Historical Ecology of Amazonian Lizards- Implications for Community Ecology

8/12/2019 Historical Ecology of Amazonian Lizards- Implications for Community Ecology

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Nordic Society Oikos

Historical Ecology of Amazonian Lizards: Implications for Community EcologyAuthor(s): Laurie J. Vitt, Peter A. Zani and Maria Cristina EspsitoSource: Oikos, Vol. 87, Fasc. 2 (Nov., 1999), pp. 286-294Published by: Wileyon behalf of Nordic Society Oikos

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OIKOS 87: 286-294.Copenhagen999

Historical cology f Amazonianizards:mplicationsorcommunitycologyLaurie J. Vitt,Peter A. Zani and Maria Cristina sp6sito

Vitt,L. J., Zani, P. A. and Esp6sito,M. C. 1999.Historical cologyof Amazonianlizards: mplications orcommunity cology.- Oikos 87: 286-294.Nineteen peciesof lizardssimultaneouslytudied t the Curud-Una n the centralAmazon of Brazil separateon the basis ofmicrohabitat se and prey ypes.There sno relationship etween microhabitat iche breadths nd dietaryniche breadths,speciesusing hegreatest iversityf microhabitats o notnecessarilyse thegreatestdiversity f prey ypes.A pseudocommunitynalysisrevealed hatthe izardassem-blage is structured ithrespect o microhabitat nd diets. A comparison fdietaryoverlapswithphylogeneticimilaritiesndicates hatmuch of thevariationndietarysimilarityn this assemblage s associated withphylogeneticimilarity. similaranalysiswithmicrohabitat verlapsyielded o relationshipo phylogeneticimilarity.These results uggest hat much of the structure n thisparticular ssemblage shistorical,.e.,not thedirect esult f ongoing pecies nteractionst the ocal level.Because this assemblage s comprisedof lizards from a diversity f higher axa(families),we suggest hat historical actorsmaybe more mportantn determiningstructuren phylogeneticallyeeply rooted assemblagesthan in phylogeneticallyshallow-rootedssemblages.L. J. Vitt,Oklahoma Museum of Natural History nd Dept of Zoology, Univ. ofOklahoma,Norman,OK 73019-0606,USA ([email protected]). P. A. Zani, Dept ofBiology,Univ. f Oregon, ugene,OR 97403-1210,USA. - M. C. Esp6sito,Departa-mentode Zoologia, Museu Paraense EmilioGoeldi/CNPq/MCT,Caixa Postal 399,66017-970Belem,Par6, Brasil.

It has long been championed hat species nteractions(e.g.,competition, redation) re theunderlying echa-nismsresponsible orobserved structuren ecologicalcommunitiesCody 1974, Schoener 1974, 1986, Codyand Diamond 1975). Extensive omparativend experi-mental tudies n a diversityfvertebratesnd inverte-brates have indicated that species interactions ausedivergence n diets, activity imes, and habitat use(Schoener1968, 1974,Losos 1992, 1994,Dunham 1980,Pianka 1986). In some instances, ven withinverte-brates, ivergenceppears to have takenplace relativelyrapidly e.g., Losos et al. 1997). This viewof communityecology stresses resent-day actorswhichhave clearlybeendemonstratedo be importantnmany ases (e.g.,Losos et al. 1997). Recently,however, number ofstudieshave suggested hathistorical ifferencesmongspeciesplaya role n determiningresent-daytructure

of communities e.g., Mayden 1987, Brooks andMcLennan 1993, Cadle and Greene 1993, Farrell andMitter1993). Most such studieshave examined arge-scale data sets to demonstrate hat at least some of thestructure reviously ttributed o present-day peciesinteractions ay have an historical asis (e.g.,Mayden1987).Such studies ed Ricklefs nd Schluter1993: 350)to state, ntheir verview f book chapters ealingwithhistorical cology, ...taxonomic components f com-munityorganization can be comprehended nly byplacing he ocal communityn tshistoricalnd biogeo-graphic context.To some extent, his may apply tofunctional omponents f community rganization swell".This ed us to examine he tructuref a phyloge-netically eeply rooted izard assemblage o determinewhether ivergence istoriesmight ave been mportantdeterminantsf present ay communitytructure.

Accepted 8 February1999CopyrightC OIKOS 1999ISSN 0030-1299Printed n Ireland- all rights eserved286 OIKOS 87:2 (1999)

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Lizards hat at nvertebratesretypicallyhoughtfas generalists,ating rey vailablen themicrohabitatsin which hey ive Rand and Myers 990). t is wellknown hat izard iets iffernd that here re someextreme ood specialistsPianka 1986,Vitt nd Zani1996,Vitt t al. 1997). t has also been uggestedhatmuch of the ecological ariation mong species ntaxonomicallyiverseizard ssemblages ay ephylo-geneticallyr evolutionarilyonservativeThorpetal.1994,Vitt ndZani 1996), utquantitativevidenceslacking.Usingdata on an Amazonianizard ssemblage,eprovide vidence hat izard iets n taxonomicallyi-verse ssemblageshat re structuredith espectoresources ave an historicalasis,thathabitat ichebreadthdoes not necessarily redict ietary ichebreadth,hatmore loselyelatedizardsatmore imilarprey, hat hemost losely elatedpecies o notneces-sarilyive n themost imilarmicrohabitats,nd thatecological ifferencesllowingoexistencensome pe-ciesmayhavebeen nplacebefore hese peciesnter-acted. fthis s the ase, nsitu peciesnteractionsayhavebeen f ess mportancendetermininghe resentday tructurefthis ssemblagehanhistoricalnterac-tions hat ontributedospecies orting.urs s the irststudy o quantitativelyxamine tandard iche xes(time,pace, ood) f ommunitytudies n izardsn nevolutionaryramework.thertudies avefocused nsingle ichexes, uch s microhabitate.g., osos1994).Wefocus nthe womostmportantichexes, ood ndspace microhabitatse).We consider ime elativelyunimportantn thisizard ssemblageecause llexcepttwo pecies rediurnal ndoverlap roadlynactivitypatternss inother mazonianizardssemblagese.g.,Vitt ndZani1998a). ecauseweuse ndividualnimalssampledntheir aturalnvironmentss focal oints ordata collection,urresults eflect resent-dayonse-quencesf he ombinationfhistoricalndpresent-dayinteractionshat ed to ecological ifferencesmongspecies.We lso ommentnthe easons or n historicaleffectnthis ssemblage.inally, e uggesthat ssem-blages fanimalsomprisedfhistoricallyeep-rootedtaxa shouldbe more onservativeith espect o theimpactfongoingpeciesnteractionsn nicheegrega-tion than ssemblages ithmorerecent hylogenetic(divergence)istories.

Methods nd materialsFielddatacollectionFielddatawere ollectedvery ayfrom 3Februaryto 27 April1995bythree esearchersn terra irmeAmazon owland ain orest ear heRioCurua'-UnatAgropecuariareviso TDA, approximately01kmSand 18 km E of Santarem, ard, Brazil 309'2.4"S,

54'50'32.9"W; ereafter,urud-Una). ortions f thearea had beenselectivelyogged8 yearsbefore hestudy, ut theforest ppeared tructurallyimilar oundisturbedorest. he canopywas dominated yBrazil nut trees (Bertholletia excelsa Humb. andBompl) xtendingoapproximately0m off round.Individualizards f 19 species Table 1) were hefocal oints or ata collection. econductedaphaz-ard searches hroughhe forest ach day recordingmicrohabitatssedby izards.Microhabitatategoriesin whichwe found izardswere:Fe-Gr forestdge,grass; e-G forestdge,ground; e-L=forest edge,leaf; Fe-LL forest dge, leaf litter; e-M forestedge,mud;Fe-TBL= forestdge, runk,ranch,imb;Tf-Gr treefall ap, grass; Pf-Gr primary orest,ground; f-L primary orest,eaf; Pf-LL primaryforest,eaf litter; f-O primary orest, ther;Pf-TBL= primary orest, runk, ranch, imb;Sf-Grsecondaryforest,grass; Sf-G secondaryforest,ground;Sf-L secondary orest,eaf; Sf-LL sec-ondary orest,eaflitter; f-MMS litter econdaryforest, an-madetructure;f-TBL secondaryorest,trunk,branch, imb; Sb-TBL streambed, runk,branch, imb; Sb-G streambed, round;Sb-Lstreambed,eaf; Sb-LL streambed,eaf litter; b-M = streambed, mud; Sb-TBL = streambed, trunk,branch, limb; Sb-W streambed,water; Tf-Gtreefall,round; f-LL treefall,eaf itter;f-TBLtreefall,runk, ranch,imb.Our microhabitatatarepresenthe ctual ccurrencef ndividualizardsnavailablemicrohabitats,hey o notrepresentelativedensities f lizards.A totalof 1079 observationsnmicrohabitatseweremade.Four hundredndfiftyndividualsf the 19 lizardspecieswere ampledor ietarynalyses.izardswerekilledwith neumaticiflesrhand apturedndkilledwith lethal ose of sodium entabarbital.tomachswere emovedromizards, reytemseparated,den-tifiedothe owest ossibleaxonomicategoryusuallyfamily),ounted,ndmeasuredor engthnd width.Volumes f ndividualreytemswere stimated iththeformulaor prolatepheroid:Volume= 4/3c(1/2ength)(1/2idth)2For analyses fdiet,we grouped rey tems nto28broader ategoriesepresentingorphotypese.g.,Pi-anka 1986). These are: OD = odonates; GC =grasshoppers nd crickets;BL = roaches; MA=mantises; TE = termites; CO = beetles; HO = ho-mopterans; HE = hemipterans; EL = butterflies ndmoths; SP = springtails; TH = thrips; HY = hy-menopterans non-ant); FO = ants; DI = flies;DE =earwigs; I = miscellaneousnsects;L = insectarvae;AR = spiders; PH = harvesters; AC = mites; IS =isopods; CH = centipedes;DI = millipedes;MA = mis-cellaneous arthropods;EW= earthworms;MO =

OIKOS 87:2 (1999)



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mollusks; VE = vertebrates;EL = lizard eggs; SS =lizard hed kin. here s no evidence rom ast tudiesthat lizards discriminatemong families r specieswithin he bove categories.urthermore,nalyses ftropicalizard iets sing xpanded ersus educed ietdata have produced imilar esults t the communitylevel e.g., Vitt nd Zani 1996, 1998a,b). The onlydetectableifferences that ietaryverlapsre slightlylowerwith reater umbersf morenarrowlyividedprey ategoriese.g., amilyevel dentifications).ore-over,fpatternsre detectable ith he bovedietarycategories,urtherivision f prey ategories ouldresultngreateretectableifferences.onsequently,fanythingur analyses re conservative.

Community analysesAs a measurefproportionalse ofprey nd micro-habitat ypes or izard species,we calculated ichebreadthswith the reciprocal f Simpson's 1949)measure:

nB= l/Z pi/ iwhere is theresourceategory,is theproportionfresource ategory used bythat pecies,nd n is thetotalnumberfresourceategoriesseePianka1986).

Values ary rom ne exclusivese of single esourcestate) on (evenuse of all resourcetates).We usedSpearman ankcorrelationnalysis o determinefthere as a relationshipetween ietaryndmicrohab-itatnichebreadthsi.e., do lizardswith hebroadestdiets also occur in the greatest diversity fmicrohabitats?).To test the hypothesishat izard speciesuse re-sources andomly ith espect o what s usedby theentire ssemblage,we conducted seudocommunityanalyses n volumetricietary ata andmicrohabitatuse data. The pseudocommunitynalysis asbeende-scribednd testedndetail lsewhereWinemillerndPianka1990).Nevertheless,e brieflyeviewt here.Amatrixfproportionaltilizationaluespi)was calcu-lated romhe ommunityatrixlizard pecies s rowsversus rey ypes r microhabitatypes s columns) ydividing esource se of each category y thecolumn(resource) otal for each species row). A matrix felectivityaluesei)was calculatedJacobs 974)fromthesamedata and scaledto varyfrom to 1 withcolumnsummingo1.0 Winemillernd Pianka 990).Thegeometric eansgi) ofpi andeiwere alculatedresultingn a third onsumer-resourceatrix. hisprocedurealances pposite iasesresultingrom seofproportionaltilizationsr electivitiesncalculationsofoverlap Winemillernd Pianka1990).Thegidatawere hen ubstitutedorpi data in the formula orecological verlap ielding:

Table 1. Lizard species occurringt the Curud-Unasite,microhabitats,nd niche breadths ormicrohabitatsBin) and diets(Bd).Species Microhabitats B. BdNorops uscoauratus um&ril Arboreal;trunks, imbs,branches 1.55 6.12and BibronNorops trachydermaope Arboreal-terrestrial,ow vegetation,eaf itter 6.90 5.35Noropsortondi ope Arboreal;treetrunks nd limbs 3.19 5.16Dactyloa punctataDaudin Arboreal;treetrunks, imbs, nd canopy 2.85 2.48Enyalius eechii Boulenger) Arboreal-terrestrial,ow vegetation,eaf itter 1.28 2.70Tropiduruslica (Linneaus) Arboreal; argetreetrunks nd limbs 1.00 2.86Tropidurus mbra Linneaus) Arboreal;moderate-sized reetrunks nd limbs 2.15 1.09Uranoscodonuperciliosus Arboreal;smalldiameter reetrunks, imbs, nd vines 1.13 7.06(Linneaus)Gonatodes umeralis Arboreal;treetrunks 2.59 11.15

(Guichenot)Coleodactylusmazonicus Terrestrial;eaf itter 1.90 6.81(Andersson)Hemidactylusmabouia Scansorial;flat urfaces, suallyman-made tructures 1.00 4.48(Moreau de Jonnes)Thecadactylus apicauda Arboreal;treetrunks nd limbs 3.39 3.13(Houttuyn)Ameiva meiva Linneaus) Terrestrial; round, eaf itter 1.67 6.39Cnemidophorusemniscatus Terrestrial; round,grassy pen areas 5.26 3.37(Linneaus)KentropyxalcarataSpix Terrestrial-arboreal;round, eaf itter, ead branches n treefalls,ow 8.55 3.59vegetationLeposomapercarinatum Terrestrial;eaf itter 1.28 5.94(Muller)Cercosaura cellataWagler Terrestrial;eaf itter 2.50 2.35NeusticuruscpleopusCope Terrestrial-aquatic;treambanks 1.48 4.65Mabuya nigropunctataSpix) Arboreal-terrestrial;ree trunks nd limbs,palm fronds,eaf and palm litter 3.82 6.16

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Z gig g1ki i=/n n

where ymbolsre the ame s abovebutwith andkrepresentingizard pecies.Matricesfoverlap aluesfor ll species omparisonsere alculatedormicro-habitat nd dietaryata.To examine hetherhe ver-lapvalues ased nrealdatadifferedrom hatwouldbeexpectedased n a randomamplingf he ata,weperformedwopseudocommunitybootstrap)nalyses.Inthe irstnalysis,scramblederos", ll valuesntheoriginal imatrixwererandomized 0000times ndoverlapswere alculated or achrandomization.orthisnalysis,he umberf esourcesndniche readthsremainedntact. he econdnalysis,conservederos",was denticalxcept hat ddressesnthe imatrix ithzeroswere ot andomized ith espectoposition.henumberfresources,iche readths,ndzero tructureof thematrix eremaintained.esultingverlaps ereranked, ith rank f 1 indicatingearest eighbourand higher anksrepresentingorrespondinglyoredistanteighboursnnichepace.This llows tatisticalcomparisonsfoverlaps esultingrom he seudocom-munitynalyseso the eal verlapst all ofthenearestneighbouranks. tatisticalignificanceas based onthepercentfpseudocommunityverlapsallingelowthe eal ommunityt each ankWinemillerndPianka1990).

PhylogeneticnalysesTo test hehypothesishat heresanhistoricalompo-nent o resource seof izards,wefirstonstructedmatrixfphylogeneticimilarityalues omputedsingthe"Distance" rogramf theCOMPARE computerpackage E. P. Martins) rom hefollowingreeforwhich ehad dequate iet ata: (((((Noropsfuscoaura-tus: 1,N. trachyderma:): 0,N. ortonii: ): 1,Dactyloapunctata: ): 1, Enyalius eechii: ): 1, (Tropiduruslica:1, Tropidurusmbra: ): 1,Uranoscodonuperciliosus:):2): 2, (((Hemidactylusmabouia: 1, Thecadactylus api-cauda: 1): 1, (Gonatodes humeralis:1, Coleodactylusamazonicus: ): 1): 3, (((Ameiva meiva:1, Cnemidopho-rus emniscatus: ): 1,Kentropyxalcarata: 2): 1, (Lep-osoma percarinatum:1, Cercosaura ocellata: 1): 1,Neusticuruscpleopus: ): 1): 1, Mabuyanigropunctata:4): 1) 1). This ree epresentscompositeromeveralphylogeneticnalyses n portions fthetree. amilylevel elationshipsrebased n cladisticharacternal-ysisof Esteset al. (1988)with heexception f theIguaniaconsideredocontain hreeamiliesyEstes tal.). Familial elationshipsithinguania rebasedona cladistic nalyses y Frostand Etheridge1989).Relationshipsf lizards n thefamiliesolychrotidae

andTropiduridaere basedonconsensusreesGuyerand Savage1992 ndFrost 992, espectively)hereasrelationshipsithin heGymnophthalmidaere basedon a rootedWagner etworkPresch 980).Teiidrela-tionshipsrebased n Presch1974). or a summaryfrelationshipsorother amiliesepresented,ee Estesand Pregill1988).Branch engthsre indicated ynumbersndnestingf thephylogenys indicatedyparentheses. e thencompared hecorrelationsfdietaryndmicrohabitatverlaps or pecies airs ophylogeneticimilarityaluesusingMantel's andom-ization estMantel 967,Manly 991).n this est, llof theelementsf one matrix rerandomizedeforecomputingorrelationsetweenmatrices.henumberof randomized orrelationoefficientsqual to orgreaterhan he bservedorrelationormshe asis orthe tatisticalest asedon 10000 andomizations.

ResultsMicrohabitatiche readthsBin) vary rom .0 e.g.,Coleodactylus mazonicus o as highas 8.6 (e.g., Ken-tropyxalcarata)Table1).Overlapsnmicrohabitatsevary rom .000 o as high s 0.961, utmost re ow(Table 2). The use of microhabitatsy izards ftheCurud-Unas not random nd structurexistswithrespect omicrohabitatse Fig. IA). DietsofcentralAmazonianizards ary mong pecies n prey ypes(completeistingfspecies' iets an be obtained romthesenior uthor). ietary ichebreadthsBd) varyfrom .09 nthe ntspecialist . umbrao as high s11.15nG.humeralisTable1).Overlapsnprey sevaryfrom .002 o as high s 0.871, utmost re ow Table3). The izard ssemblages structuredith espect ouseofprey ypes Fig. lB). Species o notrandomlyselect ood rmicrohabitats,ather,hey isproportion-ately ivide subset f allresourcesvailable. here sno relationshipetweenmicrohabitatichebreadthsand dietarynichebreadthsr,= - 0.07, P = 0.77, n=19); izards hat se a diversityfmicrohabitatsonotnecessarilysea diversityf prey ypes.A significantositive orrelationxists etween i-etary verlaps ndphylogeneticimilarityP = 0.028);themore loselyelated wo pecies re, hemoreikelythey reto eat similar rey.However,here s not asignificantelationshipetweenmicrohabitatverlapsand phylogeneticimilarityP = 0.088); morecloselyrelatedpecies o notnecessarilyive n similarmicro-habitats.hus, iets f izardsnthis ssemblageeflectwhatancestors f these izards te (Fig. 2) whereasmicrohabitatssedbythese izards o notnecessarilyreflect icrohabitatssed byancestors f each izardspecies. significantositiveorrelationxists etweendietaryverlapsndmicrohabitatverlapsP = 0.018);microhabitatse does, to someextent, eflectreychoice.

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DiscussionNumerous studies of lizards (Schoener 1968, Pianka1973, 1975, 1986,Pianka et al. 1979),birds Cody 1974,1983), mammals Brown 1973, 1975), and fish Wine-miller1989, Winemiller nd Pianka 1990)have shownthat species within ssemblagestypically eparateonone or more niche axes. In New Worldtropical izardassemblages, he niche axis "time" appears unimpor-tantbecause nearly ll speciesare diurnal nd overlapbroadly in time of activity e.g., Vitt and Carvalho1995, Vittand Zani 1996, 1998a, b). Evidence n thisstudy and others show that lizard species generallyseparate on the basis of food or microhabitat.Whenlizard species eparateon thebasis oftime, t s usuallybecause a given ssemblage ontains largenumber fnocturnalgekkonid izards e.g., Pianka 1986). More-over,at leastone study uggests hat timemaynot bean important iche axis in lizard assemblages Hueyand Pianka 1983).In this tudy,wehaveshown hat pecies eparate nthe basis offood and place (microhabitat). t least twononexclusive ypotheses an account forthis apparent"partitioning"f resources: ) specieshavediverged sthe resultof species nteractions t a local level overlimitedresources nd 2) species coexist because theywere differentn resourceuse patterns o start with.The firstmplies resent-day rocesses uch as competi-tion in determining tructure e.g., Schoener 1968,Cody 1974,Connell 1975, 1980). The second invokesthe role of historical vents e.g., sortingof speciesoccurred ased on differencesn resource se patterns)in determiningresent-daytructuree.g., Brooks andMcLennan 1993, Cadle and Greene 1993). We haveshownthat there s a strong ssociationbetween hedietsofAmazonian izards nd their volutionaryimi-larities.Rather than present-daynteractions ausingobserveddietarydivergencemong thesespecies, t isapparent that present-day ood preferences ave anhistorical asis. The lack ofassociationbetween hylo-geneticsimilaritynd microhabitat imilarity mongthese species and the significantelationship etweendietary nd microhabitatimilarityuggest hatmicro-habitatchoicemay be more labile than food choice:lizardsmayselectmicrohabitats ased on the kinds ofprey vailable nthem ather hanfeeding n a randomset of prey vailable in specificmicrohabitats.Microhabitat ransitions ithin lades are more fre-quent than food typetransitionsFig. 2); phylogenydoes notappearto constrainmicrohabitat se as muchas itconstrains rey hoice. Similar esultswere ppar-ent in an Ecuadorian Amazon lizardassemblagecon-taining ome of the same species Vittand Zani 1996)but quantitative omparisonswere not made. Recentstudies emonstratingreydiscriminationn scleroglos-san lizards based on chemical cues (Cooper 1995a)providesa mechanism orprey choice in the lineage

290 OIKOS 87:2 (1999)



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containing ore hanhalf fextantizard pecies.naddition,eneral ietaryifferencesetweenguanians,which at primarilyymenopteransnd beetles,ndscleroglossans,hich at primarilynsect arvaeandspiders,redeeply ootedn thephylogenynd at eastpartiallyied o foraging ode Huey ndPianka1981)which s also deeply ooted n the volutionaryree flizards Cooper 1994, 1995b).Evolutionarily,ietarychoicemay e more eritablehanmicrohabitathoice.Lizards hould e expectedo switchmicrohabitatsofindpreferredood; they houldnot be expected oremainnmicrohabitatsith ow food ualitydefinedas prey atenby ancestors hatresultedn increasedrelative itness) henpreferredood is available nnearby atches nless he ostto switchmicrohabitatsis high Charnov 976).

0.8 -A.0.7 - Microhabitatommunitynalysis0.6 -0. 0.5 -- 04 -'B 0.3 -o 0.2 -0.1-100 -80 C

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1 2 3 4 5 6 7 8 9 1011 12 13 1415 16 1718Rank of nearestneighbournniche paceB.

0.6 - Dietary ommunitynalysis0.5 -0.5 - X real verlapsCU0.4 -A scrambledero verlapseL --- conserved erooverlapso 0.2-0.1 -0.060 - scrambled ero50 - -A- conserved eroCL 40 -02 30 -20 -10-0 A... AAAAAA.1 2 3 4 5 6 7 8 9 10 11 1213 1415 16 1718Rank ofnearestneighbournniche pace

Fig. 1. A) Results of pseudocommunitynalysis on Amazo-nian lizardmicrohabitats howing hat structure xists n theassemblagewithrespect o microhabitat se. Scrambled erooverlaps re significantlyower thanreal overlaps t all rankswhereas onserved ero overlaps re lower only at rank 2. B)Pseudocommunitynalysis n Amazonian izarddiets howingresults imilar o thosefor microhabitats.

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Historical Ecology of Amazonian Lizards- Implications for Community Ecology