Occurring Island Reptiles

Page 1of50 between this versionandtheVersionofrecord. Pleasecitethis articleasdoi:10.1111/geb.12716. through thecopyediting, typesetting, paginationandproofreadingprocess, whichmayleadtodifferences This istheauthormanuscript acceptedforpublicationandhasundergone full peerreviewbuthasnotbeen authors all Itescu of behalf Yuval on Sincerely, Biogeography Ecology and Global in for and at level publication ready now is our manuscript find you will hope We comment. each with of we how dealt by description point a find point Below please accordingly. manuscript the of these and revised most followed We suggestions. important their for and referee the editor the and thanks review supportive were see We to happy the comments. added her own Santos, Dr. Ana Editor, the and referee anonymous a by was new reviewed manuscript suggestion). The (Ref. GEB20170100; title modified according referee’s to modified title reptiles’ (Ref. GEB20170100; island co-occurring of our manuscript version attach aWe revised McGill, Dear Prof. Biogeography and EcologyGlobal Chief in Editor McGill, Professor Brian 07.12.2017

Global EcologyandBiogeography . . ‘Inconsistent patterns of body size evolution in in size evolution body of patterns ‘Inconsistent

suggestion and it would strengthen your results, but I will leave it up to you to decide if youyou decide to if I to up your it but leave will results, strengthen would andsuggestion it a good is I this think selection. based model you AIC that use and new reviewer suggests This your submission. previous greatly since has improved Iyour it and think paper, share view of his I of reviews. round first the in involved not was a that by referee evaluated was Your manuscript your manuscript. happened as with it to, like would we takes that longer it process.editorial Sometimes for the delay the in I apologize First, want to Author: the to Comments Ana Santos, Editor: TO AUTHORS COMMENTS EDITOR'S last paragraph, as part of your takehome message. your message. of takehome as paragraph, part last the you in theory former that rejecting you are stress to In need little. particular, although very discussion, the in also rewording needed is of this theory”. Some former rejects/contradicts “this that also reptiles.” And Aegean on evolution size body driversuniversal of are no “There like saying something sentence a straightforward need start to conclusions abstract the main the in Similarly, “Divergent”. than rather coherent” or “Non “Heterogeneous” I where title, use the heterogeneity such e.g. in put to front, in text the more, tweaking a bit it you I highlight should why think is That timeliness. and its of paper the importance the with line I’m in 100% so yougreat job, a correlations make on each based species separately explain to trying rather than species among differ responses that By highlighting rules”. “island of general lack the front and about make a in paperyou this put choseI’m to glad really so result, important most the is your species all predictors over of power predictive general lack the me To of ones. and a few small comments, got significant I’ve two more data. species with the analyses on additional the eliminate good to was decision a it me to so straightforward, an robust seems version this but round, first get the I review a to nice is work. didn’t This Author the to Comments 2 Referee: . Author the to Comments 1 Referee: TO AUTHORS COMMENTS REVIEWER Editor Dr. Ana Santos, islands. on evolution generality of theory size the contrast our results that discussion of the paragraph concluding the and in abstract the in and highlighted appendix scores a new in AICc included we below, As we specify comments. supporting for editor the her thank We universal. not rule is island the that generalyour conclusions more ofsuggestion highlighting his follow to you is do I to advise What strongly manuscript. the these analyses in include should

Page 2of50 Page 3of50 predictor across all species? Here the total weight of a predictor for a species it would be the sum be sum the would it a species for Hereof weight a predictor total the species? all predictor across file. supplementary a new in for species each (AICc 2) ≤ models best paper in cite this now We modest. too (2) is parameters additional adding to applies AIC penalty I which concur, results, perhaps raw the not although supplement, the in AICcprovide results after but readingit, I with certainly live could a biostatistician I’m not unconvincing. Since bit

low sample size, as some of yours are. Here, the pick would be to compare models according to models compare be to would yours pick the Here, are. of as some size, sample low with for datasets suited particularly are AICc And scores model. the in of variables amount the way, penalizes for it you correct the in used AIC I whether wonder DinizFilho responseyour to a use AIC to not scores out, stand reasoning I the found will conclusions the so general drivers, lack the of on insights sound provides scores based R2 on of models your selection you that in I agree with would is Although selection. model remaining text the in problem main the me To it. to according changed text We the suggestion. forThanks this results from a different approach. approach. from a different results similar includes it if be more trustworthy yourwill But work informative. is often it how about you just tells it of relationships, the signs homogeneity the a on pick is provide certainly not does approach this What paradigms. analytical are across results that consistent show to useful I quite find So still AICs”. used they if had general patterns be would there they Surely use R2s! “but readers saying I other but hear also can metaanalysis”, the with that showed we“but have youII saying can hear think of species. the regardless informative be a to general is predictor, it closer the 16, to closer the across species, all And is. a variable more the informative 1, closer to for the are AICw, models tested all Given that present. where is models it all of of AICw the per AIC weights all summing why just not so tested, are parameters weights same the if are AIC but datasets, based different on among comparable models not AIC are scores weights? based AIC on similar something why doing your not So metaanalysis. is here, approach sound is frequentist a why reason actually real the your and results, compelling of most the me To the AICc of scores present and now get themselves, our impression readers the let to suggestion referee’s the accept we Nevertheless, ours. as such for studies selection model criteria based over information selection based model prefer for pvalue to good why example a serve as our results think as we point regarding this added also a of paragraph discussion We text. the and second, the want it), you not do still diseases of infectious best the (like models informative group a of non of best the AIC may lowest the be simply with why: model the first, explains compellingly and clearly http://onlinelibrary.wiley.com/doi/10.1111/13652664.13060/abstract) anygood?” model ‘best’ the criteria, is but usingselection information “Model 2017, Ecology, of Applied (Journal paper et al.’s recent Nally and Mac informative, most the is model which scores determine AICc) to (or AICon even relying comfortable with not are weof revision), round previous the in referees reply the to (and in text the weAs in stated be confusing. would quickly, and so approach the information abandon youI shouldn’t think species. Anyway, for several model best as the competing probability say,you’llwith, or 30% 20 havesure models I’m informative. most the is model a that probability the particular is, that Akaiketheir weight;

Now added. added. Now as a valuable contribution for GEB in my opinion, and these other analyses can be published can be published analyses and these other my in for opinion, GEB contribution as a valuable out and stands and straightforward, clear is work The current sound. is few species aanalyses on more the detailed out of leaving decision I the that say. really to think I little Besides have that weights analysis. AIC the include we to 5, prefer not Appendix in AICc based models the show do although we predictor. Therefore, of effect the of the direction the information no about provides it mentions, rightly as referee the especially since, confusing, somewhat is metaanalysis the in shown what we show have to using weights AIC that think but suggestion this for referee the thank We comment). (see first my result your important most highlight to similar, or “Strikingly” with paragraph I’d this L366. start text. the from sentence the we omitted interspecific differences, rather but the right), own its in studyquestion an interesting study perhaps (but focus of this are the not evolution size body differences in and since intersexual the readers, byconfusion our any avoid want to we or only Since males. females only we if test even maintained species, is different in size affect body factorsdifferent that conclusion, general our but studythe species, some in of exist do single to predictors response and their females in males differences between misleading: a bit be may it phrased as we sentence originally the that realized now We incorrect) grammatically is (it sentence this and rephrase material), (as supplementary results these show L242243. Please book 1967 inclusive rather more the than and Wilson, of MacArthur paper for 1963 the citation the just and retain (1957) for Darlington a citation we so added mechanism, (often contentious) any to point than rather islands, nature only of depauperate the emphasize want to actuallyWe ibt. just ETIB than rather mention rephrase and just theories, need cite No these latter to anagenesis. than be more important to starts wherezone cladogenesis radiation determine a area and increasing or that isolation with, familiar you one be may mention richness, to species increases also island the heterogeneity within habitat as,such e.g., that topics, many other includes (ibt) theory biogeography island (ETIB). Currently Biogeography Island Theory of Equilibrium you & Wilson’s MacArthur refer to here certain, Line74. be To Sentence rephrased. reptiles…” all data for the analysing informative, are more predictors which determine to and metaanalyses linear models use “We analyse” for I’d “We substitute In 52 line you use. of analyses kind which you tell don’t In Abstract. methods the follow: comments minor Some elsewhere.

Author Manuscript

Page 4of50 Page 5of50 continuously to the caption to understand the figure. figure. the understand to caption the to continuously have jump to not reader the does so Y axes, variable the in names put Figures and Please 2 3. Done. for sentences. or separate 2 3 out canIprobably make read. to and difficult fragmentary sentence, is it this L483486. Rephrase prediction. Heaney’s failed supporting that first the we not are as wording, other strong such use “strikingly” or to appropriate not is it specifically here that we think but suggestion, this for referee the thank We The names of the variables on the Y axes are now clearly marked in figures 2 & 3. & 2 3. figures clearly in marked now are Y the axes on The of names variables the

Research paper Research Ann Arbor, MI 48109, USA48109, MI Arbor, Ann 4NS,UK E1 London snakes. Key words: 7 Greece 84, 6 5 4 USA 3 02138,MAUSA Cambridge, 2 1 Pafilis Panayiotis Valakos D. Sagonas Kostas ([email protected]), Slavenko Alex Donihue Colin ([email protected]), Itescu Yuval reptiles. island co-occurring in size evolution body of patterns Inconsistent School of Biological and Chemical Sciences, Queen Mary University of London,of University Mary Queen ChemicalSciences, andof Biological School USA TX 79409, Lubbock, Tech TexasSciences,University, Biological of Department 76203,Denton, TX ofTexas,North University Sciences, Biological of Department Israel6997801, TelAviv University, TelofAviv Zoology, School Department of Biology, National and Kapodistrian University of Athens, Athens 157- Athens of Athens, University andKapodistrian Biology, ofNational Department School of Natural Resources and Environment, University of Michigan, Ann Arbor, Arbor, Ann Michigan,of Universityand Environment, Resources of Natural School Department of Organismic and Evolutionary Biology, Harvard University, HarvardBiology, andofEvolutionary Organismic Department

Author Manuscript

6 ([email protected]), Johannes Foufopoulos Johannes ([email protected]),

1 body size, competition, island area, isolation, lizards, predation, predation, lizards, isolation, area, island competition, size, body

([email protected]), Rachel SchwarzRachel ([email protected]), 1 ([email protected]), Stephanos A. Roussos A.Stephanos ([email protected]), 6 This article isprotected by copyright. All rights reserved. ([email protected]), ShaiMeiri ([email protected]),

Global EcologyandBiogeography

5 ([email protected]), Efstratios Efstratios ([email protected]), 2 ([email protected]) 1 ([email protected]) 1 1 7 ([email protected]), ([email protected]), 3,4

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Author Manuscript 73 of references: Number maintext:in the of words Number 300abstract: inthe of words Number [email protected] 6409403; +972 +972640phone:9811, fax: Israel; Aviv6997801, Tel Author: Corresponding title running Short This article isprotected by copyright. All rights reserved. : Body size evolution in insular reptiles reptiles insular in evolution size Body

Yuval Itescu, Department of Zoology, Tel AvivUniversity, Zoology, of Department Itescu,Yuval Global EcologyandBiogeography

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predictors to directly assess the effects of island area, isolation (both spatial and isolation islandofarea, effects assess tothedirectly predictors species. single separately,foreach and snakes lizards and for reptiles, all for informative arewhichpredictors determine to metaanalyses and models linear used We bodysize evolution. on competition andpredation interspecific temporal), largeof aarray quantifiedalso We 273 islands. species across eightsnake and representingeightlizard individuals 10,000 nearlydata on surveycollect to literature Methods studiedtaxa Major period Time Location conditions.insular similarly to respond species cooccurring whether and patterns are, the common whatevolution, bodysize primarily drive characteristics island which tested andthesame archipelago in cooccurring species reptile 16populationsof insular multiple across bodyvariation studied size We evolution. bodysize promote these conditions thatand islands, remote onsmall, relaxed pressures are predation interspecificand competition that predicts Theory evolution. ofbodysizedrivers patterns andboth poorly understand stillwe research much Aim ABSTRACT Animal body sizes are often remarkably variable across islands, but despitebut islands,variable across often remarkably sizes are body Animal Author Manuscript

We combined field work, museum measurements, and a comprehensive comprehensive aand measurements, work, museum field We combined Aegean Sea islands. SeaAegean

3 54 53 52 51 50 49 48 47 46 45 44 43 42 41 40 39 38 37 36 35 34 33 32 31 Page 8of50 Page 9of50 particularities interact to drive the course of sizeofevolution. interact the course to drive particularities eachpredictor snakes.While andlizardsbetween andwithindifferfamilies patterns reptile bodysize evolution on islands. Instead, species’ autecology and island andspecies’ autecology Instead, islands. on evolution bodysize reptile ofprocesses and patterns oversimplify generalizations thatoverarching conclude onislands. We trajectories bodysizeevolutionary general of theories contradicts This conclusions Main isolation. with decreases richness,and predator and competitor area, with increases generally size snake we whereas tested, predictors the byofany hardly areaffected lizards group, aAsrecovered. trend was nogeneral leastspecies,one at sizein body influenced Results

Author Manuscript Body size varies with different predictors across the species we studied, andwe speciesstudied, the differentacross predictors varies with sizeBody This article isprotected by copyright. All rights reserved. No factor emerges as a predominant driver of Aegean reptile sizes. reptileof driverAegean predominant aemerges as No factor Global EcologyandBiogeography

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become smaller and more isolated (e.g., Darlington, 1957; MacArthur & Wilson, &MacArthur Wilson, 1957;Darlington, (e.g.,moreisolated andsmaller become oftenperceivedas islandsis on Trait evolution contradicted.and proposed been havedrivers patterns and and multiple of attention, has received decades populations body sizes. Additionally, he hypothesized that food limitation is the most important important most isthe limitation thatfood he hypothesized Additionally, sizes. body differentattrends different produces but all sizes, important at isequally predation

Authoronbody factors theseecological selection indirectof suggesting islands, on variation bodysize for explanations Alternative themainland. onislandslargeand competition interspecific and islands, on mediumsized predation islands,on small agent selection oftheeffect that Heones.hypothesized to small also than animals to large important more isones limitation thatto food and large animals smallthan to important more iscompetition interspecificthat He hypothesized animal. and theislandfocal of thesize with changes theseoffactors of each thatthe effect suggested (1978) Heaney 2005).1982;Lomolino, Melton, 1978; (Case, onislands sizeevolution drive body to Manuscript thought processes commonly are these ecological resourcelimitation, with Together 1982). (Melton, competition intraspecific stronger consequently and densities population promotehigher in turn, which, pressures, predation and competition interspecificrelaxed to experience thought animals are insular Therefore, 1963). islands as speciespoor becoming depauperate, generally are faunas 2015Insular & Winkler, 2014;ItescuLeisler Meiri, al.,2007; et 2016 cf. Svenning, & Faubry Lomolino, 1973;2005; size (Van body Valen, a totendmedium evolve animals rule”, insular suggesting “island isthe pattern debated The 2008).most et al., Köhler 2005; Lomolino, 1973; Valen, (Vantaxa across consistent and predictable strong, variation insular among bodysize of evolution in pathwaysthe general Identifying INTRODUCTION This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography

5 90 89 88 87 86 85 84 83 82 81 80 79 78 77 76 75 74 73 72 71 70 69 68 67 Page 10of50 Page 11of50 becoming smaller on both smaller and larger islands (Heaney, 1978). However, MeiriHowever,1978). (Heaney, larger islands andsmaller on both smaller becoming large mammals thatwhileof decreases mammals small of size thebody predicted smalleron increase size small in that suggested species (2005)Lomolino size. body be completed (Anderson & Handley, 2002). However, accelerated trait evolution on traitevolution accelerated However, 2002). Handley, & (Anderson completed be

Authorto optimum anchangetowards unidirectional aallow toyetpassed not has isolation since time wheresufficient systems bodysize in with associated be to thought is Temporalisolation differ.wouldnot islands farand on sizesclose body 1999), & Ives, SeaAegeanreptiles,Foufopoulos (asofthe case in unlikely is dispersal overwater where that, suggested (2002) Handley &Anderson 1979). Arnold, 1978; (Heaney, pressures and competitionpredation as such factors reflecting indirect, possiblysize ison body isolation spatial of The2000).effect (Heaney, likely more makes rates gene (i.e., and flow) immigration reduces al.,2010), et Handley,Meik &2002, (e.g., Anderson nearest mainland thefromdistance island as Manuscript isolation,usuallycalculated Spatialtime. inspacein and defined be both can Isolation island area. to sizeof linearresponse or a for prediction, Heaney's nofor support (2005) al. found et islands, largestareintermediatesized sized on mediumanimals that predicted further He1998). Taper, Marquet & 1982;also (seeMelton, withareaincreasing increases (1978) Heaney (Meiri,2007). lizards (Boback, and 2003), snakes 2006), 2005, al., (Meirietmammals in relationships such for support found however,minor studies, Other islands. on such becomesmaller species largewhile islands isolated more and to animal affect thought characteristics island thearetwomain isolationand Area Palkovacs,2003). 1994; Levins, (Adler& proposed have beentraits,alsohistory life on directselection via size, This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography in situ in adaptations adaptations

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population densities, and consequently stronger intraspecific competition and strongercompetition intraspecific densities,consequently and population aggressiveness (Pafilis et al., 2009; Donihue et al., 2016), which in turn favors large large in favorsturn which 2016), et al., Donihue et2009; (Pafilis al., aggressiveness Authorhigher promote thought to iscompetitors) also predators interspecific and from (both releaseEcological 1978). etal., seeDunham but 2003; Hasegawa, Case,1978; 1970; Schoener, 1966; (Soulé, sizechanges promotes nicheand shifts allows competition interspecific Relaxedal.,2007). et 1978;Vervust (Heaney, adaptations antipredatory onselectionsizerelated byrelaxingdirect specieslarge in decrease size and species small in increase drive issize thoughtpredationto Release from2016). et al., Donihue 2010; Meik Meiri,et al., 2007; 2003; Boback, 1989; &Moriguchi, Hasegawa 1966; (cf.Soulé, results providedinconsistent unclear.studies Previousis evolution size reptilebody affectisolation and area island How2008). 2007,Meiri, 2003; Guyer, Manuscript&1978;Boback e.g., Case, (see characteristic cladespecific abe to seems mainland theto on islandscompared diminish or tosizesgrow tendbody reptile Whether (Caribbean snakes and geckos ( lizards smallesttheworld's oneand on hand, dragons Komodo giantand tortoises insular forms: extremesized their for wellknown are Reptiles species.small stronger in effectshowing a possibly with the later competition, interspecific predation toand apply should area for suggestedisland pattern the therefore and interspecific competition andpredation reflectsarea island suggested, Heaneyas Furthermore, isolation). of highdegrees lowand larger at and isolation, at intermediate smallest arespecies intermediatesized (i.e., that well as isolation possiblyfor is true area prediction island for (1978) Levins,1994),& Heaney's (Adler area island decreasing as changessame phenotypic theto drive isexpected isolation As2015).increased (Aubret, suggested been also islandshas isolated recently

Brookesia This article isprotected by copyright. All rights reserved. Tetracheilostoma chameleons; Hedges & Thomas, 2001; Glaw et al., 2012) and2012) etal., Glaw Thomas,2001; & Hedges chameleons; Global EcologyandBiogeography threadsnakes; Hedges, 2008) on theother. 2008) onHedges, threadsnakes; Sphaerodactylus

7 139 138 137 136 135 134 133 132 131 130 129 128 127 126 125 124 123 122 121 120 119 118 117 116 115 Page 12of50 Page 13of50 potentially enables us to distinguish between patterns driven by the island conditions conditions bytheisland driven betweenpatterns tous enables distinguish potentially

2016), making this system ideal to study size evolution on islands. We aimed to test to aimed Weevolution onislands. size study to ideal thissystem making 2016), Author etal., Itescu size, documented maximalreaches the even some species in (and greatly alsovaries thesepopulationsislands on in reptile Body size composition. faunal isolation inand area, widelyThese islandsvary theSea. Aegean in 273islands from snake species), (eight eight lizardandspecies individuals16reptile of10,000 nearly fordata bodysize encompassing coverage databaseunprecedented ofa assembled We speciesspecific. those thatstudiedare and we design Thisstudy (Meiri al.,2008). species et cooccurring such among uniform relatively butremain etc., primaryvegetation,productivity, climate, latitude, as Manuscriptsuchregions different across vary that factors islandspecific eliminatetoone allows samearchipelago the within cooccurringspecies Comparing archipelago. single a within reptile species multiple ofpopulations multiple island agents across selection potentialof multiple quantifyingthe effect questionsdirectly by these approached we Therefore,2011). Pringle, &Losos 2011;2010, Cox, & (cf.Calsbeek species single fordebated even a sometimes influentialmostis is factor Which conditions. insular to respondsimilarly species cooccurringwhether and patterns are, common the whatevolution,body size driveprimarily characteristics island which unclear remains It1978).al.,(Dunham genuset or 2010) etal., Meik1966; Soulé, (e.g., species single of a populations across ofpredictors some the effects or 2014), examined et al., Itescu 2007; 2003; Meiri, Guyer,& Boback 1978; Case, (e.g., populations or species island and mainland studieseither contrasted evolution reptilebodysize date, insular To 2003). Palkovacs, 1982; (Melton, isadvantageous whereearlysizematurity is expected smaller although sizes, This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography

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Mediodactylus kotschyi Mediodactylus

Koenig in Bonn, Natural History Museum of Crete, Goulandris Natural HistoryNatural Goulandris Crete,of Museum Natural HistoryBonn, in Koenig AuthorAlexander Forschungsmuseum Zoologische München, Staatssammlung (Zoologische collections museum in eight specimens measured further We (19842016). years 33 periodicallyover spring summer duringandthe field specimens in measured We collection Data 2008). etal., (Valakos islets verysmall even inhabiting thegeckowithSea islands,inhabitAegean species reptile Fifty islands. varyacross greatly resource availability composition and faunal Consequently, 2005). al., et(Fielding areas maquis agricultural and sclerophyllous evergreen ‘phrygana’), Manuscript called (locally scrub Mediterranean patchworkdwarfof aare landscapes the and 2010) Poulakakis, (Lymberakis & diverse are geologicalhistories Their area. in ofsix magnitude orders varyingacross several islands thousand has Sea Aegean The system Study METHODS 2008).etKöhler al., 2005; (Lomolino, taxa across consistent are islands on patterns 4. size Body 1978); (Heaney, sizedependent ispredation of not effect the while species large thanstronglymore species small competition affects Interspecific (Heaney,3. 1978); and competition predation of isolation, degrees intermediate and islands largestintermediatesized specieson are Medium 2.sized 2005); Lomolino, 1978; (Heaney, thefocal species for predators competitors and fewer harbor space),and time and(inisolation in increase in area, decrease as islands smaller species become largewhile size in species increase Small 1.hypotheses: several This article isprotected by copyright. All rights reserved. and thelacertid and Global EcologyandBiogeography Podarcis erhardii being most common, common, mostbeing

9 186 185 184 183 182 181 180 179 178 177 176 175 174 173 172 171 170 169 168 167 166 165 164 Page 14of50 Page 15of50 p=0.23; p=0.23;

of isolation for islands isolated during the past 20,000 years (since the end of thelast theend (since pastof years 20,000 during the isolated islandsisolation for of AuthorPeriods tools.Earth Google usingwere calculated Distances sinceisolation. time the index, isolationtemporal island alargerand the closest fromthedistance studied also we Ives,Therefore 1999).& (Foufopoulos archipelago AegeaninSea the especially (Itescu,2017), isolation effective quantify notadequately maythis index systems island landbridge isolation.for spatial However, ofindex as mainlandan nearest the testedfrom thedistance Meirial.,2005),we 2005;et Lomolino, 2003; Boback, included; e.g., bodysize of (studies biogeographyislandstudies most Following analyzes. further inall field data literatureand museum,pooled therefore We Manuscripterhardii ( sourcesbetween differencesdata found we no withsufficient species two the For source.data each and females for males five of five at least bodysize data recorded wewhich islands from only compared islands. We several forin thefield measured specimens museum collections and in measured specimens bodysizeof mean the we compared nottheresults,bias did data sources of multiple our use ensure To the literature. fromand thein recorded field datawe wasfrom calculated species each bodymassfor Mean snakes. for length total and (SVL)lizards, for length snoutvent reportedsizeindices: commonly themost used 2). We Appendix 1, islands(Table 273 species originating from snake lizard of individualseightand eight 9951adult for size bodysex recordedand We 1 seefora list). sources;Appendix 97 extracted from were (data possible as reptiles islandAegean as many forbodysize information recorded and surveyedliterature we Finally, comprehensively History). ofNatural Museum Yale Peabody and University, HarvardZoologyat of Comparative Museum History, Natural of Museum National French Museum, HistoryNaturalBritish Museum, : field mean SVL=61.05 mm, museum=60.43 mm, n=38 islands, t=1.22, n=38islands, mm, museum=60.43 mm, meanSVL=61.05 : field M.kotschyi This article isprotected by copyright. All rights reserved. : field=43.35 mm, museum=43.68 mm, n=25, t=1.05, p=0.30). p=0.30). t=1.05, n=25, mm, museum=43.68 mm, : field=43.35 Global EcologyandBiogeography P. 10

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population. Predators were defined as all mammals and reptiles likely to prey upon prey to reptiles mammalslikely and as defined all were Predators population. Service Navyof Hydrographic the maps Hellenic bathymetric withregional wasconnected itwhich to landmass thelast andaisland focal between potentially compete with adults of smaller ofspecies. withadults compete potentially ofpredatory counts valuesthecombinedand richness birdpredatory between thecorrelation tested we richnessvalues predatorour affect significantlydid not birds

Author species oflargethat assumingjuveniles snakes),snakes (forother or lizards) (for otherlizards defined as rare.were tooCompetitors themare listfor faunal specific and byisland reptilespredationarthropods, on either on reports since centipedes) scorpions,spiders,(e.g., arthropods predatorypotential quantify not could We our fromdatabase. counts bird to confidentexclude feel therefore (p<0.01). We 0.90 (r)was coefficient correlation 2017).The (Itescu et al., lists birdpredatory reliable reasonablywe have did islandsforwhich 41 across reptiles predatoryand mammal ManuscriptthatTo excluding ensure lists. islandspecific reliable to itcreate impossible making sites, breeding their hunt awaythemtofrom well allows migration seasonal their and islands across mobility since their excluded wereBirds (Appendix species4). focal of speciesfocal each competitor and predator potential the count to observations our own field and theliteraturebased onlists faunal islandspecific We assembled 3). (Appendixliterature the from assembled wereLGM the than isolatedearlier islands of isolation Temporal finesuchresolution.a at isolation time estimate accurately can method presumeour donot and acknowledge wevalues didalthough round not we resolution, yearand one to werea calculated Estimations (www.hnhs.gr/geoindex/). resolution fine were depthsdrawn from Maximum 2011). et al., Foufopoulos 1999; Ives,&Foufopoulos (seeLGM the changesince sealevelof charts specific depth themaximum fordata by crossing calculated –were maximum"LGM") glacial This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 11

236 235 234 233 232 231 230 229 228 227 226 225 224 223 222 221 220 219 218 217 216 215 214 213 212 Page 16of50 Page 17of50 positive relationship where Heaney's prediction holds, since it asserts small species sincespecies itsmallholds, asserts Heaney's where prediction relationship positive

transformed body mass. Here we expected to find a significant negative trend for for negative trend tosignificant a wefind bodymass. expected Here transformed Author against log coefficients correlationof the the value absolute regressedwe classes, size all important isacross predation equallywhile species largethan for species important small for more iscompetition thatinterspecific prediction Heaney'stest To 2005). et Meirial., (seepositiveshowslopes species and zero, large to equal slopes have would species slopes,mediumsized bodysizepredictor show negative would tofind a expected ofspecies. bodymassWe each logtransformed against variables, sixpredictorof andthe bodylength) each body (i.e. betweensize relationship ofthe coefficient the correlation regressed we bodysize thespecies by affected Manuscriptvariablesis predictor of ofthe species and each sizeevolution between relationship thethat prediction (1978) Heaney's testTo curvilinear. islinearor richness) competitor andpredator richness, isolation,time since island, larger closest the fromdistance the distancemainland, area, (islandfrom islandcharacteristics six the ofbodysize ofwith each therelationship whether species each for explored first We difficult. isextremely manner invasive non the in ina sexfield their determiningreliably ofsince sex regardless individuals means. Onlyfor femaleand male bysize mean averaging bodypopulation calculated we size dimorphism, sexualto due sizebiases avoidTopredictor variables.in the variation maximizehence, and islands ofthe rangeaswell as populations of species numberand us maximize theto allowed This approach islands. at least eight sampledacross population per individuals at least ofthree hadwe measurements which reptile species for 16 examined We Analyses This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography Ablepharus kitaibelii Ablepharus kitaibelii weof amean all used 12

260 259 258 257 256 255 254 253 252 251 250 249 248 247 246 245 244 243 242 241 240 239 238 237

package (Laliberté, 2011). Finally, we examined for each species which of the sixof the which species examined weforeach Finally, 2011).(Laliberté, package it iscomparable across since these analyses in predictorvariable the mass as body being the best of a collection of poor models; Arnold, 2010; Mac Nally et al., 2017). MacetNallyal., 2010;Arnold, poorofmodels; collection aof thebest being levelswere significance when pvalues>0.05 (i.e. hadsize with associated poorly

that our general conclusions are robust for using different modelselection approaches. approaches. modelselection differentusing robustfor are ourconclusions general that Author5toAppendixhighlight in species for models each bestAICcbased present the we Nevertheless, conservative. moreproved pvalue approach maligned themuch Thus, merelyscores thelowest(models with them noninformative making estimated), werewhich predictors had oftenscores with the lowest models because the scores) orAICc (AIC selection model criteria for information theAkaike using avoided We quadratic terms. linearand both using(α<0.05), on pvalues based procedure selection model backwardstepwise bya followed species, eachfor regressiontest multiple Manuscripta performedwespecies. thisendTo models bestacross selected the compared and size onislands body predicts its that best themodel comprise characteristics island Rthe 'metacor' wereusing performed Themetaanalyses we analyzed. valuesr the standardizeto all species, predictors) for tested (against body size log10transformed of regressions from thecorrelationcoefficients used separately. group We each for sizes, effect as all species in variable predictor eachsize ofagainst body regressions linear of the (r)of coefficients correlation metaanalysis randomeffect Laird DerSimonian We conducted separately. lizards)(snakes andsuborder each across generalorin studied we reptiles all bodysize across significantly affects characteristic islandwhetherany to explore approach metaanalytic took thena We 2016). al.,etFeldmanMeiri, 2013; (Feldman& length body than comparisons specific inter for suitableismore andtherefore bodyshape, driven bybiases eliminating taxa, usedholds. We predictionpredationif the no trend for andcompetition interspecific This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 13

285 284 283 282 281 280 279 278 277 276 275 274 273 272 271 270 269 268 267 266 265 264 263 262 261 Page 18of50 Page 19of50 predictorbody size relationships were nonsignificant, and no predictor was importantwas predictor no nonsignificant, and wererelationships size predictorbody butmost species, leastone of bodysizeat with wasthe wecorrelated tested predictors theanalyzedvalues. standardize to species all for predictors) Following model selection, selection, model Following 6).statisticsAppendix in model univariatesee (Fig. 1;modelsfull univariate inwastested area when size between and area, twonegative) positive and (three linearrelationship asignificant showed species only five and significant, area island snake (the reptilespecies 16 ofoutone2).In only (Table within families separately,evenand lizards and snakes within 16 species examined, the across greatlyvaried predictorof each sizes effect modestus and snakes Onlyspecies.the the16studied signs) across trend and variablesincluded ofterms (in models 15 different found 1). We (Table all species for ofEachspecies. the highlyacross inconsistent bodysizewere for models best The RESULTS tested(against bodysize log10transformed ofregressions usedvaluesfrom ther we variabledependent the(r)as coefficients we correlation analyzed where allcases In all analyses. normalityintest) ShapiroWilk transformation; normalbefore not were 16species ofsix of the distributions(residual size bodyneeded, where and isolation, timesinceislandarea, log10transformed weparametric oftests assumptions the meet Tosamemodel. in 5)the ≥ factor (variance inflation others with colinear highly werewhich predictors discarded We predictor. tested per points islands) (i.e., data threeof amaximum include to modelsby limiting overparameterization avoided We

Authorammodytes Vipera Manuscript and and

Natrix natrix Natrix This article isprotected by copyright. All rights reserved. shared a similar model. For two snake species ( species snake twoFor model. sharedsimilar a ) no predictors were significant. Explanatory powerand Explanatory significant. were )predictors no Global EcologyandBiogeography T.fallax Telescopus fallax Telescopus still showed the same quadratic pattern, pattern, and quadratic thesame stillshowed ) was a quadratic model of bodyofmodelfor quadratic size was )a Elaphe quatuorlineata Elaphe Eirenis Eirenis 14

309 308 307 306 305 304 303 302 301 300 299 298 297 296 295 294 293 292 291 290 289 288 287 286

predictor, negatively correlated with body size in three species and positively so in in so positively species threeinand bodysize with correlated negatively predictor, significantfrequently most thewas distance the mainland snakes, from For predictor. p=0.23. R p=0.23.

DISCUSSION DISCUSSION Authorpattern).thelizard of opposite the (i.e., island larger theclosest was thedistance from size body snake affect significantly did not that variable only predictor The sinceisolation. with time and mainland the fromthedistancedeclined with and richness, predator and competitor with well as island as area, withincreased significantly however,bodysize, Snake that). weakatone (and a lizards on (positive) effect generalhave a to seemed island larger theclosest fromdistanceoverall. the reptiles bodyOnly in on size effect significant a wehad of tested thepredictors none revealedthat4) (Table metaanalyses The ManuscriptR p=0.40. (slope=0.06±0.07, predation bothof the importance showed that (n=16) bodymass against regressions richness lengthcompetitor body and richness lengthpredator bodyfrom coefficients valuesof the correlation the absolute 2). Regressing 3, (Table Fig. nonsignificant were inthisanalysis richness competitor andrichnesspredator area,island for results The since isolation). time largerandtheisland closest from distance mainland, the (distance from indices threeisolation thefornegative relationship asignificant found we mass,body its log against species study each in variable eachpredictor against bodyregressionof size (r)fromthe coefficient thecorrelation regressedwe When one. significant most wasthefrequently island area lizards for fact, In in four). negative species and(positivetwo onlyin in lizards emerged relationships linear significant

2 =0.10) for body size variation is not sizedependent (Fig.3). not issizedependent variation bodysize for =0.10) This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 2 =0.05) and interspecific competition (slope=0.08±0.06, (slope=0.08±0.06, competition interspecific =0.05) and 15

333 332 331 330 329 328 327 326 325 324 323 322 321 320 319 318 317 316 315 314 313 312 311 310 Page 20of50 Page 21of50 patterns emerged, except that most predictors were uninformative for most species (a (a species most for wereuninformative predictorsmost that except emerged, patterns between body size and island area, Boback’s (2003) metaanalysis revealed no revealed metaanalysis (2003) area, Boback’s island bodysize and between

snakes in general support the findings of Meik et al. (2010, 2012), but for most most for but2012),(2010, ofet al. Meik the supportfindings ingeneral snakes Author( rattlesnakes inspeckled correlation positive a2012)strong(2010, found et al. Meik them,between and correlation negative correlation found (1989)a Moriguchi & snakes, Hasegawa For inconsistent. often are resultstested, directly arereptile bodysize on and ofisolation area effects the when However,2005).2004;Lomolino, Ziv, & (Melton,1982; Filin intensify inhabitants their traitsofon the environment theinsularof effects the isolated, moreandgetsmaller islandsthatas biogeographyis inisland perception common A increases Manuscriptisolationtemporal geographicasand smaller become totend species large while islands,isolated more largeron become totend species reptile said,small That snakes. in and lizards those in conditions to environmental on size ofislands body response thein strikingdifferences revealedalso results Our reptile body onsize. insular effect nogenerallikely have richness competitor and richness, predator area,Island species). across inconsistentare (i.e., patterns direction particular a in reptiles studied the ofbodysize drives insular significantly indices isolation ofnonethethree that showed sizesof effect the thismetaanalysis withofInfinding, sorts).line consistency fewconsistent ofVery factors. different combination bya were affected species allnearly showedthat species across best ofmodels thea andcomparison species, of the majority even affected variablesconsistently of thepredictor None significance. and (linear, curvilinear) shape sign,varyingin variables predictor the of effects with studied,we thespecies diversityamong pattern great found studied. We we factors theto respond sizes theirway in idiosyncrasy the greatshow reptiles island Aegean This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography Crotalus mitchellii Crotalus ). Our metaanalysis results ).Ourfor metaanalysis 16

358 357 356 355 354 353 352 351 350 349 348 347 346 345 344 343 342 341 340 339 338 337 336 335 334

patterns in the predicted way. Moreover, isolation, regardless of the index tested,theindex of isolation,Moreover,regardless predicted way. in the patterns surprising somewhat iseightspecies) of (for sevenout lizards ofmodels best proved false. Predator and competitor richness did not affect body size of most of ourofmostof size notbodyaffectdid richness competitor and Predator false. proved biased.

the biotic interactions directly would result in stronger patterns. This, however, however, This, stronger patterns. in wouldresult directly interactions biotic the Authorof the testing effect thatwe2005), expected (Lomolino, size withbody associations significant found andbiotic effects for proxies as isolation area islandand used haveothersAs interactions. biotic of weak effects only wefound few, Surprisingly, size isnot island on variation forsizepredation of thattheimportance prediction his butspecies,largesupports strongly than species smallmore influences competition thatinterspecific prediction(1978) refuteHeaney's results also Our archipelago. theSea Aegean leastin on islands, at reptilebodyofvariation patterns size on overallhas noeffect 2005), Lomolino, 1978; theory(Heaney, to in contrast however, Manuscript area, Island 2005). Lomolino, 1973;(VanValen, size evolution body reptile driving in ofisolation islandrole highlight aresults These patterns. theopposite shows bodysize reptile drives examined we theisland ofcharacteristics that none clear Itis of inreptiles. case the predictions (1978) Heaney's ofmost support cannot We trend.nogeneral to species points the relationship across changes of direction thethatthe fact However,2016). see butet al., 2007,Donihue Meiri, 2004; al., 1978; etet al.,LososDunham 1966; (Soulè, studies previousof results considering theinofisland area inclusion The frequent relationship. negativewe a found whereas isolation or geographic temporal size either and betweenno association found (2010) al. and et (2003) Meik sizeBoback ishumpshaped). body with relationship butthe is, (for to model selection bodyofaccording size predictor important particularly ais notarea island species)oftheouteight (seven studiedwe species This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography Telescopus fallax Telescopus it 17

383 382 381 380 379 378 377 376 375 374 373 372 371 370 369 368 367 366 365 364 363 362 361 360 359 Page 22of50 Page 23of50 predation pressure index in the literature (e.g., PérezMellado et al., 1997; Cooper et et 1997; Cooper et PérezMelladoal., literature(e.g., the in pressureindex predation body size evolution on islands are intraspecific competition and resourcelimitation and competition intraspecific onislandsare evolution size body

Author2008).2007,Meiri, 2003; 2003; Boback, Hasegawa, Moriguchi, &1989; Hasegawa 1987; Shine, (Schwaner,1985; snakes forwell, asespecially onislands sizes body withincreased reptiles associated in usually are preyavailability and diversity, preyof inpreysize, terms resources, Richer 1993). &Schwaner, see Case but 2013, etal.,1978; Keehn (Case,islands sizeson withlarger toassociated be thought also is ofstudy someourspecies, characterizes and conspecifics resources against defending involveswhich Territoriality, 2009).al.,(Pafilis et pressureislow predation where competition, intraspecific ofstronger large under conditions asize to evolveneed the byhas explainedbeen on islands gigantism example,For 1982). 1978; Melton, (Case, Manuscript to shape oftenthought this arenot intest but didstudy thatwe importantfactors Two islands. thanpredatorpoor, ratherpredatorfree, on only apparent are changes evolutionary issignificant that possibility Another 2014). (Meirietal., intensity competition ofindex richness abe weak may Similarly,competitor 2017). et al., Itescu 2005; (Meiri et al., intensitypredation poorlyrichnessreflects predator 2004), al., use aas suspectits common that despite we Therefore, competitors\predators. twodominant orone pressurethan selection astrongerimpose necessarily donot predators and competitors possibly inefficient, many, thatmaybe implies richness predatorof and thecompetitor weak effect think We isgreater. richness competitor andpredator largersizeswhere tendencytowards showeda metaanalysis agroup),the generalas (i.e., inreptiles or for notlizards for but snakes, For model). bestintheir included factors hadthesespecies of (less the thanquarter a species study This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 18

407 406 405 404 403 402 401 400 399 398 397 396 395 394 393 392 391 390 389 388 387 386 385 384

predators are rare, foraging and basking times may increase, thereby allowing allowing thereby mayincrease, times basking foragingand rare, are predators founder effects. orresponse) genetic adaptive (in to contrast responses phenotypic populations is that where adult mortality rates are low (e.g., where predation is low) wherepredation are rates(e.g., mortality low where adult isthat populations se per islands (Kolbe et al., 2012). Thus body size variation is not necessarily orsolely necessarily not variation is body size Thus2012). et(Kolbeal., islands youngandremote small,in patterns, especially bodysize in shaping role have a also may effects Founder &2008). HasegawaMori, 1989;(King,lives their throughout growreptiles longerandsurvive individuals because attained are adultsizes larger variation across for size body nonadaptive explanation alternative Another underselection. truetrait gape size with the being spandrel, an evolutionary than more no inis variationsnakes size that body proposed (2009) al. et Vincent 2012). (Aubret, bodysize variation populationlevel drive may pressure) also predation sizebiased ratheronthanbodysize growth rates on directselection Ofgrowth.course, enhanced where Additionally, 1993).Shine, Madsen 1991; & 1976; Forsman, (Case, differences size body nonadaptive mayproduceresource availability in variation from resulting populations insular variability growth rateacross plastic that indications Thereare growth).faster permits food more (e.g., changesor plastic adaptation) (i.e., genetic changes of varyresult as a ratescan growth individual example, For patterns. reptilebodysize on have agents selection suggested commonly effect minor thesystem, ourconsideringinstudy relevant seem especially possibilities These plastichabitatdrivenas such possibilities out alternative rule cannot one adaptations, underof a framework iscommonly onvariationexamined islands bodysize While 2009). etal.,(Pafilissizes largerselectingbody for consequently reptiles, insular drive in cannibalism may resource limitation Furthermore,

431 430 429 428 427 426 425 424 423 422 421 420 419 418 417 416 415 414 413 412 411 410 409 408 Page 24of50 Page 25of50 patterns found for snakes follow the common predictions regarding insular evolution evolution insular regarding predictions thecommon snakes follow found for patterns island. larger theclosest distancefrom from apart factorsmost respondsto size body

eggs), or that the variation of one group is adaptive, while that of the other is led by theotheris led of that groupwhile isadaptive, oneof thatthevariation or eggs), Authortheir lizards and uponpossibly prey snakes,but quality food offer for which islands, of onratsremote theabsence (e.g., differently snakes lizards and that affect factors ofor combination reflects factor ofanother isolation theeffect thateither speculate we However, other.the thanstrongly moregroupone indices affect isolation different reasons discern the pointwe thiscannot At islands.remote on more larger grow to species) smaller (i.e., lizards tendencyofgenerally by driven probablythe is largerisland theclosest thedistancefrom trendfor the negative In contrast, increase. factorsas dwarfism these largertowards species) (i.e., generally of snakes tendency bythe driven andare isolation time since themainland distanceof from the cases thetrendin thenegative Likely, complex. somewhat bodyis size onreptile isolation Manuscriptof thattheeffectoverall suggest theseparately, groupsof two each found for we part. Thethemostpatterns 1978), in species (Heaney, predictedsmallfor patterns the notlizards do follow of however,those Interestingly, 1978). (Heaney, specieslarge in Generally, sincethe withisolation. andtime themainland fromdistance with decline richness,predator to and richnessand competitor withincreaseisland area, totends It snake In contrast, closestisland. thewithfromlarger declinetodistance tendency weakfor except a island toconditions, general responseno shows bodysize Lizard each other. topicture opposite anexactlyshow they we found Instudied. fact, we respond to the factors sizes howtheir in differ level, markedly thesuborder at snakes, is lizards resultsand that fromour emerged that pattern important and interesting An near patterns future. the in nonadaptive and between adaptive discrimination 2017)better may Raia,allow & (DinizFilho approaches noveland adaptive, This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 20

456 455 454 453 452 451 450 449 448 447 446 445 444 443 442 441 440 439 438 437 436 435 434 433 432

preys upon its prey species wherever they coexist, and that we can correctly identify we identify correctlycan that coexist, theyand species wherever prey upon its preys species predator thata(e.g., someassumptions general richness)necessitates predator not allow did species thestudied available for currently phylogenies populationlevel body sizes because for some islands we only had data from few individuals. Weindividuals. few onlyfromdata wehad islands some for because sizes body levendis Podarcis strauchi, Blanus schweizeri, Macrovipera

Authormean populationlevel assessing of accurately theexpensecome at may however, This, values). in predictor variance the maximizing also species (thus each for islands thenumberof maximize to aimedwe therefore population and the was analysis ofmain unit island.Our per sampled individuals islandsand sampledof numbers the between tradeoff exists inherentthe rest,an For beproperly analyzed. to islands speciesSeveral(e.g., species. certainfor some populations and infor samples short quitewe came still reptiles examined 10,000 adult samplesizes.Withalmost small is to face, challenging mostdrawback, and theimportant mostPerhaps results. our biasednot havethey we islands, areconfident species and regarding all assumptions Manuscriptthesekept we Sinceconsistently predators). andimportant competitors all richness, (e.g., competitor ourofpredictors of some Thenature patterns. evolutionary of investigation generallyfurther facilitate will which studies, molecular level island importance of weTherefore,the highlight thisaspect. robustlyexamine to us the However, nonphylogenetic models. and phylogenetic bycomparing addressed beto needs onvariationislands size body shaping in condition sharedancestral of noted.beThe role shouldseveral andcaveats methodologicalchallenges and logistic alwayswillonecreate thiswideas scope ofa such study a comparative Conducting 2007lizards). for Meiri, andsnakes for 2003 Guyer, & Boback rule,island(e.g.,cf. the beenbefore haveshown islands onevolution body size snake oflizard Inconsistentpatterns and isnonadaptive. signalsor founder effect strong This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography ) simply occur on too occur simply few ) on 21

481 480 479 478 477 476 475 474 473 472 471 470 469 468 467 466 465 464 463 462 461 460 459 458 457 Page 26of50 Page 27of50 because of any theoretical views about its merit, but because we prefer to err on on the errprefertobecause webutmerit, its viewsabout theoretical of any because 5). (Appendix We uninformative as rejected method based thatthepvalue predictors pOften, <(at0.05). predictors onlysignificant with thethanbest model predictors

side of caution. side Authorvaluesnot p use thus We islands, important. actuallyare on sizeevolution animal with relationship tenuous mosthave the that thatvariables, many madehaveinfer us would liberal more thatiseven methodselection model aUsing2017). et al., Benjamin (e.g.,2013; Johnson, enough isnot 0.05conservative p= anything, if that, think –including predictors AICcmore thelowestcontained with models the however, fewer lowestcontain AICc the with models, groupof undistinguishable or model, the didno 5).In Appendix case in thoseTable2to in models one (cf.based pvalue the Manuscript than lessconservative far simplyprovesmethod theAICcIn cases many cases). these in bad(i.e. equally goodequally thatare predictors thatmultiple arethere show only ofsize body in variation the weexplains tested the of none predictors that inferusing pvalues we example, For inference. biological problematic forbe can alonescores onAIC relying thus and pooroverall often are AICc) models AIC(orprefer. toThebest approach selection ofmodel which issue thestatistical for example a providecompelling results Our ourresults. from conclusions valid robustlydraw to usthisenables think reptiles.Wefor leastscope, at unprecedented bothof speciesare of these sampledwithin each populationswe insular of number the weand species numberofexamined the Moreover, valid.are conclusions ourconfidence have wespecies,thus the bestsampled few even across apparent speciesare found we across inconsistencies patternsandgeneral the Yet, cases. some inresults our havemay affected powerstatistical that low possibility the acknowledge This article isprotected by copyright. All rights reserved. Natrix natrix Natrix Global EcologyandBiogeography and Eirenis modestus Eirenis . Using AICc, we could could AICc, we Using . 22

506 505 504 503 502 501 500 499 498 497 496 495 494 493 492 491 490 489 488 487 486 485 484 483 482

predictors of animal trait evolution, and into the need to quantify relevant ecological ecological quantifyneedrelevant tothe into of and trait animalevolution, predictors isprobablyisland bodyon size evolution that thus conclude We reached. being study inthis least at island byconditions, identity than speciesby more affected be

collections they manage. We thank the editor Dr. Ana Santos, the reviewer Dr. J.A.F.Dr.reviewer Santos,the editorAna Dr. thethank manage.they We collections intheworkingwhile Crete), MuseumHistory (Natural Papadimitrakis Manolis AuthorLymberakisandPetros Koenig, Bonn), Alexander Forschungsmuseum (Zoologische Böhme and Butt, UrsulaWolfgang Dennis Rödder, München), Staatssammlung (Zoologische Glaw Franzen,and Frank Michael of the assistance appreciate deeply We work.help with field all their for Werheimer KerenOr and Vezyrakis AlexandrosTamar, KarinTallowin, Sviri, Oliver Sharoni, Achiad Eylon Savion, GayaMosses, MichaelErez Lewin,Maza, Amir Levinkind, Shani Kurzits, Menachem LamaKhoury, SimonJamison, Hartmann, Anat Goshen, Golan,Eden Efrat Gainsbury, AlisonFreud, NirTamar Einav, Brown, Stav AvivBejerano, thank We ACKNOWLEDGMENTS Manuscript systems. study differentfor effects monotonous as island characteristics from paradigm, away evolutiontheinsular of majorrethinking afor theneed elucidates study This size evolution. of processes and patterns the complex oversimplify generalizations and islandspecific, and species generalizations from did,prevent tounjustified weas simultaneously, mechanisms drivingpotential severaltestingof the importance highlightsspecies, across greatly differedmodelsthe best that and thespecies,of majority aaffected consistently tested wethe predictors of that none Thefact taxa. different across general are islands bodyofon size patterns thatevolutionary theoryasserting the This contradicts system. to seemsvariation bodysize reptileThus, environment. insulartothe idiosyncratically respond speciesdifferentlarge aextentthat to indicate thisof study results The This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 23

531 530 529 528 527 526 525 524 523 522 521 520 519 518 517 516 515 514 513 512 511 510 509 508 507 Page 28of50 Page 29of50 processes driving the evolution of morphological and life history trait variation ofvariation traitlifehistory and of morphological evolution drivingthe processes REFERENCES REFERENCES Authorfauna.insularon of theecology environment insulartheof implications bythe fascinated alsoHe is animalsspecifically.and general insularin animals theand patterns theinbiogeographic interested candidate isaPhD Itescu Yuval BIOSKETCH 2.in inAppendix isavailable thisstudy analyzedthedata All ACCESSIBILITY DATA Manuscript6. Appendix 5. Appendix 4. Appendix 3. Appendix 2. Appendix 1. Appendix INFORMATION SUPPORTING declare. interests to ofhave noconflict The authors #1005/12. (ISF) grant Foundation Science Israel byan study funded Thewas (Permit 20305/824). Change Climate andEnergy, Environment, of Greek Ministrythe from withpermission conducted was work All of thispaper. improveearlier versions to contributedgreatly that reviews, constructive and providinginsightful for tworeviewers andanonymous DinizFilho,

A list of potential predator species included in our analyses.ourincluded in species predatorApotentiallist of LGM. before islandsisolated time of isolation Literaturefor sources Full statistics of singlepredictor linear and curvilinear models. curvilinear models. linear singlepredictor and Fullof statistics scores. AICc species basedon each for Best models usedanalyses.Data for data. size body listliterature containing forReference This article isprotected by copyright. All rights reserved.

Global EcologyandBiogeography

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Naturalist Machery, E., Maxwell, S.E., McCarthy, M., Moore, D., Morgan, S.L., Munafó, M., Munafó, S.L., Morgan, D., M., Moore, McCarthy, S.E., Maxwell, E., Machery, A., R., Little,Lupia, J.,D.,List, Laibson, M.,Kirchler,Jones,M., Jeon,J.H., J.P.A., G.,Ioannidis, Imbens, K.,Imai, D.J., H., Hruschka, Hoijtink,T.H., Ho, L., Held, L.V.,Hedges, J.D., Hadfield, A., Greenwald, D.P.,Green,E., Green, S., Goodman, R., E.I.,Gonzalez, George, M.,Forster,A.P.,Field, Fidler,F., E.,Fehr, C., Efferson, K., A.,Easwaran, Dreber, Z., Dienes, P.,Boeck,De T.D., Cook,Clyde,M., C.D., Chambers, D.,Cesarini,C., Camerer,L., Brown,B., K.A.,Brembs, Bollen, R., Berk, E.J., B.A.,Wagenmakers, Nosek,M., J.O.,Johannesson, D.J.,Berger, Benjamin, snakes. neonate insularsize ofinrates body evolutionary theandIsland F.(2015) colonisation Aubret, size? onselectionbirth of consequence anonadaptive snakes tigerin variations size on islands: are adult evolutionBodysize F.(2012) Aubret, Criterion. Information using Akaike's modelselection andparameters Uninformative (2010) T.W. Arnold, 286 adaptations. islandand systematicstortoises: giant (1979)their IndianOcean E.N. Arnold, rate. evolutionaryand selection, sloths:biogeography, in Dwarfisminsular (2002) Handley,R.P.&C.O. Anderson, Biology of Review Quarterly syndromerodentpopulations. in Theisland(1994) R.Levins, & G.H. Adler, Author Manuscript127145. , , 179

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Biological Sciences Biological in evolution size body Ecology andBiogeography Ecology Gardens, Kew, London, Surrey, UK. London, Surrey, Kew, Gardens, (2005) Mathew, B. &N.J. Turland, J., Fielding, thetuatara. and snakes,lizards,amphisbaenians ratesof diversification and sizesBody S.(2016) Meiri,I. R.A., Mayrose, & N., Pyron, A.,Sabath, Feldman, Society theLinnean of snakes. in allometry Meiri,Length–mass S.(2013) & A. Feldman, 187 EvolutionaryPattern. aObscured Have Basic Extinctions HumanDriven Rule: theIsland ofResurrection Svenning,(2016) J.C.& S. Faurby, Tale. Cautionary A Island in SizeLizards:Body J.W.(1978) D.W.Gibbons, & Tinkle, A.E., Dunham, lizard. island of an ecology onthefunctional intraspecific aggression and ofavailability impact food the testing or Feedfight: (2016) A. Herrel, J. & Foufopoulos, Brock,K.M., C.M., Donihue, brain– andgenetics quantitativeRule, (2017)Island P.Raia, J.A.F. & DinizFilho, York.New Sons, & Wiley (1957)P.J. Darlington,

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614 613 612 611 610 609 608 607 606 605 604 603 602 601 600 599 598 597 596 595 Page 32of50 Page 33of50 Research AuthorIzuIslands.on the thesnakes of historytraitslife and size habits,body in food Geographicvariation H.(1989) Moriguchi, & M. Hasegawa, Herpetology of study. longterm field a from gigantic? Evidence be to longer orlive snakegrowfaster insular gigantic Doesa (2008) Mori,A.& M. Hasegawa, IzuIslands. the of snakes on thelizard and the studies of areviewreptiles: insularterrestrial of diversification Ecological (2003) M. Hasegawa, Manuscript( chameleons leafof species new microendemic andminiaturized discoveryreptiles: of smallest world'sthe Rivaling (2012) Vences,T.M.& Townsend,J.,M. F., Köhler, Glaw, 119129. islands. of Mediterranean ratesreptiles from extinction elevated and Climate (2011)change A.R. Ives, A.M. & Kilpatrick,J., Foufopoulos, to extinction. vulnerability andattributes history islands:life landbridge on extinctions Reptile (1999) J. Ives, A.R. & Foufopoulos, 253267. preysize.of effects populations: adder among body maximumsize dimorphismand sexual size in Variation (1991) A. Forsman, change. bodymass and dispersability of loss unifying the theory of evolution: insular New (2004) Ziv,Y.I.& Filin, , 7

, 5968., Brookesia

, 3 , 145154., This article isprotected by copyright. All rights reserved. ) from northern Madagascar. Madagascar. ) northern from Global EcologyandBiogeography Evolutionary Ecology Research Ecology Evolutionary The Journal of Animal Ecology Animal ofThe Journal The American Naturalist TheAmerican PlosOne Current Herpetology in East East Herpetology in Current The American Naturalist TheAmerican Global Environmental Environmental Global South American Journal Journal SouthAmerican , , 7 , e31314. e31314. , , , 6 , 115124. 115124. , , 153 , , , 125., 60 , , 177 , , 28

, , 635 634 633 632 631 630 629 628 627 626 625 624 623 622 621 620 619 618 617 616 615

Zootaxa Asia predation, drives lizard tail loss on islands.ontail loss lizard drives predation, Itescu, Y., Schwarz, R., Meiri, S. & Pafilis, P. (2017) Intraspecific competition, not competition, Intraspecific P.(2017)Pafilis, & S. Meiri, Y.,R.,Schwarz, Itescu, disagree. Turtles general? rule island Isthe S.(2014) Meiri, & P.C.H. Pritchard, P.,Raia, N.E., Y.,Karraker, Itescu, characteristics island of Y.(2017) Itescu, 173. the Indies. West diminutivelizardfrom new a vertebrates: limitamniote size in the lower(2001) At Thomas, R. S.B.& Hedges, Leptotyphlopidae: (Squamata: threadsnakes of species new two insnakes: size limit Atthelower S.B.(2008) Hedges, 5974. biogeography. ofmodel island equilibrium the onperspective largescale longterm, a disequilibrium: Dynamic(2000) L.R. Heaney, ( squirrel tricolored the mammals: from evidence sizeof insular body Island and (1978) area L.R. Heaney, ofJapan. Society

Author ManuscriptHerpetological The414432. pp. Goris), and R.C.T. Hikida M.(ed. Matui, by , , 1841

, 130. , Evolution on islands: testing the extent and consistency of the of effects and consistency the extent testing islands: Evolutionon This article isprotected by copyright. All rights reserved. Callosciurus prevosti Callosciurus . Ph.D. thesis, Tel Aviv University, Israel. AvivTel thesis, University, Ph.D. . Global EcologyandBiogeography Global Ecology and Biogeography and Ecology Global ) of Southeast Asia. SoutheastAsia. ) of Journal of AnimalEcology of Journal Leptotyphlops Caribbean journal of Scienceof journal Caribbean Global Ecology andEcologyBiogeography Global ) from the Lesser Antilles. theLesserAntilles. from) Evolution , ,

23 , 689700. , , , 86 , 32 , 6674. 6674. , , , 2944., 37 , 168 , , , 9 29 , ,

654 653 652 651 650 649 648 647 646 645 644 643 642 641 640 639 638 637 636 Page 34of50 Page 35of50 Herpetologica ofSciences Academy National Bulletin Herpetological Author masses. bills and beyond ofwarblers: Evolutionisland H. (2015) Winkler, & B. Leisler, http://CRAN.Rproject.org/package=metacor (2011) E. Laliberté, Science with lizards. aexperiment field differentiation: despiteadaptive persist effects Founder J.B.(2012) Losos, Spiller,& D.A. T.W., Schoener,M., Leal, J.J., Kolbe, Manuscript&Ecologyin Evolution Trends broken. cannot rulesbe (2008)Island R.W. &MoyàSolà,Wrangham, S. M., Köhler, populations. snake mainland island and among variation sizeBody (1989) R.B. King, 278. Lake. Pyramid around Island the mainland and Anaho Nevada'sof thereptiles divergencebetween bodysize desert island: ona Evolution (2013)& Feldman, Gienger,C.R. C.R., C.M. N.C.,Tracy, Nieto, J.E., Keehn, evidence. statisticalstandards for Revised (2013) V.E. Johnson, housegecko, Turkish theforsizes(2016) Record S.P. & Pafilis,Meiri, M., Moses, Y.,R.,Schwarz, Itescu, , Journal of AvianBiologyof Journal 335

, 10861089. 10861089. ,

, , 45 , 8488. , This article isprotected by copyright. All rights reserved. Metacor: Meta-analysis of correlation coefficients. R packageR of coefficients. correlation Meta-analysisMetacor: Hemidactylus turcicus Hemidactylus , , 138 Global EcologyandBiogeography , 2426. , , , , , 110 23 , 46 , 67. 67. , , 1931319317. 1931319317. , , 236244., , from Aegean islands, Greece. islands,Aegean from ,

Journal of Zoologyof Journal Proceedings of theof Proceedings , 291 The , 269 , .

671 674 673 672 670 669 668 667 666 665 664 663 662 661 660 659 658 657 656 655

Ecology & FloresVillela, O. (2012) Notes on activity, body size variation, and diet in insular diet andin body variation, size on activity, Notes O.(2012) FloresVillela, & AuthorE. K.,MociñoDeloya, Setser, Lawing,M.J.,A.M., S., Schaack,Ingrasci, J.M., Meik, (Viperidae:rattlesnakes speckled ininsular size evolution Body (2010) A. PiresdaSilva, & Lawing,A.M. J.M., Meik, landmasses. across extremes size body of mammalian size patterns area:and On(1998) P.A. &M.L. Taper, Marquet, snakes. in grass European dimorphism size sexual bodysizes and plasticity in (1993) R.Phenotypic Shine, T.& Madsen, Manuscriptany is‘best’modelgood? but the criteria, information using (2017) Model selection Yen,J.D.L. & J.R. Thomson, Duncan,R., R.P., Nally, Mac zoogeography. ofinsular theory Anequilibrium (1963) E.O. & R.H.Wilson, MacArthur, diversity. herpetofaunal Aegean’s of evolution the an archipelago: claiming Threecontinents N.(2010) Poulakakis, P.& Lymberakis, populations. lizard fieldexperimental inselection natural and shifts behaviour Predatorinduced (2004) D.A. Spiller, T.W. & Schoener, J.B., Losos, lizards. island in selectionnatural and predation (2011) Competition, R.M. Pringle,J.B. & Losos, rule. island thegeneralityof vertebrates: insular in evolution (2005) size Body M.V. Lomolino, , DOI: 10.1111/13652664.13060. 10.1111/13652664.13060. DOI: ,

Journal of Biogeography ofJournal

Nature Evolution This article isprotected by copyright. All rights reserved. , 475 , 17 , E1E2., Global EcologyandBiogeography , 373387., Evolutionary Ecology Evolutionary Crotalus mitchellii Crotalus , 32 Evolution , 16831699. , , 47 Diversity , 321325., ). PlosOne , 12 , 127139. , , Journal ofApplied Journal 2 Nature , 233255., , 5 , e9524., , , 432 , 505508. 505508. , 31

695 694 693 692 691 690 689 688 687 686 685 684 683 682 681 680 679 678 677 676 675 Page 36of50 Page 37of50 Biogeography Naturwissenschaften theory Evolutionary history approach. history lifeislands:a size on body in shifts Explainingadaptive E.P. (2003) Palkovacs, inlizard. insulara gigantism with associated are food availability high and competition Intraspecific(2009) Valakos, & J. E. Foufopoulos, S.,P.,Meiri, Pafilis, island and(1982) R.H.size Body Melton, island. onalarge size Borneo–small of(2008)K.M. Mammals Helgen, & C.P. S.A., Groves, E., Meijaard, Wich, S., Meiri, dimorphism. sexual to isunlinked speciesof number competitor The(2014) D. PincheiraDonoso, & P. Y.,Itescu,Raia, J., Foufopoulos, Pafilis,M.,P., A.E., Kadison,S.,Novosolov, Meiri, insular carnivores. in bodysize andArea,evolution isolation (2005) D.& Simberloff, T. Dayan, S., Meiri, lizardbody sizes. ecologyof and Evolution S.(2008) Meiri, 16 lizards.island in evolutionSize S. (2007) Meiri, 43 Mexico. Sea Cortés, thewesternof rattlesnakesfrom speckled Author Manuscript702708. , 556560. ,

, , 17 , 724734. , Oikos This article isprotected by copyright. All rights reserved. , Journal of AnimalEcology of Journal , , 6 Ecology Letters Ecology 96 , 113126., , , 11071113. 11071113. , 103 Global EcologyandBiogeography , 3744., Journal of Biogeography of Journal , , 8 , 12111217. 12111217. , Peromyscus , , 83 Global Ecology andEcologyBiogeography Global , 13021312. 13021312. , : a pattern and a hypothesis. hypothesis. a andpattern: a Global Ecology andEcology Global , , Herpetological Review Herpetological 35 , 10871094. 10871094. , , , 32

, 715 714 713 712 711 710 709 708 707 706 705 704 703 702 701 700 699 698 697 696

by G. Grigg, R. Shine and H. Ehmann), pp. 3546. Royal Zoological Society of New ofSociety ZoologicalRoyal 3546.pp. Ehmann), H. Shine and R. Grigg, G. by Naturalist American Zoology of Journal performance and behaviour between recently diverged populations of divergedpopulations recently between behaviour and performance mirror differences in predation pressure. predation indifferences mirror morphology, in Differences (2007) R. Damme, I.&Van Grbac, B., Vervust, nature. ofbalance theand(1973) PatternValen, L. Van Main. am Frankfurt J.(2008) Foufopoulos, P. & Maragou, P., K.,Lymberakis, P.,Sotiropoulos, Pafilis, E.D., Valakos, 100 oflizard. radiation a theinsular Trends (1966) in M. Soulé, ( tigersnakes Australian populationsof ofandmainland island comparisons Ecological (1987)R. Shine, Wales. South Australia.of South offshoreislands on snakes,blackof structure tiger (1985) Population T.D. Schwaner, convergence. and speciesdisplacement sympatric particularof the sizes with patternsIndian in West (1970) Size T.W. Schoener, populations. insular and of continental Astudy lizards.preliminary Mediterranean in extinction and (1997)autotomy Tail P. LoC. V., Corti,& Cascioa, PérezMellado,

Author4764. , Manuscript This article isprotected by copyright. All rights reserved. , , 243 , , 104 The amphibians and reptiles of Greece ofand reptiles The amphibians Notechis , 533541. 533541. , , 155174., Global EcologyandBiogeography : Elapidae).: Biology of Australasian frogs andreptiles frogs of Australasian Biology Oikos Herpetologica , 116 , 13431352. , Anolis Evolutionary theory Evolutionary lizards. II. Correlations II. Correlations lizards. , , The American American Naturalist The 43 . Edition Chimaira, Chimaira, Edition . , 233240. , Notechis ater niger Notechis ater Podarcis sicula Podarcis , , 1 The , 3149. 3149. , (ed. 33 ,

736 735 734 733 732 731 730 729 728 727 726 725 724 723 722 721 720 719 718 717 716 Page 38of50 Page 39of50

Author Manuscriptreptilepredators). aspotential theirindicating sourcesfor references 4 Appendix Appendix - 1 Appendix sources fordata references containing Appendices http://precedings.nature.com/documents/3360/version/1 size. gapeon selection orspandrel to response snake,inadaptive a dwarfism and gigantism Insular (2009) M. Hasegawa, Herrel,&A. T.,A., Mori, Kuriyama, S.E.,M.C., Brandley, Vincent,

3 Literature sources for isolation time of islands isolated before the LGM the beforeislands isolated of time isolation Literaturefor sources . A list of potential predator species included in our analyses (with analyses included our in species predatorApotentiallist of .

Reference list for literature containing body size data size body literature containing forReference list This article isprotected by copyright. All rights reserved. NaturePrecedings Global EcologyandBiogeography , , . .

. . 746 745 744 743 742 741 740 739 738 737

60 59 58 57 56 Snakes Lizards ru Family Group positive quadratic relationship;quadraticpositive ( sizes) are given are sizes)Tablein 2. 1.Table Boidae Gekkonidae Viperidae Colubridae Scincidae Lacertidae Summarythedataofand effectseachof predictoron eachspecies.(+)marks positive a correlation; a()negative correlation; (

Natrix natrix Elaphe quatuorlineata Eirenismodestus Lacertatrilineata Podarcismilensis Podarcisgaigeae Eryx jaculus Hemidactylusturcicus Mediodactyluskotschyi Dolichophis caspius Ablepharuskitaibelii Podarciserhardii Vipera ammodytes AuthorViperaxanthina Telescopus fallax ManuscriptOphisops elegans pce Ilns Individuals Islands Species

Populationsizemeanrefersrange SVL tolizards inlength total and in snakes.

∩

1 52 5.7–7.3 6.0 52.87 – 75.23 5323 118 1 5 6.5–900 91.6 846.9 566.25 – 910.00 1036.20– 1525.61 55 70 NS 11 69.9 11 8.4 107.33 – 141.85 57.37 – 77.67 231 641 16 14 7 7 4.4–5.9 3.3 42.04 – 59.89 270 27 6 01 35 98 2.8 33.52 – 49.88 2071 86 1 2 054 860 625.2 1085.40– 1886.00 72 11 5 5 213 9.5 81 NS 28.1 281.33 – 592.25 152 1.415 35.66 – 44.84 180 24 2 6 7.3–933 44.9 477.43 – 903.33 56 2.0 12 40.10 – 48.53 143 14 3 360 4.0 19.8 386.00 – 543.00 33 4.1 8 53.65 – 67.09 116 8 4 297 1.3 52.5 269.79 – 516.33 48 8 5 407 433 260 NS 276.0 490.78 – 1493.33 51 8 )negativequadratica relationship; (NS) nonsignificant correlation;Fullstatistics modelsallof (with sample This article is protected by copyright. All rights reserved. Population mean size rangesize (mm) Global EcologyandBiogeography meanbody mass (g)mass Species Area NS NS NS NS NS NS ∩ + + +

mainland fromthe Distance NS NS NS NS NS NS NS NS NS + +

Distance closestfrom larger island

NS NS NS NS NS NS NS NS NS NS NS NS NS NS + + +

Time since isolation NS NS NS S NS NS NS NS NS NS NS NS NS NS NS NS NS U + +

rdtr Competitors Predators NS NS NS NS NS NS NS NS NS NS NS N S

U ) a) 35 NS NS NS NS NS NS NS NS NS NS NS NS

∩ + +

Page 40of50 749 748 747 Page 41of50 Snakes Lizards Predictor n Species Family Group Lacertidae Viperidae Colubridae Scincidae Boidae Gekkonidae

transformed in species marked with (*). withmarked in species transformed 2. Table Natrix Natrix natrix* Elaphe Eirenis Lacerta Podarcis gaigeae* Podarcis fallax* Podarcis Podarcis Ablepharus Ablepharus Dolichophis Dolichophis Eryx jaculusEryx Hemidactylus Mediodactylus quatuorlineata modestus trilineata milensis elegans Ophisops Telescopus erhardii* kitaibelii caspius Vipera turcicus

Author Manuscriptkotschyi Best models by species. n is island sample size. Body size was log10 Bodysize.is samplenisland bymodels Bestspecies. 11 distance from distance mainland the 11 none 8 16 8 1 none 11 14 14 14 12 12 118 24 distance from distance mainland the 24 11 11 15 distance from distance mainland the 15 7 log area 27 86 86 8 This article isprotected by copyright. All rights reserved. predator richness predator predator richness predator predator richness predator distance from distance mainland the log area logisolationsince time log area log time logisolationsince time from logdistance largerclosestlandmass from distance mainland the log area (log area)^2 log area competitorrichness log distance from logdistance largerclosestlandmass from distance mainland the log area competitorrichness (^2) competitorrichness log area distance from distance mainland the Global EcologyandBiogeography

0.14 7.28 2.06 2.23 0.08 0.08 0.07 ()<0.01 0.09 0.09 0.55 447.95 326.08 0.01 ()<0.01 ()<0.01 0.38 0.38 2.81 0.86 0.67 0.67 3.06 26.20 5.77 .1 .9 63 10 <.1 0.26 <0.01 1.01 46.37 0.59 1.91 Slope SE Intercept SE P P SE Intercept SE Slope 27 04 18.9 75 <.1 0.77 <0.01 37.59 1482.29 0.48 2.79 .3 .1 91 07 <.1 0.30 <0.01 0.73 39.13 0.01 0.03 <.1 00 27 00 <.1 0.63 <0.01 0.04 2.74 <0.01 ()<0.01 0.05 3.01 0.68 0.74 0.02 0.02 <0.01 0.04 0.21 116.38 88.43 <0.01 <0.01 <0.01 0.12 0.81 0.44 0.28 1.00 3.56 0.55 3.2 7.45 136.02 3.80 48.63 .3 0.19 2.53 .4 0.26 2.04 712 391.42 2751.25 .0 <0.01 1.80 96 0.88 39.64 16 1.28 41.61 325 89.63 1302.57 36

0.01 0.03 0.03 0.03 <0.01 0.01 <0.01 0.04 0.03 <0.01 0.01 0.04 0.01 0.02 <0.01 <0.01 0.05 0.04 0.01 <0.01 <0.01 751 750 0.43 0.75 0.51 0.43 0.58 0.07 0.10 0.36 0.94 Adjusted R 2

ru Fml Seis Predictor n Species Family Group

xanthina*

0.71 6.94 Slope SE Intercept SE P P SE Intercept SE Slope 0.19 1.86 94 4.39 19.42 37

0.01 0.01 753 752 0.65 Adjusted R Page 42of50 2

Page 43of50 Predator richness richness Predator area Log Author Manuscript 0.30 0.03 0.13 richness Competitor 0.33 0.08 isolation since Time Log -0.20 island larger closest from Distance Log mainland the from Distance Predictors bold. in highlighted areSignificant results (g). bodyitslog studymass against 16 species of the variable in predictor each each and 3. Table

Results of the regression of the correlation coefficient (r) between body size size (r)between body coefficient thecorrelation regressionof of the Results

-0.30 0.12 0.22 0.20 0.03 0.29 0.29 0.03 0.20 0.22 0.12 -0.30 0.20 0.14 lp S Itret E R P SE Intercept SE Slope -0.30 0.09 0.36 0.15 0.01 0.42 0.42 0.01 0.15 0.36 0.09 -0.30 0.13 0.11 0.04 0.18 0.27 0.08 0.08 0.27 0.18 0.04 0.11 0.13

0.11 0.13

0.18 0.11

0.19 0.21

0.11 0.32

0.17 0.07 2

758 757 756 755 754

Competitor richness richness Competitor richness Predator t Log (years) (years) (km) island larger closest from distance Log (km) mainland the from Distance (km area Log Predictors ime since isolation since ime 2 ) 0.108 0.106 – 0.313 0.16 0.082 0.326 – 0.173 0.27 0.27 0.173 – 0.326 0.082 0.16 –0.313 0.106 0.108 )

H for are 4. Table

Author Manuscript 0.042 0.224 – 0.142 0.33 0.077 0.134 – 0.282 0.24 0.24 0.282 – 0.134 0.077 0.33 –0.142 0.224 0.042 0.17 –0.204 0.070 0.16 0.372 – 0.129 0.068 0.130 0.06 –0.055 0.433 0.201 0. 0.096 Mean r Range (r) P Mean r Range (r) P Mean r Range (r) P P (r) Range r Mean P (r) Range r Mean P (r) Range r Mean

0 Metaanalysis results for all reptiles, only lizards and only snakes. P valuessnakes.P only onlyand lizards reptiles, all for results Metaanalysis : r=0. Significant results are highlighted in bold. in highlighted resultsare Significant r=0.: 147

– –

0.3 0. 284 Global EcologyandBiogeography 2 0

0.06 0. 16

0.152 -0.028 – 0.323 0.05 0.05 0.323 – -0.028 0.152 0.003 0.088

0.190 0.278 Lizards Lizards – –

0.196 0.108

0.49 0.19

-0.263 -0.536 – 0.058 0.05 0.05 –0.058 -0.536 -0.263 <0.01 –-0.281 -0.777 -0.581 0.410 0.146 – 0.620 <0.01 <0.01 0.620 – 0.146 0.410 0.124 0.361 – 0.128 0.17 0.17 –0.128 0.361 0.124 0.423 0.179 – 0.618 <0.01 <0.01 0.618 – 0.179 <0.01 0.423 0.645 – 0.182 0.443 Snakes Snakes 39

763 762 761 760 759 Page 44of50 Page 45of50 Dolichophis caspius Dolichophis Full trend lines indicate a significant relationship (p<0.05). relationshipsignificant a indicate trend lines Full species xanthina Vipera ; quatuorlineata milensis turcicus Hemidactylus Figure1.

Author Manuscript c, d; Total length for species i, j, k, l; Log total length for species m, n, o, p.o,n,m, species Loglength l; totalk, for species i,j, for lengthd; Total c, ; f – Body size as a function of island area: ofarea: island sizeas a function Body

Ophisops elegans Ophisops

. m

Body size index is:index size Body – – This article isprotected by copyright. All rights reserved. ; Natrix natrix Natrix j ; ; – c – Eryx jaculus Eryx Podarcis erhardii Podarcis Global EcologyandBiogeography ; g – ; n ; Lacertatrilineata –

SVL for species a, b, e, f, g, h; Log SVL h; g,e, b, speciesf, a, SVL for ; k Telescopus Telescopus fallax – Eirenis modestus Eirenis ; d – a Podarcis gaigeae Podarcis – Mediodactylus kotschyi Mediodactylus h – ; o Ablepharus kitaibelii Ablepharus –

; l Vipera ammodytes Vipera – Elaphe ; e – – Podarcis ;

b for for – – ; ; i p – –

771 770 769 768 767 766 765 764

significant. richness. (km); (km); themainlandfrom distance – show:Panels species. each ofbody (g) mass mean log the andvariable predictor seetext) against each snakes,length totalfor lizards, for SVL 2. Figure d

Author Manuscript(years);sinceisolation timelog

n=16 species in all cases. Trend lines are shown only if they are statistically onlyaretheyifshown arelines Trend in all speciescases. n=16 The relationship between the correlation coefficient (r) of body length (mm;bodylength(r) of coefficient correlation thebetween relationshipThe

c – log distance from the closest largertheisland closestfromdistance –log e – predator richness; –predator a –

log island area (km arealog island f – competitor – 2 ); 41 b

779 778 777 776 775 774 773 772 Page 46of50 Page 47of50

Author Manuscript species. ( richness predator against seetext) snakes,length totalfor lizards,SVL (mm; for bodylength of 3. Figure

a The relationship between the absolute value of the correlation coefficient (r)coefficient theof value correlation absolute thebetween relationshipThe ) and competitor richness ( richness ) competitor and This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography b ) and the log mean body mass (g) each ofbody logmass mean theand )

784 783 782 781 780

Podarcis erhardii; Podarcis gaigeae; eg Podarcis– – milensis;Ophisops– elegans; f dPodarcis – quatuorlineata; m – Natrixnatrix;– m quatuorlineata; Telescopus– pfallax;– n ammodytes; Vipera– o Body xanthina. Vipera Figure 1. Body size as a function asizeislandMediodactylus –a Figure of area: Body 1. as kotschyi;bHemidactylus– turcicus; – c size index is: SVL for species a, b, e, f, e, forb, species h; g, for a, sizeLogis:SVL specieslength SVL index d;for species Total c, i, l; j,k, Log – Ablepharus kitaibelii; Ablepharus – Eirenis– i k –Dolichophis lmodestus; j caspius; Elaphe –jaculus; Eryx –

AuthorFull p. length trend total o, forspecies lines significant n, relationship m, a indicate (p<0.05). Manuscript This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 81x60mm (300 x 300 DPI) (300 30081x60mm DPI) x

Lacerta trilineata;Lacerta h

Page 48of50 Page 49of50

length for snakes, see predictor text) lengtheach forsnakes, against species. variable (g) each of bodylog the mass mean and Figure 2. The relationship FigurecorrelationforThe lizards,the 2. between SVL (r) bodycoefficient total lengthof (mm; Panels show: a – log– islanda show: Panels (km2); (km);bdistance –log – area c mainland the fromthefrom distance closestlarger(km); d- log island time since(years);competitor– f isolation e predator– richness;

Author Trendlines species richness.n=16 cases. are shownin all only ifstatistically are they significant. Manuscript This article isprotected by copyright. All rights reserved. Global EcologyandBiogeography 81x60mm (300 x 300 DPI) (300 30081x60mm DPI) x

Figure 3. The relationship Figurevalueabsolutecorrelation The the the 3. of between coefficientbody lengthof (r) (mm;

Author Manuscriptseeforlizards, text) length total SVL forsnakes, against competitorpredator richnessand (a) richness (b) This article isprotected by copyright. All rights reserved. and the log mean bodylogspecies. the mass (g) mean each and of Global EcologyandBiogeography 81x60mm (300 x 300 DPI) (300 30081x60mm DPI) x

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