidaehadthegreatestabundancerepresenting23.36%ofthetotalindividualsfoundacrosshabitats.ThisresultissimilartothefindingsofMoore(2015),whofoundPholcidaewasthemostabundantfamilyacrossaltitudesinanadjacentgeographicalregion.

Figure5.Examplesfromthe3mostabundantfamiliesintheregion,amorpho-speciesfromthePholcidae,TheridiidaeandTetranathidaefamily.ThefamiliesTheridiidaeandTetranathidaewerethesecondandthethirdmostabundantfamiliesrepresenting18.24%and17.95%ofthetotalregionalpopulation(appendixC.)ThefamilyTheridiidaewasfoundtobeabundantinEcuadorianCloudForestsbyRobinson(2018),inastudyonthebiodiversityofspidersacrossaltituderange.

26.20%25.56%

12.14%

8.63%6.07%4.47%3.00%2.88%2.56%2.00%1.28%1.28%1.00%1.00%0.96%0.96%0.64%0.32%

0.00%5.00%

10.00%15.00%20.00%25.00%30.00%

pholcidae

pisaurid

ae

tetran

athidae

theridiid

ae

saltic

idae

ctenidae

lycosid

ae

thomisidae

corin

nidae

heteropodidae

scytodidae

Unknwn

agele

nidae

anyp

haenidae

araneid

ae

theraphosid

ae

symphyto

gnath

idae

deinopidae

A) Primary Family Dominance Curve

Figure6.TheFamilyDominanceCurvesofeachhabitattype,exhibitingthefamilieswiththegreatestrepresentationinthehabitat.ThemostcommonfamilyfoundintheprimaryhabitatswasPholcidae,whichrepresented26.20%ofthetotalindividualsfound.PisauridaeandTetranathidaemadeupthesecondandthirdlargestcomponentofthesample,representing25.56%and12.14%respectively.Theremaining14familiesandthefewindividualsthatcouldnotbeclassifiedintofamilieseachrepresentedfrom0.32%to8.63%ofthetotalcomposition(figure6A).ThemostcommonfamilyinthesecondaryhabitatwasPholcidaewhichrepresented38.44%ofthetotalindividualsfound.PisauridaeandTheridiidaemadeupthefollowinghighestpercentagesrepresenting33.13%and25%respectively.Therestofthefamiliesandthefewindividualswhocouldnotbeidentifiedrepresentedfrom0.31%to6.56%oftheindividualsinthehabitat(figure6B.)ThemostcommonfamilyinthecultivatedhabitatwasTetranathidaewhichrepresented40.18%ofthetotalindividualsfound.TheridiidaeandAraneidaemadeupthefollowinghighestpercentagesrepresenting21.11%and12.02%respectively.Therestofthefamiliespresentedfrom0.29%to10.26%oftheindividualsinthehabitat(figure6C.)

38.44%

33.13%25.00%

6.56% 4.06% 3.44% 2.1… 1.88% 1.25% 0.94% 0.94% 0.31% 0.31%0.31%0.00%5.00%

10.00%15.00%20.00%25.00%30.00%35.00%40.00%45.00%

Pholcidae

Pisaurid

ae

Therid

iidae

Araneidae

Tetra

nathidae

Thomisid

ae

Ctenidae

Saltic

idae

Scytopidae

Corinnidae

Lycosid

ae

Therophosid

ae

Anyphae

nindae

Unknwn

B) Secondary Family Dominance Curve

40.18%

21.11%12.02% 10.26%

5.57% 4.40% 2.93% 1.76% 0.88% 0.29% 0.29% 0.29%0.00%5.00%

10.00%15.00%20.00%25.00%30.00%35.00%40.00%45.00%

Tetra

nathidae

Therid

iidae

Araneidae

Pisaurid

ae

Saltic

idae

Ctenidae

Lycosid

ae

Corinnidae

Dysderid

ae

Deinopidae

Thomisid

ae

Unknwn

C) Cultivated Family Dominance Curve

ForboththeprimaryandsecondaryhabitatsthePholcidaefamilydominatedthenumberofindividualsintheecosystem.ThisfindingwassimilartothefindingsofMoore(2015),whofoundinuncultivatedregionsthePholcidaefamilywasthemostabundant.Thecultivatedhabitathadthemostsignificantdominanceofasinglegroup(Tetranathidae)comparedtotheotherhabitatswhichshowedalmostequalpresenceofatleasttwofamilies(figure6.).TheabundanceoftheTetranathidaefamilyinthecultivatedareawassoapparenttheywereconsideredapartofthethirdmostabundantfamilyintheregion(appendixc.)

Figure7.TheCaveHabitatDominanceCurve,exhibitingthefamilieswiththegreatestrepresentationinthehabitat

Thepopulationsinthecavehabitatshowadramaticdominanceoftwogroups.ThefamiliesPholcidaeandGnaphosidaepresented96.67%ofthepopulation(figure7).Comparedtothethreemaintransectsthathadincreasedecologicalresources,thecavehabitatscouldonlysupportafewfamilies.Theresourcesandecologicalinfrastructureofcavehabitatsisinhospitabletomanyorganisms,andthepresenceofafewspecializedfamiliesisexpectedundersuchconditions(Mammolaetal2017).GuildCompositionAcrosstheregionthespaceweb,sheetwebandorbweaverguildswerethemostabundant.ThetwomostabundantfamiliesPholcidaeandTheridiidaeareclassifiedasspacewebbuilders,whilethethirdmostabundantfamilyTetranathidaehuntsintheorbweaverguild(table1).

60.00%

36.6667%

1.67% 1.67%0.00%

10.00%20.00%30.00%40.00%50.00%60.00%70.00%

Pholcidae

Gnaphosid

ae

Therid

iidae

Therap

hosidae

Cave Family Dominance Curve

Figure8.HabitatGuildComposition,displayedinpercentagesoutofthetotalnumberofindividualssampled.ChartArepresentsprimaryguilds,ChartBrepresentssecondaryguilds,andChartCrepresentscultivatedguilds.Themostabundantguildintheprimaryhabitatwerespacewebbuildersmakingup35%ofthetotalguildsrepresented.Thesecondgreatestusagewasrepresentedbysheetwebbuilderswhomadeup26%oftheindividualsinthehabitat(figure8A).ThefamiliesbuildingspacewebsincludePholcidaeandTheridiidae(table1),withPholcidaerepresentingthehighestpercentageofindividualsinthehabitat.Themostrepresentedguildinthesecondaryhabitatwerespacewebbuilders,makingup53%ofthetotalindividualsinthehabitat.Thethreemostabundantmorpho-specieswereclassifiedasspacewebbuilders.Thesecondmostrepresentedguildweresheetwebbuilders,makingup28%ofthetotalindividualsinthehabitat(figure8B.)Thefirstandthirdmostabundantfamiliesinthehabitat(PholcidaeandTheridiidae)wereclassifiedinthespacewebguild,whilethesecondmostabundantfamilyPisauridaewasclassifiedinthesheetwebguild(table1).Themostrepresentedguildinthecultivatedhabitatwereorbweavers,makingup53%ofthetotalindividualsinthehabitat.Thesecondmostabundantguildwasthespacewebbuilders,makingup21%ofthetotalindividualsinthehabitat(figure8C).Themost

Ambush Hunter3% Ground Hunter

2%

Space Web53%

Orb Weaver9%

Other Hunters5%

Sensing Web1%

Sheet Web28%

Unknw1%

Secondary Habitat Guilds

Orb weaver53%

Space web21%

Sheet Web10%

Other Hunter10%

Ground Hunter5%

Ambush Hunter1% Unkwn

1%

Cultivated Habitat Guilds

Space web 35%

Orb weaver14%Other Hunter

13%

Ground Hunter7%

Sensing Web1%

Sheet web26%

Ambush Hunter3%

Unknwn1%

Primary Habitat Guilds

abundantfamilyinthehabitatTetranathidaeisclassifiedundertheorbweaverguild,whilethefollowingmostabundantfamiliesTheridiidaeandAraneidaefallintothespacewebandorbweaverguilds,respectively(table1).Only3guildswerefoundinthecavetransects,withthespecialistandspacewebguildsmakingup99%ofguildactivity(figure9).Comparedtotheguildcompositionofthethreemaintransectsthecaveecosystempresentsahighlevelofecologicalstressresultinginonlyafewguildsbeingabletoadapttotheextremesubterraneanenvironment(Mammolaetal2017.)

Figure9.CaveGuildCompositiondisplayedinpercentagesoutofthetotalnumberofindividualsfound.Therichnessandevennessofguildsincreasedfromthecultivatedtotheprimaryhabitat.Habitatswithlessanthropogenicinterventionwereabletosupportahigherrangeofguilds,resultinginhighermeasuresofbiodiversity(table2).Guildswithmorespecializedhuntingtechniqueswerefoundtobemorerepresentedinmoreunalteredhabitattypes(Michalko2016,Cardosoetal2011).Habitatswithhigherecologicalpressure(Mammolaetal2017)orincreasedlevelsofdisruptionwillonlybeabletosupportsimplisticguildcompositions,asthesehabitatslimittheamountofresources,structureandrefugeneededtosupportcomplexguildactivity(Cardosoetal2011).

Space Web72%

Specialist27%

Sensing Web1%

Cave Habitat Guild Composition

VegetationandLandUseImpactThetypeofvegetationineachhabitatcouldbeclassifiedinto9categories.The8guildscouldbearrangedintothetypesofvegetationtheywerefoundusingtohunt.

Figure10.Vegetationandstratapresentinthefourhabitats,withdesignationoftypeandtheguildsfoundusingtheinfrastructure.Vegetationvariabilitywasfoundtodecreaseacrossthefourhabitatssampled,parallelingadecreaseinthebiodiversityandactiveguildsinthehabitat.Boththeprimaryandsecondaryhabitatsshowedthehighestrangeinvegetationtype,whilethecultivatedhabitatshadonly4outofthe9classifications.Thecaveecosystemsshowednovegetationandonlyonestrataclassification(figure10).Thespeciesrichness,familyandguildpresencebetweenthemainthreetransectsandthecavesamplesshowedadramaticdecrease,withonly6morpho-species,4familiesand3guildsbeingfoundthroughoutthecavessurveyed(appendixA.)Astheecologicalstressorsincreased,classifiedindiminishingvegetationvariability,thediversityofthespiderpopulationsdecreasedproportionally(Mammolaetal2017).Theavailabilityanddiversityofvegetationdirectlyimpactedthebiodiversitypresentacrosstheregion.Asvegetationandstratacomplexityincreasedthenumberofactiveguildsincreased.Thiscontributestotheincreaseofbiodiversityasthesupportofmoreguildtypesleadstothepresenceofmorespecies(Cardosoetal2011).Asbothofthesecondaryandcultivatedhabitatshadbeenpreviouslyalteredandbothhadvaryingdegreesofdensevegetationtype,similarspecieswouldfindthemhospitable.Theywerealsoclosertogetherinlocation,meaningmorespecieshadtheabilitytocrossoverfromhabitattohabitat.Spidersinadaptiveguildssuchasspacewebororbweaverswillbeabletothriveinvaryinghabitattypes,asbothrelyonvegetationstructurebutcanadapttovaryingvegetationtobuildwebs(Schüpbach2003).Astheprimaryforestandthecultivatedfieldshadthegreatestdegreeofimpactdifference,itisexpectedtheywouldsharetheleastamountofmorpho-species(table3).

Thesignificantdifferencebetweenanthropogenicimpactofunalteredprimaryforestandcultivatedfieldswouldsupportstheassumptionthatspeciesinprimaryforestwouldnotfindagriculturalareasashospitable(Uetz1991).Thecultivatedtransectshowevershowedarelativelyhighbiodiversitymeasure(table2).AsfoundbyMarc&Canard(1997)agriculturalzoneswithlittletonochemicalimpactwillbehospitabletoarangeofspidersandtheirrespectiveguilds.Theyalsofoundhigherspiderbiodiversityincultivationhasthepotentialtoactasaneffectivepestcontrol,andwiththeintersectionofarangeofguilds,cantargetinsectsinalloftheirlifestages.Thepercentageofbiodiversityfoundinthecultivatedtransectsrepresentacultivatedareawithgoodagroecologicalmanagement.Cultivatedlandthathasbeenabandonedtoacertaindegreewillalsopresenthigherlevelsofbiodiversity,asthelackofchemicaluseandconsistentdisturbancepermitsspiderpopulationstoincrease(Martin1982).ConclusionSpiderbiodiversityaroundtheworld,andspecificallytheneotropics,isstillabiologicalfrontiertobefullyunderstood(Basset2012).Presentinganewfrontierofnewscientificdiscoveries,spidersdisplayanintriguingrangeoftaxonomyandecologicalinteractionsthatrivalscomparativetop-levelpredatorsonothertrophiclevels(Basset2012,Silva1992).Thisstudyfoundspiderbiodiversitydecreasedwithincreasedanthropogenicimpact.Primaryforestswerefoundtosupportmorespiderfamiliesandprovideinfrastructureforincreasedguildactivity.Themostimpactedregions,thecultivatedhabitats,showedadecreaseindiversity,withafewspeciesdominatingtheecosystem.Theclassificationofvegetationtypeshowedthatadecreaseinvariabilitycorrespondedtoadecreaseinbiodiversity.Inboththecultivatedandcaveecosystems(imposingbothartificialandnaturalecologicalpressures)fewerfamiliesandguildscouldbesupported.Thisdemonstrateshowecologicallymindfullandmanagementisvitalinpreservingspiderbiodiversity,asincreasedecologicalpressurelessentheresourcesneededbythemorecomplexguilds.Theuniquebiodiversityfoundintheprimaryforestsshowedhowpreservationofpristineenvironmentsisvital,especiallyinthecaseofspiderswheremanyspecieshaveyettobedescribed(Platnick2013).Thecultivatedhabitatbiodiversitydidshowhowmindfulagriculturalpracticescanmaintaindiversespiderpopulations,simultaneouslycreatingeffectivepestcontrol(Jeanneretetal2003).Spiderbiodiversityacrosshabitatscallsforincreasedscientificexploration,asknowledgeofdiversityacrosstrophiclevelsisnecessarywhendevelopingcompleteconservationprotocols.BibliographyAvezedo,G.H.F,etal.(2014.)Effectivenessofsamplingmethodsandfurthersampling

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AppendixA:Thenumberofmorpho-species,families,guildsandindividualsfoundineachhabitattype.Habitat Morpho-species Families Guilds Individuals

PrimaryForest

73

17

6

313

SecondaryForest

64

13 7 320

CultivatedFields

68 11 6 341

Caves 6 4 3 60

AppendixB:TheFamiliesandguildsfoundineachhabitattype.Habitat Families GuildsPrimary agelenidae AmbushHunter anyphaenidae GroundHunter araneidae Orbweaver corinnidae OtherHunter ctenidae SensingWeb deinopidae Sheetweb heteropodidae Spaceweb lycosidae Unknwn pholcidae pisauridae salticidae scytodidae symphytognathidae tetranathidae theraphosidae theridiidae thomisidae

Unknwn

Secondary Anyphaenindae AmbushHunter Araneidae GroundHunter Corinnidae OrbWeaver Ctenidae OtherHunters Lycosidae SensingWeb Pholcidae SheetWeb Pisauridae SpaceWeb Salticidae Unknwn Scytopidae Tetranathidae Theridiidae Therophosidae Thomisidae Unknwn

Cultivated Araneidae AmbushHunter Corinnidae GroundHunter Ctenidae Orbweaver Deinopidae OtherHunter Dysderidae SheetWeb Lycosidae Spaceweb Pisauridae Unknwn Salticidae Tetranathidae Theridiidae Thomisidae

Unknwn

Caves Gnaphosidae Specialist Pholcidae SpaceWeb Theridiidae SensingWeb Theraphosidae AppendixC:RegionalDominanceCurveandFamilyPercentComposition

AppendixD:BiodiversityComparisonsHabitat Simpson Entropy Comparison Difference % increase

P 3.86 C/S 1.68 61.31% S 2.74 C/P 2.8 72.54% C 1.06 S/P 1.12 29.02% Habitat Gini-Simpson (transformed) Comparison Difference % increase P 25.58 C/S 11.47 67.23% S 17.06 C/P 19.99 78.15% C 5.59 S/P 8.52 33.31%

23.36%

18.24%

17.95%

9.65%7.63%

4.25%

2.51%2.22%

2.12% 1.35%0.97%

0.77%0.68%

0.58%0.58%

0.29%0.29%

0.29%0.19%

0.19%0.10%

0.00%

5.00%

10.00%

15.00%

20.00%

25.00%

Pholcidae

Therid

iidae

Tetra

nathidae

pisaurid

ae

Araneidae

Saltic

idae

Lycosid

ae

Ctenidae

Gnaphosid

ae

Thomisid

ae

Corinnidae

scytodidae

Heteropidae

Therap

hosidae

Unknwn

Agenele

nidae

Anyphae

nidae

Dysderid

ae

Deinopidae

symphyto

gnath

idae

Miturgi

dae

Regional Family Dominance Curve