Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. custo ffns uevso ftepoet n ocpuldsg.A rt h rtdato the of draft first the wrote to AS contributed both design. SM conceptual and revisions. and Availability: AA to Data project, substantially analyses. contributed the authors statistical of all the and supervision performed manuscript, funds, and designed of and acquisition data collected AS Authorship: of Statement title: running Short type: Article Israel a.slavenko@sheffield.ac.uk Aviv, author; Tel Corresponding History, – Natural * of Museum Steinhardt The USA – Honolulu, 4 Museum, Bishop Israel P. Aviv, Bernice Tel University, – UK Aviv 3 Sheffield, Tel Sheffield, Zoology, of of University School Sciences, – Plant 2 and Animal of Department – 1 tropical of Slavenko radiation Alex a in partitioning niche morphological lizards We with correlates species. poorly morphospace lowland of Competition than areas specific limbs occupy shorter conditions. to skinks with thermal on sub-optimal and morphospace selection in overall strong thinner, thermoregulation and impose efficient smaller, habitats other, facilitate average, highland each that in on to climates closer are, harsh tropical and that largest skinks narrower hypothesize predictor world’s Highland stronger become the a niches Guinea, is decreases. elevation increases New richness, elevation of occupancy distances species As fauna with and predictably overlap skink occupancy. change reduced the metrics morphospace and morphospace on of niches, niches some predictions narrower while cause these to that, to show lead tested We We predicted to island. expected is species. is different species richness of competitor species niches of in between increase presence dictating an the pressures Therefore, However, selection contract. ecological performance. and to microhabitat efficient on for based niches, adaptations ecological morphological species’ represent to expected is Morphology Abstract 2020 5, May 3 2 1 Slavenko lizards Alex tropical of radiation a niche in morphological partitioning with correlates poorly Competition e vvUniversity Aviv Tel Museum Bishop Sheffield of University The 1,2* Letters 1 le Allison Allen , le Allison Allen , opopc attoigo rpcllizards tropical of partitioning Morphospace 3 hiMeiri Shai , 2 n hiMeiri Shai and , 2,4 1 3 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. irhbttue ilatvt,rpoutv ilg n it(lio 92 lio re 96 Allison morphology, 1986; in Greer variation & much Allison display 1982; and Gui- (Allison island New diet the Papua and on of biology habitats fauna reproductive skink all activity, the almost diel in in use, partitioning occur microhabitat and Skinks occupancy (PNG). morphospace nea of Rabosky patterns 1989; examined (Greer We skinks more. Australian many in and loss 2010), and Skinner reduction limb 2009), Ricklefs al. eai irhbtt ntrepn tikeak Lvn&MPal18,18;MKno ude2002; Rundle & McKinnon evolution 1986; morphology 1985, al. et trophic McPhail & as (Lavin variation such stricklebacks Morphological McGee radiations, threespine adaptive classic in R traits. most of microhabitats 1998; morphological the cases pelagic (Sturmbauer of of impressive cichlids Some are most African traits. partitioning the in different niche of many to some for leading underlies occur 1974; divergence can always Pianka of partitioning not 1956; niche examples is Wilson to displacement leading & character divergence community-wide (Brown Trait diet such Meiri adaptation and said, 1981; morphological Boecklen That times in & between activity 2006). manifest (Simberloff reduced Grant use, thus manifest to & microhabitat is thought Grant Competition of are 1980; 1960). partitioning which Slatkin be Hardin via of to 1934; generating niches thought all (Gause force are similar space – similar species major occupying niche too Competing are taxa a the 2010). they closely-related is of Losos if 1998; partitioning and co-occur Travisano & ecology, to and Rainey unable in diversification 1974; promotes (Schoener processes species opportunity fundamental among Ecological most the diversity. among phenotypic is partitioning Niche selection sub-optimal strong in impose thermoregulation Introduction efficient habitats facilitate highland that in morphospace climates of areas harsh conditions. specific that limbs while thermal of occupy shorter hypothesize that, with to predictor We and show skinks thinner, overall stronger We on and smaller, a average, species. other, on each is island. lowland are, to elevation skinks closer tropical than Highland and richness, largest narrower decreases. become these world’s species occupancy niches tested morphospace the increases with We elevation Guinea, As predictably species. lead occupancy. New change different to morphospace of expected of metrics is niches fauna morphospace richness between skink species some distances in the and increase overlap on an reduced Therefore, predictions and contract. niches, to competitor narrower niches of presence cause to selection the to However, ecological predicted and performance. is microhabitat efficient species on for based adaptations niches, morphological ecological dictating species’ pressures represent to expected is Morphology Abstract Tables: Figures: in included References: and project text: R in an Words as packaged abstract: are in analyses Words the run to code Keywords: and study the S1. in Appendix used data All 05,saeadsz ieetaino rae nilsaoe Shee 94 oo 90 08 oo & Losos 2008; 1990, Losos 1974; (Schoener anoles Antilles Greater of differentiation size and shape 2005), 08 Lerner 2008; tal. et 1 5 cnia,toia clg,PpaNwGie,caatrdslcmn,cmeiieexclusion competitive displacement, character Guinea, New Papua ecology, tropical Scincidae, 03,bl vlto fHwia oecepr (Freed honeycreepers Hawaiian of evolution bill 2013), 88 4425 tal. et 150 01 n awnsfice Lc 97 rn 99 Abzhanov 1999; Grant 1947; (Lack finches Darwin’s and 2011) uber ¨ tal. et tal. et 01 tat&Lss2013). Losos & Stuart 2011; 99 R 1999; 2 br&Aas20) dpaint eti or benthic to adaptation 2001), Adams & uber ¨ tal. et 97 Lovette 1987; tal. et tal. et tal. et 07 in 2009; Wiens 2007; 04 Herrel 2004; 02 Reding 2002; et Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. l h nlsswr efre nRv.. RCr em21)adRcd saalbei pedxS1. Appendix in available is code R and 2019) species Team each Core of (R distribution v3.6.1 scheme. R the categorization in across same performed the were a.s.l.) using analyses m ”lowland” the or (in All ”highland” cell elevation either from per (mean to forest either elevation ”lowland” species montane to mean lower assigned or cells averaged define and assigned who we and (2015), a.s.l.) Similarly, cell Shearman m each & above. for Bryan 1000 a.s.l.) in m categorizations (3 [?] (in habitat System elevation elevation Information mean (mean Resource calculated Guinea then ”highland” New We Papua 2008). it. the Shearman in from & data occur elevation species obtained which We ( recorded PNG and of richness, grid species with projection skink cells Berhmann total excluding to equal-area cells, caliper km 620 digital 25*25 prising a a using unto taken maps Roll were the from measurements maps cases All distribution barring obtained instead. body, We the body of the mm. side of 0.1 left side nearest the right the from the ( taken from body damaged all taken in were were variation were limbs limbs history quantify of left natural to Measurements the in measurements 1a). where species, morphometric (Fig. recognized 16 shape currently recorded and 79 we size comprising specimen skinks, each PNG From of collections. specimens 1860 measured We collection Data Methods hypotheses: following (Fig. the test skinks to PNG PNG, of in specimens recognized currently 1860 110 from of measurements out morphological species used 79 We comprising 2018). 1a), Meiri 2009; 2007, Cmeiieecuin(as 94 lo 96 adn16)adlmtn iiaiy(aAtu & (MacArthur similarity competition limiting and diffuse 1960) to Hardin due 1946; decrease Elton will 1934; (Gause species exclusion between Competitive overlap 5. niche more increases, becomes morphospace richness the skink as decrease As will 4. species between and distances distances increases, relative richness the skink but As richness, with 3. greatly very not will morphospace the of size overall The ( narrower 2. become will niches increases, richness skink As 1. o hnighbtt n niomna odtosi hpn ih attoigi hslarge this in partitioning role niche morphospace strong shaping that a in suggesting show assemblages, conditions lizards. also lowland environmental of We to and radiation compared skinks. habitats morphospace reduced, changing PNG likely of a for both area therefore in occupying distinct, assemblages was partitionining a highland Competition with niche and elevation, morphospace. with of strongly of changes driver occupancy areas strong similar sympatric a and occupy richness, not with species correlate poorly allopatric metrics and occupation morphospace that allo- species’ chance. phylogenetic here sympatric of by show and whereby We that expected pattern, filtering than with opposite habitat similar more the Alternatively, more overlap create 1g). is may will morphology morphology (Fig. species morphology its proximate ones of species spatially distance sympatric between and of the relatedness with distances 1f), than than longer mean (Fig. be relatives species the will patric allopatric Therefore, species of sympatric co-occurring. and those from species and focal species a similar of 1e). prevent morphology (Fig. will 1974) 1972, 1967) (Pianka Levins areas species-rich in species competing closely 1c). between (Fig. overlap change reducing will into partitioned remains is “pie” “pie” 1d). the morphological (Fig. proverbial that saturated “slices” the the of of size size the the is, but That same, will. the species various between overlaps 1b). Fig. morphospace; of area e.g. < 0 adcvrt rvn deeet.Frec el etlidthe tallied we cell, each For effects. edge prevent to cover land 50% isn iis,i hc aealmaueet o htspecimen that for measurements all case which in digits), missing tal. et 21)fralteseisicue ntesuy eoverlayed We study. the in included species the all for (2017) 3 i.e. niiulseiswl cuyasmaller a occupy will species individual , sensu < Tallowin 00masl,bsdon based a.s.l), m 1000 tal. et rd ca. dto;Bryan Edition; 07,com- 2017), 00mand m 1000 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. hog h rgnwt endsac/vra ihntefclseis ag ntexai,admean and axis, x the on range regression species’ linear a focal its (b) the outside and range, within and species’ distance/overlap range, overlaps focal and mean its the distances for within outside with mean and the species cells origin range species, all focal each the species’ for across the focal through comparing, the species from t-tests within (a) overlap focal cells tests: and the between two which performed distance from then and mean distances We range, mean the range. its within the calculated their are calculated then in cells We and morphospace which species of ( each range. areas cell for its similar each tallying sharing outside by this species are did between cells We competition compare ”highland” 1f-g). of to either (Fig. signals analysis distributions to PERMANOVA for a species tested used we each species. we Finally, lowland assigning section), cell and After collection” per highland ”data occupied between lowlands. the morphospace occupancy in and overall morphospace described the highlands (as if the ”lowland” examined in or also we species analyses, between described scores shifted skewness previously PC deviation, the of cell. standard distributions to each the the Similar in against in elevation scores changes mean examined PC of and of generated the models cell we kurtosis Furthermore, regression each quantify and linear for cells. to fitting PC highland each by metric and elevation on lowland a with between scores centroids PC as PC the used performing the of by in we histograms cells shift lowland which the a to from there’s cell, highland function if compared each ’adonis’ examine We the for with cell. each analysis scores PERMANOVA in a PC occupied morphospace of of centroid area commonest the calculated the further from We function we ’calc.relimp’ model, the full in the implemented was as model) model to null best (the contribution the none etc. If the ( or score. decomposing orderings richness), AICc by different or lowest over importance elevation the variable (mean with as them relative model richness of the compared species selecting one and scores, model), elevation AICc (full mean Factor using predictors Inflation with Variance of both metric, strongly, 1) either each co-vary to had not for 0 do models from (they regression variables (ranging of linear predictor ’hypervolume index centroids multiple similarity the ran the Jaccard (using pairwise then the between species We mean as of density distance defined the pair kernel Euclidean cell, each (c) the ’hypervolume the 95% of cell; in the volumes as Gaussian pairs (using the total the defined the species species the in cell, all of (b) as occurring between the pair cell; overlap defined species each in the of cell, all pairs in volumes of the species species the individuals different in all all by of the between species occupied of coordinates distance all volumes morphospace the by of morphospace around volume the occupied estimate mean on morphospace the based of (a) cell, volume each 1b-e): for (Fig. metrics species morphospace four calculated We morphospace species each of of analyses individuals Spatial of coordinates the around estimate function) morphospace. density of in used probability We portion the skinks. significant estimate PNG statistically to of ’hypervolume a morphospace the the explained (using define which species to axes used were PC PCs those These the only analyses. by downstream select data measure- for the morphometric to variance size-adjusted from the 1993) 15 function in (Jackson by ’prcomp’ and dimensionality the them reduced method mm), We using dividing (in SVL). 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Data may be preliminary. hra enpiws vra hne h es,a infidb h ieecsi tnadzdregression standardized in differences elevation, the with by most signified average the as the changes least, volume the volume), Total changes (total overlap overlap). decreases pairwise (mean pairwise between mean (mean assemblages decreases overlap average whereas increases species by the and morphospace by distance), occupied in pairwise occupied (mean morphospace species decreases morphospace morphospace of of in species area area between distance average total in the positions the increases, similar volume), elevation occupy species as from average, average, differ on species. not On that allopatric do suggesting or ranges 4), differ sympatric their (Fig. not within either The ranges species as do other their 1f-g). morphospace range, and outside respectively; species’ species species 0.02, (Fig. each within focal of +- between axis the parameters those overlap 0.98 y outside and the and distances parameters the when 0.01 mean the overlap, on +- the against pairwise (0.99 range compared 1 mean species’ from are and focal significantly range distance the species’ pairwise outside range focal mean species’ distance/overlap the both focal mean for the within and slopes distance/overlap axis, regression that mean with suggests x origin, This the the through S1). species. on regressions (Table distances sympatric linear ranges higher from ran their both occupancy then We outside have morphospace than (31.2%) in ranges species differ their 24 significantly species within only species with overlap However, species few lower with significantly ranges. have overlap their (39%) lower outside species and within 77 than species of ranges from 30 distances their and occurring higher ranges, significantly within species pairwise have their and (51.9%) outside mean species species than each 77 elevation. ranges and between of mean their overlaps 40 volume, as allopatry, and or important total distances sympatry as pairwise volume, either mean nearly in compare mean richness to species for t-tests is is on richness overlap elevation Based pairwise skink Mean mean whereas than mean for richness. – Only important with increasing richness 1). more with skink increases (Table times increase distance with overlap mean distance 17.7 decrease and pairwise pairwise overlap, to volume, mean mean pairwise 2.4 and morphospace whereas mean volume total and elevation, morphospace volume, volume, total mean morphospace morphospace with Mean Mean decrease elevation. all volume). Model both total distance by 3b). (for explained pairwise 1). (Fig. metrics 64.2% (Table Peninsula all Papuan elevation to in variation the mean overlap) mid-elevations have in metrics and in Mts morphospace richness peaking Stanley all mostly skink Owen for the models pattern, northern regression the and complex linear along Cordillera most Multiple and Central the Cordillera the shows Central peaks of the overlap volume slopes in high pairwise niche lower the the Mean Mean is Mean on in and 3a,d). 3c). decrease 3). 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Data may be preliminary. h vrl pta atrso pce ihesadmrhsaeocpnyaentlkl ocag greatly. change to likely not are occupancy morphospace and into richness species species cryptic of of ( patterns complexes species spatial wide-ranging overall similar splitting the morphologically of be and will allopatric changes mostly taxonomic if However, ( underestimated determined. described. greatly be potentially is to al. skinks competition, need extent Guinea experience et elevation, New The therefore Papua with and of another. are and diversity PNG, one taxonomic spatially across cell Third, with change other ecologically km each co-occurrence interact with of PNG. 25*25 them interact in patterns of cell a cells whether same few all and meaning the within cell, for 1982), in same (Allison true occur species elevation the isn’t which and that in likely to microhabitat very co-occuring by species separated and all are tested despite all Kiandi here that Mt that on not species is assumes skink that instance, here assumption For an employed others. syntopic, we than and sympatric strongly method more space the trait Second, of axes which some these, measurements (Irschick affect as may morphometric performance such competition Traits the locomotory 1974). by on (Greer captured habits impacts Macrini not arboreal documented their genera are well with the that from association have space skinks in likely of niche cases lamellae, species of other several subdigital several instance, axes For competition. in other of here. described be effect used been weak may has the there for what structure First, Furthermore, explanations to assemblage possible similar 1). driven several skinks, Table are having PNG 3; There similarity in weak. limiting (Fig. Therefore, or competition S1). interspecific exclusion Table ( for competitive & metric 4 of habitat a and evidence (Fig. conditions richness, environmental little species for is we occu- proxy by However, a Morphospace there is elevation, 1974). skinks. by it Pianka influenced PNG than strongly 1957; in type, more (Hutchinson partitioning be niche competition to ecological diffuse appears competition-driven of adapt pancy of will effects morphology evidence the species little to increases, found competitors due of similarity number reduce the as to that, predicts theory ecological partitioning Classic niche competition-driven of evidence Little lowland from Discussion species Similarly, power. permutations, S5). explanatory 999 low is on PC2 very Fig. There based of has (PERMANOVA centroids S2; model morphospace S5). the Table the of and 5b-c; Fig. increasing areas PC1 distinct S2; (Fig. become of 0.001, Table occupy thus elevation centroids cells 5b-c; the cells in highland with the increase and (Fig. values, in an values modal centroids the with modal of in decreasing the shift distributions standard and around pronounced The the smaller centered a and average, PC1 also increase. more on of PC2 and kurtosis have, right-skewed and of cells skewness more kurtosis highland the increases, meaning and elevation skewness cells, As lowland deviation, limbs. than shorter scores with skinks PC2 thinner based lower (PERMANOVA S3). and cells (Fig. lowland scores from highlands PC1 the morphospace the from of permutations, from species area cells 999 species different that highland on than significantly means from a morphospace This occupy species of cells species). encompassing area Highland highland ellipsoid larger for CI a 72.01 95% occupy species, the PNG from lowland as of species for lowlands large encompassing 106.11 ellipsoid as ellipsoids: CI times of 95% 1.5 (area the roughly Furthermore, is 1). cells (Table lowland metrics different the for coefficients e.g., Meiri R 00.Teeoe atrso opopc cuac ol oetal hnea e pce are species new as change potentially could occupancy morphospace of patterns Therefore, 2020). 2 tal. et .2;Fg 3,abi h ieec smc mle hnteasmlg-ee oprsnand comparison assemblage-level the than smaller much is difference the albeit S3), Fig. 0.027; = tal. et 03 ltot&Ishc 04,aelkl motn nmcoaia ih attoig and partitioning, niche microhabitat in important likely are 2004), Irschick & Elstrott 2003; 01 tat&Lss21) h ffcso optto nsaigncepriinn appear partitioning niche shaping on competition of effects the 2013), Losos & Stuart 2011; p 0.001, = R 2 .7 i.5.Cnriso ihadclshv,o vrg,higher average, on have, cells highland of Centroids 5). Fig. 0.17; = 6 e.g. , Papuascincus Prasinohaema tal. et p. Slavenko spp.; 96 lsi oo 1997; Losos & Glossip 1996; and Lipinia tal. et aedeveloped have e.g. 00,then 2020), Slavenko , p = Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. olns(i.S) hrfr,tedffrne ewe oln n ihadasmlgsaedrvdfrom the derived in are found assemblages much to morphologies highland so the lead and not of lowland might do between subset 1995) PNG differences diminished Hauser in the a & species Therefore, rather highland Heard but S4). However, morphospace, 1991; (Fig. of landscape. Larson lowlands area adaptive & distinct the (Baum a of innovations represent areas key several novel of in of arise evolution colonisations could driving the area regimes morphospace and selection distinct in different 2012) a differences to occupy (Pinto Such lead also could evolution cells 5). differences morphological environmental highland (Fig. instance, general, For cells in mechanisms. lowland morphospace distinct to of compared portion smaller morphospace morphospace a of occupying of as areas well distinct As occupy species highland and very Lowland morphologically to to related reason closely no phylogenetically have is as we S4). 1982) such and Simon species unknown, highland which & is dissimilar to (Greer skinks extent species of the lowlands However, radiation distinct this impossible. in currently highland assume occurs analyses many conservatism these of niche for affinities phylogenetic phylogenetic correction close phylogenetic the a by rendering explained be simply (Rodriguez may nichespecies elevations phylogenetic If high in 2009). to (Revell occupancy failing analyses and morphological independent, such phylogenetically in (Blomberg of not conservatism results are times study impact several this may phylogeny distribution in themselves, for species lowland in account the to radiation that is back distinct caveat a shifts important Another not for and sources lowlands, provided the and as past. colonised skinks the been of radiation both same have the and of extension an for are – origins lowland Skinner from from colonisations independent separate represent also for highland likely instance landmasses instance, species For skink distinct highland case. other ( form the Several remain skinks not highlands montane is taxa mostly and this of highland suggest lowlands clade and to the though, lowland several evidence, if as between is well ascertain transitions There to of radiation. of Mahler number difficult events 2005; the it Vose and is & unresolved, (Gavrilets unknown, islands are large skinks on Guinea only morphological evolve in al. differences morphologies et explain unique to where attempts anoles, wider, and (Mahler Caribbean enable large systems and island also small to may highlands. between above ascribes diversity lowlands productive and originally less the below effect” the areas in “area in between This productivity 2/1 than adaptive primary of there the ratio higher space a Additionally, niche of roughly diverse areas highlands, 2009). more Mahler novel the (Allison than 2010; occupy m area Barraclough to in 1000 & extensive potential Kisel more the indeed 2005; are providing Vose therefore areas & larger and – (Gavrilets effect” diversification, “area landscape the for by explained opportunities be might more lowlands in diversity morphological larger SlavenkoThe from data mass the skinks; occupy long-limbed Guinea which large-headed unusually New of Guinea, and Papua semi-aquatic other New than the Papua heavier, include in times these skinks – 3d,h distinct species (Figs. bumui lowland morphologically species Fojia are most lowland morphospace, do the the than of of morphospace many margins of portion Indeed, smaller S3). a occupy & species elevation highland with that found changes We morphospace occupied of size The 03,i suceri hsseai ple oNwGie.Snepyoeei ffiiiso otNew most of affinities phylogenetic Since Guinea. 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Emoia , Papuascincus tal. et oj bumui Foija Prasinohaema pce pyoeei oiin of positions (phylogenetic species 03.Wieti a etecs o oessessc as such systems some for case the be may this While 2013). p.and spp. 7 iiu gigas Tiliqua n highland and Prasinohaema neteeymrhlgclyadecologically and morphologically extremely an , ( .virens P. Prasinohaema; o vrg . ie ogr n 290 and longer, times 4.7 average (on and tal. et p.(Rodriguez spp. .semoni P. tal. et 03.Telwad nPNG in lowlands The 2013). Sphenomorphus 06,o eea species several or 2016), Rodriguez ,aense ihna within nested are ), tal. et oi bumui Fojia tal. et tal. et 08 (Fig. 2018) and Emoia 2010). 2018). tal. et as , Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. etakMlyHgmn BrieP ihpMsu) hi utnadSt akr(oiin State (Louisiana accesses Parker facilitating for Seth Zoology) and Comparative Austin of (Museum Chris Rosado Museum), Jose and Bishop Zoology), P. of (Bernice Museum University Hagemann Molly thank We partitioning niche not and due competition tested. assembly Acknowledgements pressures that clearly community selective recommend and of skinks, quantified strongly driver that PNG We be strong suggests for always a important. least should evidence be more at However, to be that, taken may and be 2006). conditions ( conclusion, environmental latter Grant partitioning Wilson harsh niche the & & of to support (Brown Grant driver results niche strongest Our 1980; same the Slatkin 2013). the always in Pianka not 1974; co-exist 1957; is Pianka (Hutchinson cannot competition partitioning 1960; competitors niche that Hardin of predicts driver 1956; theory foremost a ecological be and to 1974), considered classically is Competition retention. and even acquisition or heat Conclusions morphologies, of different food rates with passive limited species compare of for to climatic efficiency competition models thermoregulatory harsh inert the as the comparing using such performance by that interactions reduced tested to suggest be intraspecific lead suboptimal could We but in are opportunities(Irschick lowlands, particularly activity. mating that the or of climates, Morphologies for resources climates highland window mild evolution. the harsh this morphological in in of adequate drive be PNG use may in thermoregulation the faster for habitats and effective efficient montane more more of the the conditions and is, temperatures, optimal gain achieve heat to sunbathing require ectotherms gain Therefore, heat sizes improve body may al. small lizards et – (Slavenko as thermoregulation such dubious ectotherms be is (Monasterio in to (Carothers reptiles which purported climates in ratios, is cold area-to-volume relationship change in surface a this increased such behind to for mechanism lead driving evidence of The although richness species contented, 2019). (Greer indeed been ovoviviparity (Sarmiento and long as lizards, levels as such has freezing such climate (Tallowin reaching ectotherms PNG cold for temperatures in for ideal nightly elevation temperatures from with mean and far decreases with are 2009), frosts, reptiles climates (Allison occasional harsh regions in with 5-6degC These montane rate temperatures that as 1986). lapse meaning cold The low 2010), extremely (Bourke climate. as by altitude is typified dropping in skinks are gained highland Guinea m in 100 New shape every of and for size 0.5-0.6degC roughly body is on PNG selection these for in avenue length possible limb relative Another 1986; short Greer for pressure & selective highland (Allison provide the gaps might of grasslands skinks. tree-fall dense Many and in 2000). shrublands Locomotion Swain grasslands, 1986). & alpine (Melville in limbed occur shorter Greer Goodman skinks and 2000; smaller Guinea Swain be New sprint- Irschick & to Papua faster (Melville 2000; tend limbs with vegetation Swain longer dense associated & have and therefore typically Melville to and are and tend 1969; performance, limbs habitats thinner (Pianka al. locomotory Longer open smaller, capability to occupying average, jumping related 1994). species on is enhanced Losos Therefore, are, length and & limb cells speeds Jr Relative highland (Garland ing 5). from drastically, use & Skinks changes microhabitat 2 above habitat to 2015). entirely (Figs. the Shearman disappears limbs eventually and & shorter and with (Bryan shift, decreases line rainfall height canopy tree of as the the patterns stratification than reduced drop, rather with 5b,c), temperatures particularly (Fig. increases, assemblages elevation the assemblages. in As highland morphologies in of morphologies values unique modal of and distributions presence the in shift a 08 u see but 2008; tal. et 06.I iheeain,lwabettmeaue edt hr idw vial o activity. for available windows short to lead temperatures ambient low elevations, high In 2016). tal. et 05,btsm,sc as such some, but 2005), e.g. 09 Zamora-Camacho 2009; cut II Schulte , tal. et tal. et 97.Temrglto a eeteeydffiuti otn habitats montane in difficult extremely be may Thermoregulation 1997). 04.Cnesl,abra n ruddeln pce nla litter leaf in species ground-dwelling and arboreal Conversely, 2004). Papuascincus tal. et tal. et tal. et 96 Macrini 1996; 04,adteeoenest eepcal ffiin (Ortega efficient especially be to needs therefore and 2014), tal. et 8 05.Arltosi ewe oysz n climate and size body between relationship A 2005). p. loihbtrcysbtae Alsn&Greer & (Allison substrates rocky inhabit also spp., 07.Mn otn pce xii adaptations exhibit species montane Many 2017). tal. et 03 Pinto 2003; e.g. Meiri priori a tal. et tal. et 08.Ti hypothesis This 2008). 01 tat&Losos & Stuart 2011; n htiseffects its that and , tal. et 2005). tal. et et Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. ags h ahdln ersnstenl ersinwt nitreto n lp f1 f The (f) paradigm. 1. competition of ecological species’ slope the range, focal of species’ a the expectations focal and within the the higher 0 represent within are areas of lower that shaded are intercept their species that an outside focal species and with species the focal within from regression of species distances null pairs tested focal mean all the Hypotheses the range, represents between with represents (f-g) area overlap overlaps line shaded outlines. the mean all dashed blue dark and represented as of with from The centroids calculated mean distance polygons the the is mean shaded assemblage, ranges. of as purple distance the comparison calculated as in pairwise the here is species Mean for represented overlap of (d) assemblage, Mean pairs the (e) all in shape. of species hexagons. grey centroids all by from between the individuals coloured here distances by by by all occupied here here morphospace as of represented represented of calculated assemblage, blue mean volume is assemblage, the the by volume in as the denoted Mean Schematic calculated morphospace pes (b) in (b-e) is of species, – volume PL). volumes species. Total skink and PL species’ different (c) CL three individual a heel); FL, elipses. representing of all of of to each of (sum assemblage tip knee stars, limb mean length hypothetical yellow hind the – (from to of and FrL a length (wrist length circles are: of – crus red length here HiL morphospace displayed squares, – manus ML); not the arm and CL measurements – FaL – of additional pectoral knee); AL, ML AL Two representations – of to depth; digit). wrist); (sum PeW girdle longest pelvis limb to point); front of pelvic (from of tip deepst (elbow – length to the length PlD heel femur (at width; (from forearm to – length girdle depth – opening FL pelvic head FaL ear – digit); – of elbow); PlW HD longest edge depth; to point); anterior girdle (shoulder widest (from pectoral length the length – (at head PeD width width; – head girdle HL snout-vent-length; – – HW SVL snout); are: Finally, measurements A.S. The to 1. 00020309000 provided Figure who Guinea lands. no. New collaboration ancestral Papua grant gracious in their Conservation communities the BSF was to PNG local through the and access research countless accomplished Institute, and Research This only A.A., Authority, National Protection was PNG and Environment study. study the and governments, S.M. this this provincial in of to in years used the used 2012143 specimens throughout all specimens of no. the collections grant of the measurements BSF permitting by and funded collections their to i.e. ersinsoesalwrta .()Teprl hddae ersnsma vraswt the with overlaps mean represents area shaded purple The (g) 1. than shallower slope regression a ceai ersnaino kn ih1 opoercmaueet akdo it. on marked measurements morphometric 14 with skink a of representation schematic A (a) 9 i.e. ersinsoeseprta .Tu these Thus 1. than steeper slope regression Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. opoercmaueet nteP xs nteegso ahai r ceai ersnain of representations schematic are axis each of axis. edges each the along On changes morphology axes. how PC visualizing skinks the on measurements morphometric 2. Figure opopc fPpaNwGie kns h learw hwtelaig ftevarious the of loadings the show arrows blue The skinks. Guinea New Papua of Morphospace 10 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. aho h ormtisfo asad(ett ih)adma elevation. between mean relationship and the right) showing to warmer Scatterplots metrics (left (h) whereas a-d four values, richness. maps the lower from species of metrics species represent skink each four skink between and colours the (e) right) relationship Cooler of to cell; the each (left showing per cell. a-d volume Scatterplots per maps total (g) elevation from (d) values. mean cell; higher (f) per represent volume and colours mean cell, (c) per cell, richness per coefficient similarity Jaccard 3. Figure af assoigsaildsrbtoso a enpiws itneprcl,()ma pairwise mean (b) cell, per distance pairwise mean (a) of distributions spatial showing Maps (a-f) 11 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. ntesgicneo -et oprn h ercwti n usd h oa pce’rne–turquoise – range based ranges. species’ coloured between focal is difference the dot significant Each outside no for and 1. pink within of and metric slope ranges, the regression between comparing null difference t-tests significant the for represents of line significance dashed the The on ranges. their outside 4. Figure oprsn f()ma itnefo n b enoelpwt h oa pce ihnand within species focal the with overlap mean (b) and from distance mean (a) of Comparisons 12 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. iue5. Figure a ctepo hwn h oriae ftecnriso C e el ihadcells Highland cell. per 2 & 1 PCs of centroids the of coordinates the showing Scatterplot (a) 13 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. Csoe ihnec el(ahhsormi h itiuino nidvda el.Clusrpeetthe represent Colours representing colours cell). warmer and individual elevations an lower of representing colours distribution of cooler the distribution elevations. with is the higher cell, showing histogram the histograms (each of Density elevation cell (b-c) mean each group. within each scores for observations PC of 95% encompass ellipses (elevation > 00m r oordbu,adlwadcls(elevation cells lowland and blue, coloured are m) 1000 14 < 00m r oordta.The teal. coloured are m) 1000 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. aohr .. o .. aqe ..&Jki ..(97.Temlcaatrsiso e nenlizards Andean ten of characteristics Thermal (1997). F.M. genus Jaksic the & P.A. of Marquet S.F., Fox J.H., Carothers 2002-2014 period the (2015). P.L. over Shearman Guinea. & New J.E. Papua Bryan Moresby, Port Guinea, New (2008). Papua P. of In: Shearman & Displacement. J.E. Character Guinea. Bryan (1956). New https://CRAN.R- E.O. Papua Wilson in & In: 473-512. W.L. species LandscapesBrown us- pp. Terrestrial crop Canberra, on operations economic Press Influence hulls. ANU 230 set Human M). of convex and and Prebble limits Climate geometry and Altitudinal Fire, dimensional Ecologies: machines, (2010). High vector R.M. Bourke hypervolume: support data: estimation, comparative (2018). in density project.org/package=hypervolume. D.J. signal 0.85 phylogenetic kernal Harris for ing & Testing 0.71 0.56 B. (2003). A.R. Blonder Ives labile. 0.95 & more character are T. traits studying behavioral for Jr. methodology Garland phylogenetic S.P., a Blomberg reviewed: Adaptation (1991). A. macroevolution. Larson & D.A. Baum body and New biogeography of historical skinks Evolution Phylogeny, genus Crocodile and (2010). new Island G.R. Pacific a Zug in for & evolution S.J. basis size Richards structures: E.N., Rittmeyer surface C.C., pustulate Austin with shells Scincidae). Egg (Lacertilia: (1986). 652-658. skinks A.E. Guinea pp. Greer 0.15 California, & Angeles, A. Los Allison In: 0.29 and 0.67 Berkley 0.44 Biology. Press Guinea, California 0.35 New of 0.16 University 564-616. 0.05 (2009). pp. A. Singapore, Allison Editions In: 0.39 Periplus Papua. BM). of Beehler Herpetofauna & In: 4.6. AJ Section lizards. (2007). Guinea A. New Allison of ecology and -0.75 Distribution Guinea (1982). (2004). A. -0.5 C.J. Allison 0.29 Tabin finches. & P.R. Darwin’s Grant in -0.61 B.R., beaks Grant of M., Protas A., Abzhanov Elevation Mean References Richness 0.31 Skink -0.33 -0.26 0.13 Overlap Pairwise Mean Distance Pairwise Mean Volume Total Volume Mean model. Model each in predictors significant; both are for predictors calculated (all importance relative predictor each for written 1. Table e.GestJ) pigrTeHge ehrad,p.803-813. pp. Netherlands, Hague, The Springer JL). Gressit (ed. eut ftebs oesfrec opopc erc tnadzdrgeso offiinsare coefficients regression Standardized metric. morphospace each for models best the of Results Liolaemus 7 227-236. 57, , ytmtcZoology Systematic ncnrlChile. central in nvriyo au e una otMrsy au e Guinea. New Papua Moresby, Port Guinea, New Papua of University . Science Evolution au e unarsuc nomto handbook information resource Guinea New Papua h tt ftefrsso au e una21:mauigchange measuring 2014: Guinea New Papua of forests the of state The 0 1-18. 40, , 0,1462-1465. 305, , ora fHerpetology of Journal eit hln eHsoi Natural Historia de Chilena Revista 7 717-745. 57, , Tribolonotus nylpdao islands of Encyclopedia 15 p h clg fPpa atOne Part Papua, of Ecology The Sumt;Scincidae). (Squamata; < 0 116-119. 20, , ytmtcZoology Systematic .0) h w ih-otclmsaethe are columns right-most two The 0.001). R 2 es ilsi G&Cau DA). Clague & RG Gillespie (eds. Bmp4 igorpyadeooyo New of ecology and Biogeography ihesRltv motneEeainRltv Importance Relative Elevation Importance Relative Richness es ael ,SeesnJ& J Stevenson S, Haberle (eds. 0 297-309. 70, , n opooia variation morphological and ,49-64. 5, , oeua Phylogenetics Molecular r d.University edn. 3rd . es Marshall (eds. Altered Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. 20) oprsno aia s,mrhlg,cign efrac n saebhvoraogtwo among behaviour escape A. and Herrel & performance J.A.Y. clinging Meyers morphology, B., use, Vanhooydonck C., habitat Buckley of M., of comparison Ramos analysis A J., comparative Elstrott A (2005). E., (1996). Carlisle O. Ellers D.J., Irschick & J.B. lizards. Losos pad-bearing R.N., among Fisher ability K., clinging Petren key animals? 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Reilly & PC Wainwright (eds. rneo nvriyPes rneo,New Princeton, Press, University Princeton . 3,1292-1297. 131, , Lobulia uryBat n os Chipping Sons, and Beatty Surrey . rceig fteNtoa Academy National the of Proceedings Lcrii:Sicde rmhigh from Scincidae) (Lacertilia: Leiolopisma ora fAia Ecology Animal of Journal 3 389-398. 83, , 9 21-35. 59, , n t relatives. its and ora fStati- of Journal h American The Science Herpetologica Historical 313, , , Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. er . ibro .&DynT 21) omnt-iecaatrdslcmn ntepeec fclines: of presence the guilds. in weasel displacement design. character Holarctic Community-wide evolutionary of (2011). successful test T. Dayan a a & around D. variation Simberloff S., world: Meiri the of lizards Biogeography the and of Ecology Traits (2018). systems. 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Perkins Z.B., studies. comparative Rodriguez interspecific for components 3258-3268. principal 63, and , Size-correction (2009). ‘creepers’ L.J. of evolution Revell Convergent (2008). radiation. R.C. Fleischer honeycreeper & Hawaiian H.D. environment. the Pratt heterogeneous in H.F., a James J.T., in Foster radiation D.M., Adaptive Reding (1998). clade. M. vertebrate Travisano diverse & in P.B. variation most Sciences Rainey among-lineage Biological Australia’s Exceptional B: of London (2007). of radiation I.J. Society Lovette the Royal & during A.L. rates Talaba diversification S.C., Donnellan D.L., Rabosky Austria. Vienna, amphi- Computing, tropical high (2019). Team of Core causes R the reveal analyses radiation: phylogenetic Large-scale adaptive In: diversity. (2013). in bian J.J. effect Wiens & island mainland R.A. 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Western in ) aueEooy&Evolution & Ecology Nature rceig fteNtoa cdm fSciences of Academy National the of Proceedings h mrcnNaturalist American The ilgclJunlo h ina Society Linnean the of Journal Biological 6 1673-1684. 36, , . ora fTemlBiology Thermal of Journal onainfrStatistical for Foundation R Anolis Ecology 0,581-588. 106, , ,1677-1682. 1, , lizards. rceig fthe of Proceedings Nature 0 1012-1030. 50, , 6 10230- 96, , Proceedings 9,69. 394, , Evolution 56, , Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. ttsial infiatprino aine ntebto ae,tesm C r h nsweethe where ones bars). the (red are model explain stick” PCs that ”broken same PCs a the under are panel, eigenvalue) expectation (average the bottom line than the higher red In is the bars) variance. above (yellow values of explained panel, variance portion top significant the statistically In a variance. of portion an S1. in Figure size gradient. body elevational rules 2200-metre S1. Rule a Appendix Bergmann’s along lizard (2014). a G. in Biology conservation conservation Moreno-Rueda and heat & ecology, S. ectotherm: evolution, Reguera integrating F.J., conservatism: Zamora-Camacho Niche in (2005). lessons C.H. phylogenies: Graham biology. from & evolution J.J. morphological Wiens of patterns together and Australian selection from rates sexual lability Estimating and limb opportunity (2009). Ecological J.J. Wiens (2012). O. terrestrial-vertebrate Seehausen Guinea radiation. New & adaptive Papua predict L.J. (2017). Harmon reptiles. S. C.E., except Meiri Wagner all & model for F. dynamic most Kraus a matters A.C., elevation Lakes: Algar richness: Great A., African Allison the O., of Tallowin fishes cichlid in radiation. empty? speciation adaptive half Explosive of or full (1998). half C. Cryptic glass Sturmbauer displacement: (2020). character Ecological S. Evolution (2013). Meiri & J.B. Ecology & Losos & S. Y.E. Carranza Stuart ( F., skinks Kraus Guinea New Evolution A., montane and Allison of Phylogenetics radiation O.J.S., non-adaptive size Tallowin and extinctions: K., diversity reptile Tamar Quaternary Late A., (2016). Slavenko are S. reptiles Meiri squamate & T.M., P. in dominates. Doan Raia evolution insularity Y., size matters, I., Itescu Roll body Das O.J.S., D., of Tallowin G.R., Pincheira-Donoso A., patterns Slavenko Global O.S.A., Colli Pauwels (2019). L., C., climate. S. Chirio by Nogueira Meiri driven M., Z.T., & not Y. Nagy Bohm Wang D., P., A.M., Wagner Meirte Bauer U., M., A., Martins Allison M., LeBreton A., Feldman Australian of A., origins Slavenko displacement. character the Ecological and phylogeny (1980). M. Lygosomine Slatkin (2011). M.N. limb Hutchinson of & lizards. evolution scincid A.F. the Hugall and A., reconstruction, Skinner state character competition. ancestral heterogeneity, and in Rate ratios morphology size (2010). A. reconsidered: Skinner Rosalia Santa escape (1981). morphology, W. 1206-1228. between 35, Boecklen relationship & The D. Simberloff (2004). H. Biology Nunez Evolutionary clade & of lizard the F.B. Journal in Cruz occupation microhabitat J.B., and behaviour Losos J.A., communities. II ecological Schulte in In: partitioning Resource mountains. tropical (1974). T.W. high Schoener in climate of features Ecological biogeography (1986). G. Sarmiento 7 2820-2828. 27, , nulRve fEooy vlto,adSystematics and Evolution, Ecology, of Review Annual es uleme oatroM.Ofr nvriyPesOfr,U,p.11-45. pp. UK, Oxford, Press University Oxford M). Monasterio & F Vuilleumier (eds. eut fa”rknsik nlssidctn hc C xli ttsial significant statistically a explain PCs which indicating analysis stick” ”broken a of Results Lerista ora fBiogeography of Journal l aaadcd eurdt u h nlsspcae sa project. R an as packaged analyses the run to required code and data All 8 402-408. 28, , ora fFs Biology Fish of Journal Sicde Squamata). (Scincidae, lblEooyadBiogeography and Ecology Global Nature 4,106749. 146, , Lerista lblEooyadBiogeography and Ecology Global 7 408-420. 17, , 8,366-369. 487, , lizards. 8 1044-1058. 38, , ora fBiology of Journal ytmtcBiology Systematic 3 18-36. 53, , 19 Ecology ora fBiogeography of Journal 8 471-483. 28, , Liolaemus 1 163-177. 61, , 6 519-539. 36, , 5 1308-1320. 25, , ,19. 8, , 9 723-740. 59, , Iunde rpdrne:Liolaemini). Tropidurinae*: (Iguanidae: Papuascincus Science 4 1734-1744. 44, , 8,27-39. 185, , ora fEvolutionary of Journal Scincidae). ; ihattd tropical altitude High Evolution Molecular rnsin Trends , Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. r o eaielaig,adbu oor r o oiielaig.Teheadtesz ftecircle are the direction) of either size (in the loadings and higher hue that The such loading, loadings. circles. the positive larger of and for value colours are deeper absolute by colours the represented blue with and correlated loadings, are negative for are S2. Figure aibelaig o aho h opoercmaueet ntetofis C.Rdcolours Red PCs. first two the on measurements morphometric the of each for loadings Variable 20 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. 21 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. hsforbesi phus stanleyanus S4. Figure ( highland for circles blue – species the ( of lowland distribution for elevational the on based S3. Figure es ao (B) lato; sensu opopc fPpaNwGie kns(aelaig sFgr ) ihidvdascoloured individuals with 2), Figure as loadings (same skinks Guinea New Papua of Morphospace xmlso oePpaNwGie knsadtervrosmrhlge.(A) morphologies. various their and skinks Guinea New Papua some of Examples . < 00m.Teelpe nops 5 fosrain o ahgroup. each for observations of 95% encompass ellipses The m). 1000 oi bumui Fojia (C) ; rblntsgracilis Tribolonotus 22 (D) ; iii longiceps Lipinia > 00m n eltriangles teal and m) 1000 (E) ; Papuascincus Sphenomor- Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. 23 Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. pce enDsac ihnRneMa itneOtieRange Outside Distance Mean Range Within Distance Mean novaeguineae Cryptoblepharus pulla Carlia mysi Carlia luctuosa Carlia eothen Carlia bicarinata Carlia aramia Carlia aenigma Carlia Species the and means the (highland: are species Written each range. column). of (right its PC2 outside and and column) within (left PC1 S1. for Table cell per kurtosis and skewness deviation, S5. Figure o ohdsac n acr iiaiy h agro h w aus(ihrwti rotietefocal the outside or hypothesis, within exclusion (either competitive the values grey. of two predictions shaded the the are fit of that larger Comparisons the i.e. bold. marked similarity, is Jaccard range) and species’ distance both For agrdsacswti oa pce’rne rsalrJcadsmlrt ihnfclseis ranges, species’ focal within similarity Jaccard smaller or ranges species’ focal within distances larger eut of Results ctepossoigrgesoso eneeaino tpt otm etod standard centroid, bottom) to (top of elevation mean on regressions showing Scatterplots t tsscmaigma itne rmadma acr vra ihec species each with overlap Jaccard mean and from distances mean comparing -tests > 00m lowland: m; 1000 4.43 3.9 3.404 5.16 3.78 3.42 3.51 3.82 < 00m.Significant m). 1000 24 p auso h -et,a ela h elevation the as well as t-tests, the of values .0 .90.17 0.171 0.89 0.03 4.24 3.67 3.401 4.96 3.93 3.13 3.53 3.86 p aus( values < .5 r akdbold. marked are 0.05) p < < 0.045 0.001 0.62 0.42 au o itneMa acr iiaiyWti ag enJcadSmlrt usd Range Outside Similarity Jaccard Mean Range Within Similarity Jaccard Mean Distance for value

.0 0.08 0.001 0.001

0.097 0.2 0.21 0.19

0.14 .30.001 0.07 0.16 0.1 0.23 0.2 0.17 .6 .7Lowland 0.37 0.18 0.168 .7Lowland < < < 0.47 0.02 < p au o acr iiaiyElevation Similarity Jaccard for value 0.001 0.001 0.001 0.001 Lowland Lowland Lowland Lowland Lowland Lowland Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. peoopu derooyae Sphenomorphus darlingtoni Sphenomorphus cinereus Sphenomorphus brunneus Sphenomorphus aruensis Sphenomorphus virens Prasinohaema semoni Prasinohaema prehensicauda Prasinohaema flavipes Prasinohaema stanleyanus Papuascincus morokanus Papuascincus macfarlani Lygisaurus curtus Lygisaurus subalpina Lobulia elegans Lobulia brongersmai Lobulia pulchra Lipinia noctua Lipinia longiceps Lipinia smaragdina Lamprolepis nigricaudis Glaphyromorphus crassicaudus Glaphyromorphus bumui Fojia rufescens Eugongylus veracunda Emoia tropidolepis Emoia submetallica Emoia pseudopallidiceps Emoia popei Emoia physicina Emoia mi physicae Emoia mi pallidiceps Emoia oribata Emoia mi obscura Emoia mi montana Emoia maxima Emoia loveridgei Emoia longicauda Emoia kordoana Range Emoia Outside Distance Mean jakati Emoia Range guttata Within Emoia Distance Mean caeruleocauda Emoia brongersmai Emoia battersbyi Emoia aurulenta Emoia atrocostata Emoia aenea Emoia spaldingi Ctenotus yulensis Cryptoblepharus virgatus Cryptoblepharus Species 4.09 3.2 3.31 4.36 3.51 3.33 3.09 3.46 2.77 2.79 2.78 3.72 2.74 2.46 3.42 3.86 4.23 3.8 5.01 3.91 3.74 9.84 5.63 5.01 2.9 4.23 2.96 3.33 3.45 3.34 3.92 2.97 3.95 2.85 4.09 3.9 2.66 2.88 3.25 2.97 3.93 2.98 4.05 3.98 3.03 2.9 3.03 3.73 4.34 4.78 25 3.92 3.74 3.71 5 3.08 3.24 3.04 4.18 0.14 3.11 3.07 3.17 3.32 2.9 2.72 3.26 0.01 3.67 4.07 3.58 5.23 3.82 3.47 9.64 5.88 3.79 2.83 4.36 2.83 3.08 3.32 3.22 3.82 .405 0.24 0.52 2.94 4.03 2.89 .403 0.16 0.38 4.15 3.61 2.71 2.65 2.96 2.99 3.84 2.8 4.06 3.9 3.22 2.75 3.33 3.36 4.23 4.47 .40.21 < < < < < < 0.04 < < < < < < < < < < < < < < < < < 0.004 0.02 < < < 0.01 0.38 .90.25 0.09 p .30.1 0.25 0.23 < 0.06 < < 0.45 < 0.97 0.02 < < < < 0.02 < au o itneMa acr iiaiyWti ag enJcadSmlrt usd Range Outside Similarity Jaccard Mean Range Within Similarity Jaccard Mean Distance for value

.0 0.061 0.14 0.11 0.07 0.164 0.001 0.001 0.14 0.001 0.14 0.001 0.18 0.001 0.21 0.001 0.17 0.001 0.22 0.001 0.001 0.001 0.06 0.001 0.001 0.02 0.001 0.001 0.001 0.001 1.68*10 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.213 0.001 0.001 0.001 .0 0.19 0.001 0.22 0.001 0.001 0.28 0.013 0.001 0.001 0.001 0.001 0.001 0.001

0.05

0.03 0.02 0.13 0.1 0.2 0.069 0.11 0.1 0.068 4.1*10 0.04 0.16 0.217 0.19 0.21

0.18 0.22

0.15 0.234 0.23 0.18 0.14 0.245 -5 -6 0.18 0.21 0.25 0.011 0.04 0.054 0.03 .40.04 0.001 0.024 0.226 0.24 0.26 0.18 0.15 0.15 0.25 . . Lowland 0.2 0.16 0.2 0.13 0.22 0.07 0.12 0.11 0.072 1.58*10 0.09 0.18 0.12 0.221 0.23 0.23 .6 .2Lowland 0.72 0.059 0.09 0.06 0.05 Lowland 0.163 0.06 0.1 0.17 0.19 0.15 0.19 0.29 0.05 0.01 2.15*10 0.21 0.2 .50.061 0.25 0.2 0.26 .20.001 0.12 0.19 -6 -4 0.03 < < < < Highland < < 0.03 < < Lowland < < < 0.68 Highland < < < < < < 0.7 < < < 0.31 < < < < < .6Lowland < 0.001 Lowland < 0.016 0.019 < < 0.06 < < 0.38 < p au o acr iiaiyElevation Similarity Jaccard for value 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 Highland Highland Highland Lowland Lowland Highland Highland Highland Highland Lowland Lowland Highland Highland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Highland Lowland Lowland Highland Lowland Lowland Highland Lowland Highland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Lowland Highland Posted on Authorea 18 Mar 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158456536.63719873 — This a preprint and has not been peer reviewed. Data may be preliminary. model. eneeain() h aaeesaetecnri,sadr eito,sens n utsso the of kurtosis and skewness deviation, coefficients, standard regression centroid, standardized the the are are parameters Listed The distributions. (m). elevation mean S2. Table novaeguineae Tribolonotus gracilis Tribolonotus gigas Tiliqua undulatus Sphenomorphus solomonis Sphenomorphus simus Sphenomorphus schultzei Sphenomorphus pratti Sphenomorphus papuae Sphenomorphus oligolepis Range Sphenomorphus Outside Distance Mean nigrolineatus Sphenomorphus Range nigriventris Within Sphenomorphus Distance Mean neuhaussi Sphenomorphus minutus Sphenomorphus microtympanus Sphenomorphus leptofasciatus Sphenomorphus latifasciatus Sphenomorphus jobiensis Sphenomorphus granulatus Sphenomorphus fragilis Sphenomorphus forbesi Sphenomorphus Species oe umre flna ersin fprmtr fdsrbtoso Csoe e elagainst cell per scores PC of distributions of parameters of regressions linear of summaries Model 0.03 R .100 .2P1Skewness PC1 Deviation Standard PC1 0.02 -0.01 0.01 0.07 0.1 0.01 0.004 0.39 0.02 0.06 0.08 2 < p < < < < < .0 .5P1Centroid PC1 0.05 0.001 .0 .9P1Kurtosis PC1 0.09 0.001 .0 01 C Centroid PC2 -0.19 0.001 .0 .5P2Sadr Deviation Standard PC2 0.05 0.001 .0 .2P2Skewness PC2 0.12 0.001 .0 .1P2Kurtosis PC2 0.51 0.001 7.32 7.83 7.1 3.68 5.42 4.27 4.17 5.47 3.04 5.64 5.09 6.15 6.64 4 7.61 3.43 6.41 3.68 2.53 7.11 6.23 tnadzdCecetModel Coefficient Standardized 26 p . .55.26*10 0.15 0.047 0.03 0.2 0.019 0.006 0.01 0.87 7.35 0.007 7.8 6.84 3.31 0.28 0.03 5.09 4.22 4.63 5.26 3.21 0.35 5.8 0.004 5.08 6.41 6.32 3.78 7.74 4.2 6.03 3.62 2.7 7.04 6.67 auso h ersin and regression, the of values

R 2 p 0.1 < < < < < < < < < < 0.003 < fthe of au o itneMa acr iiaiyWti ag enJcadSmlrt usd Range Outside Similarity Jaccard Mean Range Within Similarity Jaccard Mean Distance for value .0 0.08 0.001 0.14 0.001 0.001 0.001 0.001 0.001 0.14 0.001 0.26 0.001 0.001 0.001 0.001

0.039 0.01 0.05 2.38*10

0.02 4.51*10 0.1 0.03 0.13 0.039 0.01 -4 -3 -3 .0 .5Lowland 0.15 0.05 0.12 0.07 0.06 0.04 0.24 0.009 .4 .8Lowland Highland 0.48 0.36 0.12 0.04 0.046 0.05 0.018 0.02 0.007 2.44*10 0.01 6.91*10 0.01 0.01 0.13 0.06 -4 -3 .3Lowland Lowland < 0.53 < < Lowland < < Lowland < Lowland 0.7 < < < < 0.05 < 0.33 0.11 < 0.01 p au o acr iiaiyElevation Similarity Jaccard for value 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 0.001 Lowland Lowland Lowland Lowland Lowland Highland Lowland Lowland Highland Lowland Highland Lowland Lowland