Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. o aeadsicinbtenoencvlai n otnna nua ytm n updte ntheir in did them or islands, lumped oceanic and volcanic systems on insular focus continental however and did 2016, their these volcanic of left al. oceanic et All have between Weigelt 2019). changes 2014, distinction al. or sea-level a al. et make Late-Quaternary et merge, Norder Rijsdijk not 2019, and 2014, 2017). dominate. 2018, Aitchison fragment al. and processes al. expand, et (Ali climatic et patterns Avila and controlled diversity scales Simaiakis shrink species is time insular ontogeny to (e.g. on shorter island’s imprint islands emerge on an causing and while (Ma), fluctuations disappear 2008), scales sea-level even distribution time major al. deep the drives et At influencing Climate (Whittaker host. settings processes they geographical geological organisms evolving by of continuously evolution with genetic entities and dynamic are Islands historical of perspective Introduction the from biota studying oceanic while from them fundamentally differ between study archipelagos distinguishing continental Our of patterns, biogeography. richness importance endemism. species the promoting LGM affecting lasting highlighting processes processes shorter systems, counteracting the of diversity, mainland terms species oceanic in the current Unlike that the with on shows islands. connectivity effect same ample increased those has promoted on configuration endemics median configurations of lasting a we underrepresentation longer species islands to the stronger endemic land-bridge where modestly and and and on islands, Particularly native species only patterns, native both mainland. diversity scenario of the affected plant and supersaturation sea-level configuration islands detected geographic Aegean median land-bridge LGM between shaped the connections The establishing largely while patterns. through changes diversity richness LGM, Paleogeographical species the centipede confirm since results degree configuration. Relationships Our lesser dramatically present Species-Area goodness-of-fit. changed the Island of the from has metrics analysing multiple investigated differs geography by on We based Aegean’s groups selection islands). chorological the Models (56 on Mixed that centipedes Linear settings Maximum Generalized and geographical Glacial using islands) Late and past 3) (ISARs) (70 and cycles; in angiosperms Glacial-Interglacial present-day patterns aim nine for of richness last we occupancy the impact species study over taxon-island shaping published median this using compiled 2) in geography We current; Through area’s fluctuations 1) the (LGM). sea-level reconstructed stands: first systems. Quaternary sea-level we three archipelago, late oceanic Aegean under of the data, volcanic on role shaping Focusing on in the systems. investigating fluctuations focused island shelf sea-level by continental largely Quaternary work but late this of systems, extend role to insular important in the highlighted patterns has biogeographical biogeography island in research Recent Abstract 2020 28, April 5 4 3 2 1 Simaiakis Greece) Hammoud Sea, Cyril (Aegean archipelago continental-shelf species a insular of structure diversity stands sea-level short-lasting and Extreme ´ nvriaed Lisboa de Crete Universidade - Museum History Natural Amsterdam van Universiteit Athens of University University Ghent 4 iteNorder Sietze , 1 osatnsKougioumoutzis Konstantinos , 5 n .EilVnLoon Van Emiel E. and , 1 2 ent .Rijsdijk F. Kenneth , 3 3 Stylianos , Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. Smikse l 07.Drn h G 2. 9k P,tegoa e ee a 15mlwrta present than lower m negligible ˜135 was mostly level is sea scale global time the BP), same ka the 19 at – (26.5 occurring locally LGM changes the the and during tectonic During oscillations the 2017). sea-level of al. the Galanidou, et importance by and (Simaiakis the caused (Sakellariou BP, changes fluctuations geographical Ka climatic the 150 of Pleistocene its last magnitude the archipelago, the Aegean by 2009). to the affected (Higgins Compared of Arc mainly 2016). paleo-evolution Volcanic been the Aegean has shaped South history strongly therefore the have recent has of tectonics and formation Eurasian) plate the and though in African even However, resulting (Anatolian, plates systems, tectonic faults three major of convergence developed the at is Aegean The 1). (Figure cycle archipelago glacial-interglacial Aegean last The the endemism during different level of sea richness median Method the species 2) on and islands To LGM, land-bridge occur. during to of 1) state, speciation effects extreme for H the needed LGM investigate We analyse time lasting levels. we the short provide hypotheses, the not than our do configuration rather test configurations glacial patterns, (LGM) spatial median richness Maximum short-lasting long-lasting species that Glacial the assuming the Late islands, influenced land-bridge short-lasting largely on and have Thus largely extreme may 2019). could the al. configuration et than archipelago (Norder patterns median third configuration the richness Our islands, endemism continents. oceanic the the On explain from patterns. richness gene-flow affecting repetitive islands drop by land-bridge level suppressed (H on is species (H hypothesis groups, speciation native hypothesis second taxonomic allopatric of well-studied as rates Our well establishing islands, two 2017). as higher bridge for current al. the archipelago of reflecting et (H diversity islands, Aegean (Simaiakis hypothesis true species the on first insular for than Our hypotheses. continental islands connectivity our centipedes. on test and impact and to area combined system angiosperms al. study island the et ideal investigate past (Simaiakis an to Sea as represents Aegean is setting the this aim in studied, 2012). Our islands well 2008, are of al. change biota et Island paleogeographic its Triantis land- and the and (e.g. continental models quantified 2017) assessed multivariate of recently quantitatively in have influence been reflected we the never as Since Moreover, has correctly chorotypes (ISARs) 2019). processes different Relationships biogeographic in al. by relevant patterns while Species-Area et driven the biodiversity archipelago; Norder interpret on connectivity Aegean to 2014, islands and the crucial bridge Tiniakou for fragmentation is and investigated analysis change, Kougioumoutzis combined been archipelago area the a never (see of of such has part that diversity effect limited and think species combined a native we or insular between The group on distinguishing 2010). species not changes single by al. sea-level a i.e., et considering limited interest: by Panitsa of remain 2020), factor studies (e.g. al. explanatory and most et single (Itescu insights, heterogeneity a species biogeographical geological on endemic crucial high focusing intensively providing by studied a Although scope been with 2018). in has al. biogeography archipelago on 2016). its et shelf Unsurprisingly, archipelagos Triantis (Strid change. continental studies (e.g. largest sea-level biogeographical typical to the for due contribute a stage of land-bridges heterogeneity ideal form one numerous environmental an islands high is Mediterranean rendering and thus Aegean the and endemism, history in The peninsulas, and geological archipelago biodiversity Anatolian complex high Aegean Its the its The 2010). to and 2020). al. Kreft Greek al. et the et extensively and (Blondel studied Itescu between Earth (Weigelt been 2008, located islands have al. biota is continental Another et island Sea lower. continental surrounding Cardillo were on 1972, often levels dynamics Diamond sea sea-level mainland (e.g. of when the effects past geo-spatial of the The in proximity 2013). mainland the the is to connected characteristic were that islands, et archipelagos, “land-bridge” Whittaker continental ( 2017, of islands al. shelf specifics et Continental the Simaiakis disregarding 2017, By (Ali overlooked 2019). 2017). potentially al. al. are et processes Veron biogeographical 2016, relevant al. et (Weigelt analyses > 3 4 oe nteMdtraen(abc n ucl 05 lr ta.20,Lmeket Lambeck 2009, al. et Clark 2005, Purcell and (Lambeck Mediterranean the in lower m 140 ocrsteeeto h uaino h rhplgccngrtosa euto sea- of result a as configurations archipelagic the of duration the of effect the concerns ) 3 yaayigseisrcns atrsi eaint w aegorpia settings: paleogeographical two to relation in patterns richness species analysing by sensu l 07 r hrceie ystigo otnna hl n a include may and shelf continental a on sitting by characterized are 2017) Ali 2 sta neimi eaieyiflecdb land- by influenced negatively is endemism that is ) 2 1 sta oentv pce cu nland-bridge on occur species native more that is ) Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. ubro te aaees h R compared We the model. parameters, power fitted (1921) of Arrhenius adjusted-R the number of using transformation models logarithmic the the using fitted were ISARs islands” land-bridge analysis (“median Inferential level sea remained 2017). median that al. at islands” those et or of “true islands”) Simaiakis groups: land-bridge as - effects different (“LGM two to LGM biogeographical the into referred during the islands mainland, mainland classified investigate the the Supplementary we To – besides from mainland, (2017 Hence, configurations. isolated al. the difference 2019). the et paleogeographical to from al. under Simaiakis the fragmentation et for by order situation work in past Norder 1) in the the - interested used mainland (MSL Appendix and we were cycles islands, LGM the Material we Aegean glacial-interglacial the the to nine Herein, of during last distances diversity. configuration the situation present-day and over species the level areas current setting, sea insular of median geographical present-day past predictors the the as between estimated metrics we these hypotheses, use our 2005). test 2004, assessment. To al. statistical et for a set Simaiakis for data (see small The morphometry too respectively). centipedes Paleo-island was species, 70 5 centipedes and and of islands (65 endemics ‘endemics’ Aegean island and single 56 ‘native’ distinguished: for were compiled chorotypes Two was matrix endemics similar multiple chorotypes similar A (MIEs) combined in endemics’ islands also trends ‘multiple We assess form to taxa). to stricto endemics and (292 Aegean sensu non-endemics and islands phyto-region with Aegean both endemics combined Aegean the We of chorotypes. same of trends taxa). the one general (91 within compare on mainland located to exclusively the islands occurring on multiple not endemics or islands but are one Aegean islands to the multiple restricted region and on are phytogeographical mainland found (PE) Greek are endemics’ the (AE) ’phyto-region on endemics’ The the both ‘Aegean on occurring The and exclusively Taxa). non-endemics’, Islands species (689 Greek ‘native are the The endemics’ on ‘Greek 2017). both The occur al. taxa). taxa that (2673 et data group database plant Mediterranean Kougioumoutzis taxonomic compiled bibliographical the vascular spread in we extensive across widely native S1 an mainland and a 3246 on are Appendix Aegean, ‘natives’, based of (see the termed 1) total archipelago in here (Figure group Aegean a chorotypes well-studied combined the including two a for islands, and are chorotypes Aegean angiosperms five 70 The on for subspecies). matrix and plant (species a compiled We been have 2016, Tan regions data and (Strid richness phytogeographical (Strid Species Aegean influences different 2017). the historical al. affinities, divergent in et and biogeographical rate Kougioumoutzis conditions environmental endemism 1997, on similar and Based from diversity resulting 2018). their identified al. for known et are Triantis centipedes and Angiosperms and 1 (Figures group the islands and organisms Cyclades increase and Target peninsula Turkey the rapid of Peloponnese into a coast fragmented the the to paleo-island between near and Cycladic connection emerged the basin 2). Islands western and Aegean 2017). the lost al. the was peninsulas, et in mainland Simaiakis formerly 70% - were respectively ca. that Kya, by Greece 1 marine area 2017). per the al. island 40% ( et of and total islands (Simaiakis most (20% of Ka Large with m/1 reduction islands. 12 basin Aegean major were Aegean of rates a the rise to sea-level in led when changes BP, This Ka dramatic 16-11 caused during observed rise transgression sea-level Subsequent 2014). al. 40tx) eas obndaledmccooye noa aledmc ru”(E group” “all-endemics an into chorotypes endemic all combined also We taxa). (450 sensu 2 ausa esr ftergons-ffi.A h oeshv h same the have models the As goodness-of-fit. their of measure a as values ti n a 19 8 aa.Fnly h igeiln neis(SIEs), endemics island single the Finally, taxa). 384 - (1997 Tan and Strid > 0km 20 2 r ietycmaal,wtotmdfiain(ratse l 2005). al. et (Triantis modification without comparable, directly are 2 eerpdysrnigadbcmn sltddrn httime that during isolated becoming and shrinking rapidly were ) 3 versus hs iha psdccneto othe to connection episodic an with those ALL ). Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. ;Fgr ) S rals a nisgicn eaieeeto I ihespten Tbe2 iue4). Figure 2; (Table patterns (Table patterns richness SIEs richness SIE Species the on and effect endemics negative emerged insignificant Greek variables an the LGM-related had Discussion for 4). loss except area Figure Area chorotypes, MSL 2; 2). (Table 4). angiosperm (Table centipedes Figure models all and 2; overall all for angiosperms was almost predictors native and in the important 2) predictor for as for Table not important variance but - of most 2), proportion (16-83% the Table large was models - a (13-67% all explained categories affinity for endemic Biogeographical high all centipedes. generally than was angiosperms for explained greater variance trend of opposite proportion the The while islands, land- true endemics on than phytogeographical than 3). taxa and Models islands (Figure native Greek taxa true more the endemic on host from regarding variance to apart observed appear of trend, is islands proportion similar Land-bridge a higher 1). followed a (Table z-curve explained the model islands; LGM function bridge power the ISAR in richness The species minimal A2.2). of was Table at analyses the 2 isolation inferential during are Appendix while and mainland Material and the Explorative (Supplementary MSL, to LGM LGM connected the the the morphometry were at to islands during Islands’ Fewer compared values size A2.1). LGM). MSL Table intermediate maximum 2 the with a Appendix represent during Material now, setting achieved (Supplementary lowest LGM areas size now, the Island minimal and (highest correspondingly: their configuration fluctuations varied current sea-level The isolation the 2). and (Figure of the Quaternary throughout extremes the dramatically two changed of archipelago cycle Aegean the climatic of last geography the that show reconstructions Our diversity. centipede Aegean in native the over-dispersion to in fauna their changes centipede Geographical and the species of centipede analysis an endemic as our it restrained of Results designated we number systematically Crete, low leverage to the vs. adjacent residuals of islets of the Because plots shown). residual (not diagnostic as outlier fitted models all (BIC), from Criterion Information pseudo-R of Bayesian (loocv) measures (AICc), multiple Criterion using potential (R Information explanatory conditional Akaike most VIF corrected provided with the that variables models removing goodness-of-fit: the and function identified predictors computing with we the by log-link Finally, variables of addressed 2013). was (VIF) All a Multicollinearity factors 2017). variables. using inflation remaining al. variance the fitted the the et with variables: Kuznetsova refitting were package, before response models 2014) (lmerTest (Murtaugh following term All error or the angiosperm richness. total the island on richness, species p-values for land-bridge species predictors distribution centipede a angiosperm present-day of Poisson was total total the sets the and island and to angiosperms, three an of compared richness whether chorotypes these geography six endemics and fitted all insular mainland We of in the richness island. change species to true of distance metrics a of predictors following same remained increase the the loss, included as plus median) area province present-day or model setting: phytogeographical (LGM of and null settings consisted effect paleogeographical the fixed model) Aegean the as as for (null the models autocorrelation models model of mixed spatial The present-day setting effect. and linear The geographical random mainland, the MSL. the generalized to and to alternative distance LGM correspond area, three present-day, models changes stages: of The geographical three richness. performance past at species the of explaining and impact compared in chorotypes the we centipede (GLMMs) investigate and diversity, To angiosperm islands. current all land-bridge on for and richness true species between area-adjusted distinguished mean the calculated also We > . eeecue rmtemdl ooti e f“ugsie u nocuie predictors inconclusive” but “suggestive, of set a obtain to models the from excluded were 0.1 2 )admria (R marginal and c) 2 l nlsswr efre nRvrin342( oeta 08.Ceewsexcluded was Crete 2018). team core (R 3.4.2 version R in performed were analyses All . 2 )R m) 2 MMnpcae atn20)adlaeoeotcross-validated leave-one-out and 2009) Barton package, (MuMIn 4 > . Drane al. et (Dormann 2.5 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. yldsadKii(asmlse l 09 aelro n aaio 06.Pesoeesalvloscillations sea-level Pleistocene 2016). Galanidou 1966, and Sakellariou (Creutzburg true 2009, Sch years al. (Pleistocene) 1990, et the of (Dermitzakis (Kapsimalis Kriti between Rodos millions and and existed Cyclades high over paleo-islands Karpathos paleo-connections as Those processes Crete, was pre-Pleistocene islands. Kythira, geological level islands true instance, by sea were For Quaternary the islands 1990). when the land-bridge Dermitzakis paleogeography, these than the and indicate earlier of may interglacials imprint fragmented This Quaternary pre-Tertiary 1972). the a Strid during larger have 1971, on as may Snogerup and distribution MIEs 1971, Rechinger larger Greuter much Aegean 1943, a 1971, (Rechinger that typically continental Quaternary 1969, had in the Runemark MIEs see beyond 1951, we Aegean reaching Rechinger-Moser what result, even to a sometimes became compared that As paleo-islands, settings 2018) 2017). pre-existing islands island al. al. oceanic oceanic et et by in shared (Simaiakis (Norder in paleo-islands islands settings oscillations contrast of of sea-level numbers The number lower Pleistocene much low MIEs. by by the fragmented explained oceanic and simply MIEs on be continental than can by MIEs shared rather islands continental Ka of since on numbers 20-11 area-loss high prominent island during dramatic more oscillations the sea-level seems from apart to Island-fragmentation patterns, due ( MIE fragmentation continental LGM observed fragmentation, paleo-islands the the Quaternary rapid for 2017). and responsible al. be repeated and et may richness the Kougioumoutzis distribution, as 2015, species’ well on al. imprint as et the its Poulakakis left that (e.g., has suggesting archipelago patterns SIEs), Aegean evolutionary (except the 2; diversity of (Figure endemic configuration LGM affected spatial the negatively LGM during has achieved 2016) connectivity Galanidou short-lasting yet and maximum, Sakellariou patterns The richness 4). species (Figure island richness continental species shaped have to seem not (H does configuration MSL long-lasting The (Whittaker fragmentation numbers island and species configurations and Pleistocene continental levels Intermediate thus endemism on gene re-established, lower underrepresentation enabled repeatedly to endemic cycles, populations leading Fern´andez-Palacios of glacial-interglacial 2007). source consequently and (Appen- state continental the and islands and a during speciation true insular to mainland hampering the isolated pointing the for previously 2008), with steeper between al. reconnections flow is et frequent slope on Triantis The ISAR reflected 0.8; islands. the pattern = as a (z - 2017), islands SIEs H al. experien- for confirming currently et pronounced 3), are (Simaiakis dix more biota equilibrium R being These (low reached – well 2018). not chorotypes as al. have ISARs et and endemic Panitsa periods, period their low-stand (e.g., from relaxation sea-level taxa Aegean result glacial a could native islands. Quaternary This cing of land-bridge the implying dropped. are influx during thus levels similar prevailing sea which continuous islands, after is states true 2010), promoting taxa, connectivity and than al. native geospatial 2012) endemics in et high fewer al. supersaturated Panitsa angiosperm the and probably et 0.326; overall were taxa (Triantis = former The native settings the (z more 3). shelf that islands significantly Appendix continental host Aegean for 1; islands East (Table range land-bridge the z-values the of ISAR within (e.g. corro- that in falls results regions to reflected 0.329) Kreft Our phytogeographical also = 82–95%). with island is (z 2019: Greek line z-value pattern al. several in This et for Valli are 3). 2014, reported (Figure results Tiniakou values our and with H and Kougioumoutzis the borate 2011, 2) as al. (Table well et capacity as Kagiampaki to predictive (2008), aimed not high al. have study had et which Our model far. islands, 2019). so continental power-function questions al. on Our these observed et to be (Norder regard might with patterns intermediate patterns investigated the diversity different islands been that MIE oceanic on possibility and that the seems SIE investigate It 2019). shapes al. Pleistocene configuration et the the (Norder sea-level of Recently, richness representative al. species 2015). more endemic et those al. (Fern´andez-Palacios Weigelt explain or et (e.g. configurations better expectations paleogeographical patterns theoretical LGM richness following whether species arose 2019), island question al. on mark et their Norder left 2016, have changes sea-level Quaternary Late 3 .LMcngrto nteohrhn,eegda ao rdco o ohntv n endemic and native both for predictor major a as emerged hand, other the on configuration LGM ). 1 n H and > 0) hspoesaet sadseisrcns repcieo ra i asv dispersal. passive via area, of irrespective richness species island affects process This 70%). 2 adbig sad otmr aiennedmctx,ytsgicnl ee endemics fewer significantly yet taxa, non-endemic native more host islands land-bridge : 2 eetees h IsIA lp ssgicnl oe hnrpre rmoceanic from reported than lower significantly is slope ISAR SIEs the Nevertheless, . 2 aus–Tbe1.Teopst rn sosre o l h endemic the all for observed is trend opposite The 1). Table – values 5 l 93 rbtentesouth-central the between or 1993) ule ¨ Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. nteAga rhplg.Oeo h otecpinlfaue fcrancnieeseis particularly (e.g. species, tidemarks centipedes centipede between the certain shore of of sea patterns features the diversity exceptional on the most reside justify in the they could (Voigtl most median) of that This barks than and One is archipelago. and insensitive (LGM Aegean archipelago. geophilomorphs, stones more paleogeography the Aegean under be in of the as to rise role in well seem sea-level significant as species by less layers centipede caused in the soil changes most litter geographical life-style deeper leaf to cryptozoic the animals as angiosperms soil their in such islands, predatory of cavities microhabitats true are to Because into on centipedes preferences layer, 2011). All types ecological soil size. diversity. habitat show island upper centipede cryptozoic that with the native suitable abilities increases of islands dispersal of these levels distinguished reflection on with Kya higher a occurring 21 support likely of than islands change most the less land-bridge is isolated to trend counterintuitive were compared our This islands islands confirms the true islands land-bridge of on on values majority centipedes endemic the of west, and absence Anatolian east H the the The the the of 2003). from or to Conispoliatis paleogeography Anatolia Greece Greece and the from continental (Perissoratis mainland Mya with with 9 from fitting either approximately Mya isolation isolated connected 5.5 permanently recently island became were of Crete islands degrees Although the low peninsula. of by most explained where probably Aegean, is 2008), al. ( et z-value low high remarkably isolation, The long size, large its 1975, diversity to 1972, Centipede due (Greuter the M´edail is history explain 2017). 2011, Crete paleogeographical also Stein- its al. in may 2013, as et occurring islands well al. Kagiampaki these SIEs as et heterogeneity, of of Trigas environmental geodiversity number and 2010, and large topographical al. complexity The topogra- formation et topographic pattern. and SIE The Kallimanis SIE mountainous in 2016). (e.g. Samos ISAR role 2013, are speciation LGM; al. important islands driven since et additional have three islands bauer an might These plays land-bridge isolation levels. topography were ecological sea that all and median suggesting which during pronounced, Samothraki, most also and phically island Samos land-bridge Evvia, a com- SIEs: was dominant 10 the (e.g., than to even isolation more inaccessible systematic virtually and were (e.g., differentiation cliffs LGM, regions These mainland the adjacent lower. during the m subsect. pronounced in 140 pre- most communities cliff were to random occupy of that up endemics by ponents mountaintops were Aegean driven of took levels Several remnants mainly differentiation islands) sea 1978). are species (18 when is which Cotandriopoulos active them Aegean cliffs, and and of limestone the therein) (Cardona most cipitous references islands in islands, and numerous other speciation 2010 inside refe- 25 and Delipetrou and place on enabled and diversification 2018 then This (Georghiou al. Plant connectivity. and drift et maximum Crete SIE. genetic Panitsa on of continental one (e.g. occur result on levels only SIEs Meanwhile, a endemism containing Aegean 2019). and as Most al. speciation LGM lower therein). et the to rences endemism (Norder during leading oceanic and size achieved thus speciation island to was gene-flow, island large contrary increased size Oceanic long-lasting, 4), island 2019). MSL al. Figure large the et islands 2; to (Norder (Table due configurations configurations elevated MSL by are past driven by are unaffected which that SIEs are data included. however, species species of SIEs MIEs the subsets Continental of or with past history working SIEs and evolutionary of short into common are importance endemics islands the the between geological highlights of reflect distances different This classification regionally dispersal frequent. by where usual were fragmented and both land-connections the oscillations is and sea-level reasons, that Bittkau Pleistocene system these 1971, continental-shelf and For a events 1969, probably in 2018). Runemark MIEs sense (e.g. old little al. make speciation the et would allopatric of Jaros through some 2009, SIEs islands, neo-endemic island Comes complex rise single mostly topographically sea-level were to In once Pleistocene rise perhaps islands. and gave that pre-existing islands paleo-endemics larger of of fragmentation on up the endemics breaking Cyclades to the pre-Pleistocene led on fragmentation geological instance island Both for induced as 2017). more, al. even et populations fragmented (Simaiakis earlier geologically the fragmented further 2 aeyta neimi eoe nln-rdeilns o aiecniee,hge -ausadR and z-values higher centipedes, native For islands. land-bridge on demoted is endemism that namely , Quinqueloculares n evda eui uigteQaenr lcain,laigt ihgenetic high to leading glaciations, Quaternary the during refugia as served and ) < .8 o l etpds vnlwrta eotdfrvrerts(Triantis vertebrates for reported than lower even centipedes, all for 0.18) iu arboreum Linum 6 hmo 05.Ol he sad host islands three Only 2005). Thomson – nl verbascifolia Inula ahmru ferrugineum Pachymerium ru and group Campanula ¨ ander 2 , Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. h urudn otnna adms.Ti onciiywt h anad vntog short-lasting, connectivity and though extreme islands even of land-bridge mainland, result islands, the the true with as Aegean connectivity this expanded This the explain mass. continental- We between land LGM a LGM configuration. continental by centipedes in the same surrounding extent, explained that lesser during the the best however, a achieved of to are show was and patterns impact results species islands that richness an angiosperm Our oceanic endemic non-endemic shows on 2019). too native angiosperm biota been of al. still insular Aegean have richness et the of may have the and Norder composition episodes to of configurations, and these 2013, estimated most diversity of BP al. archipelago, the duration Ka et on shelf the 800 Heaney impact Therefore, last significant 2019). 1989, the a al. (Porter of had et 2% have and (Norder only to sporadically level only short with sea represented Pleistocene, was low that the similarly configuration throughout a extreme an time-periods is short LGM for the of setting geographical Comes The and Bittkau the archipelagos 2005, oceanic of Comes and differentiation and Continental-shelf the (Bittkau is Pliocene/Pleistocene the this divergence during for 2009). genetic events example cumulative vicariant to prominent several fragmentation interval leading of One glacial-interglacial of state, 2009). every connected character arvensis during al. glacially repetitive Nigella existed lasting (Aguil´ee et the state longer populations considering disconnected the geographic between and when disrupting the fragmented and ka Especially of the 16 ˜20 2017). My, between most for years 2 al. since 5000 last et within time-period, the Aegean (Simaiakis over short the BP in a Ka transgression in marine 11 (e.g. events during occurred occurred island-splitting change radiations of configuration non-adaptive sequence several al. labyrinthine where et of and diversification Aegean Jaros through recent central 2008, the SIEs al. the supported et neo-endemic in Campanula have Comes formed especially may 2009, MIEs oscillations species, 2005, sea-level old neo-endemic Comes through the and isolation Bittkau of Geographical 1971, some 2018). 1969, islands, (Runemark (Runemark complex (i.e., archipelago speciation southern Aegean topographically allopatric b) the 1996a, in In Christodoulakis persist 1971). Ikaria: eastern to (e.g., In 1969, Other flora 2014), eastern islands. relict Tiniakou 2018). and Aegean a and 2011) Rodos) al. permitted al. Karpathos, (Kougioumoutzis probably et et Crete, central (Kagiampaki most (Panitsa the southern conditions and Evvia for climatic 2010) favourable diversity case occurring Tzanoudakis and addition, endemic and the Crete taxa Panitsa current actually 2010, shaping in endemic al. is in et occur role the as (Panitsa important which Aegean, of an the of play ˜45% in probably majority geodiversity nearly history patterns vast and communities: paleogeographical climate the as plant SIEs, complex such island factors are region’s consequently that Aegean and Aegean the stating species in the narrowly-ranged species studies of of turnover in proportion endemic previous result species high in the the with high a due all line occurring also of is primarily for in taxa This variance 2017) is is plant al. of et This bioregionalization less native (Kougioumoutzis affinity. proportion Aegean, the the biogeographical largest (Greuter the justifies of to the times in attributed hand, clearly patterns historic is other diversity species early metrics the the distance-insensitive or richness On in prehistoric artificially archipelago. flora affinity in or Aegean Aegean action present biogeographical the naturally the human of of these of (˜40%) role result of portion significant large a presence a variance as that the The fact islands of the 1979). fraction Aegean to small the attributed a be reached only can explained has This models) taxa. than our native in smaller the effect for times random Crete. the in 800 (i.e., found affinity about richness Biogeographical is the half example roughly for – Kastelorizo centipedes affinity Biogeographical result of LGM. a be species the as to 19 during centipedes tend hosts species from Turkey with centipedes but distance Centipede supersaturated to that Crete and are equilibrium. meaning connection Stroggyli area richness, new their and species with a Ro of total 1), Kastelorizo, to for (Table However, variables faster relaxed. geography significant faunistically present-day leading most by the taxa, determined being these mainland mainly in therefore extinctions is fast richness caused have might poseidonis Tuoba , Nigella pce ope hc sdet o-dpierdainadrno eei rf saresult a as drift genetic random and radiation non-adaptive to due is which complex species e ei 91.Tu,proso ihrtso e-ee iesc sfloigteLGM the following as such rise sea-level of rates high of periods Thus, 1981). Lewis see , , Erysimum ..Cmse l 08 ao ta.21) sarsl faturbulent a of result a as 2018), al. et Jaros 2008, al. et Comes e.g. – 7 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. l eeatGSfie,seismtie n cit ilb aeaalbeuigDyddt repository. data Dryad using available made be will scripts R and matrixes species (ref. files, grant GIS relevant research All FWO a Statement Fellowship by Accessibility Research funded the Data and Funda¸c˜ao partially UID/BIA/00329/2013 through Funds, project was National the Portuguese CH within the (FCT), PD/BD/114380/2016. from Tecnologia funding a interest. Ciˆencia received e a of SJN para work. conflicts this no during 11C5219N) declare authors models. The statistical and in problematic differences fundamentally their islands assess- incorporate continental Declarations explicitly statistical and should in make oceanic systems differences combined species islands both principal of continental the involving true noted data on and studies were The species paleogeography oceanic endemic still islands. of between of islands Aegean effect mechanisms ments the land-bridge the evolutionary of of these and patterns may evidence when geographic which biogeographical ample distance, conditions, and hold day speciation pre-Quaternary therefore present richness, the on endemic systems multiple to dependency Continental some a response for islands. and noted most relict islands We the a land-bridge Pleistocene. to species, on reflect the tuned z-values endemic of supersaturation high end island native the the continental and single chorotypes, at more prevalent underrepresentation in ever configurations endemic to extinctions (LGM) of species extreme induced signals continental lower be- to of increasing processes, waves islands that leading evolutionary migration hypothesise isolated ultimately We to allopatric when led 2005). disrupting sorts, period Geer of ( islands Quaternary der Van the “shocks” time speciation and over as and (Sondaar levels of richness the manifested mainland sea species period with the mainland their with connectivity long the affected connected greatly high with very came Quaternary of connections a the paleo-configurations These of for recent onset patterns. the mainland and with extreme the started the that from mainland endemism; separated promoting were thus islands Ma) 2010). continental is al. Aegean et endemism higher The Panitsa of island result (e.g. Single the islands then continental prevalent. is the it as by Conclusions manifest, offered seen supersaturation does geodiversity the where are endemism or is islands when debt topography continental space) and in extinction to variability islands, niche greater continental contrast vacant on to in (through supressed islands, due speciation low-stands. generally oceanic disequilibria ecological marine on species and during richness far isolation mainland and species is nearby intensely of of fragmentation, more the result driver of influenced with degree a main have difference the might connectivity as crucial changes as increased speciation sea-level a well through recent Moreover, is as area the mediated there loss, such, island biota area conclude as which of their can and at rate We islands thresholds continental the on 2019). sea-level in islands: depends greater the continental al. this influencing and et Clearly study, oceanic (Norder LGM. under between significantly the basin change than the maximum connectedness longer of far most and lasted depth how for and and whether in earlier or tested geometry stand, reached be the Native sea-level was to short-lasting connectivity lowest remains endemism. the maximum and It to continental biota on biogeography. related distinguish that historical exclusively signature that noted are of the distinct processes conditions we perspective connectivity essential Lastly, by a the thus from shaped level. left are island sea largely fragmentation endemism oceanic high reduced indeed island present and is geological the supersaturation today Quaternary by species Aegean islands pre smaller the Pleistocene time into by in deep landmasses fragmentation manifest on large island endemism natives of ( Aegean angiosperm fragmentation of islands of oversaturation more island degree much to multiple high sharing point The MIEs the islands to that islands. richness led land-bridge continental observed endemic oscillations on on also sea-level the natives endemism We hand, reduced of other to islands. richness the pointing land-bridge species islands, On mainland adjusted oceanic settings. on the mean island those to The continental than The connected lower for biota. much range once island are z-value were z-values Aegean the the that into in islands fall levels on richness endemism lower supersaturation of native conditions of and conditions both promoted 8 > )ta sosre o cai archipelagos. oceanic for observed is than 5) Ι A -ausfrttlangiosperm total for z-values SAR > 2.6 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. eiwo ehd oda ihi n iuainsuyeautn hi efrac.–Eorpy36: Ecography – performance. their evaluating study simulation a Collinearity: and 2013. it S. Lautenbach, with Carr´e, G., deal G., Southwest Carl, to 27-46. of C., methods Buchmann, of Avifaunas S., review Bacher, for A J., Elith, Times F., 3-16. 3203. Relaxation C. 20: – Dormann, of Balcanica 3199 Geolog. Estimation 69: P.N.A.S. – Kinetics: – Cenozoic. Islands. Late Biogeographic Pacific the 1972. during Aegeis J. the Diamond, of evolution The by 1990. D. exemplified Dermitzakis, as floras island s Die continental 1966. in N. Creutzburg, speciation Plant McCabe, 2008. S., C. Hostetler, Bittkau, J., A., Nigella 710-714. Tribsch, Mitrovica, 325: P., B., Science H. Wohlfarth, – Comes, J., Maximum. 63-91. Clark, Glacial 36: A., Last Phyton Carlson, The – J., 2009. islands). Shakun M. (Greece Aegean, A., Ikaria Dyke, of (E. P., flora Ikaria Clark, Aegean 399. of E. – flora The 393 1996. the 191: D. Flora of Christodoulakis, – patterns Mediterranean. distribution E. the phytogeographical within The Greece) M´editerran´eennes. 1996. insulaires D. – flores Christodoulakis, les L’end´emisme dans 49–77. 1978, : J. 2 Cotandriopoulos, mammal Mediterranea island and of M.A. structure 1549–1556. phylogenetic Cardona, 275: the in Sci. patterns Biol. Global in Proc. 2008. Diversity – A. Biological assemblages. Purvis, Region. J., Gittleman, Mediterranean Inc. M., The Press Cardillo, Univ. 2010. 36: Oxford G. Biogeogr., – Boeuf, J. Time. in J.-Y,. – and Bodiou, shift Space archipelago. J., diversification Aegean Aronson, Pleistocene J., the Late Blondel, in a speciation of increased inference from Molecular resulting 1346-1360. 2009. (Ranunculaceae) phylo- P. lat. Molecular H. s. system: Comes, island C., continental Bittkau, a in 4065-4083. processes 14: Evolutionary Ecology Aegean 2005. the P. of 0.12.2/r18. H. geography Version Package Comes, Towards R of 2019. C., – patterns E. Bittkau, inference. global M. 1116-1142. Multi-model on 94: Johnson, Mu-MIn: Soc. glacio-eustasy F., 2017. Philos. and K. K. Camb. ontogeny Barton, Rijsdijk, Rev. island R., Biol. of Quartau, – impact biogeography. S., island – model: Nuno, marine dynamic islands. B., Sensitive’ Berning, Global oceanic ‘Sea-Level 2018. C., Atlantic a E. Melo, on M. P., organisms Johnson, S. C., marine Avila, A. of 101-112. Rebelo, patterns 126: A., Bul. biogeographic Medeiros, Pol. L., large-scale Marine Silva, on P., 95–99. Madeira, impacts 9: R., Ecology change Cordeiro, of P., J. S. – Avila, area. and subsidence 1227-1241. Species thermal 41: 1921. island Biogeogr. O. oceanic and J. Arrhenius, Gal´apagos. eustasy – of the role on combined the biodiversity Exploring shaping 2014. in C. the J. and Aitchison, 984-994. R., components J. assemblage 44: Ali, biological Biogeogr. the J. of – classification types. substrates: vs island biological effects physical as founder Islands metapopulation: 2016 105-117. dynamic J.R. 76: Ali, a Biol. in Pop. fixation Th. Allele – 2009. effects. A. refuge Lambert D., Claessen Aguil´ee, R., Bibliography ´ ´ nteAga rhplg.–Pio.Tas .Sc od .Bo.Si 7 03–3096. – 3083 27: Sci. Biol. B. Lond. Soc. R. Trans. Philos. – Archipelago. Aegean the in iel arvensis Nigella ud ¨ ¨ ag iceInselbr ¨ aische line(auclca)ifre rmclrpatDA Mol. – DNA. chloroplast from inferred (Ranunculaceae) alliance ce a n elgsh egnehi.–Edud 0 20-30. 20: Erdkunde – Vergangenheit. geologische und Bau ucke. ¨ 9 Nigella Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. abc,K,Ruy . ucl,A,Sn . abig,M 04 e-ee n lblievlmsfrom volumes ice global 15296-15303. and 111: Sea-level P.N.A.S. 2014. – M. Holocene. Sambridge, the Y., to Maximum Sun, Glacial A., Last Purcell, predictions the 1969-1988. H., Model 24: Rouby, LGM: Reviews Mixed the K., Science since Lambeck, Linear Quaternary a Sea – Mediterranean in from the areas. Tests in stable floras Package: change tectonically Sea-level lmerTest island for 2005. 2017. of A. 82:13. Software Purcell, B. diversity K., Statistical H. Lambeck, Global of R. J 2008. – Christensen, W. . B., Models Barthlott, P. Effects G., Brockhoff, , A., 116-127. Kuznetsova, Kier 11: 2017. Letters 44: J., P. Ecology Biogeogr. Mutke, – Trigas, of perspective. W., A., J. macroecological – Jetz, K. revisited. Triantis, H., Archipelago M., Kreft, Aegean S. the system: Simaiakis, island E., complex 651-660. 1755–1668. Georgopoulou, a 8: of T., Div. biogeography and A. Network Ecol. Valli, Plant K., – Aegean South islands. Kougioumoutzis, the Aegean in diversity central species other plant and driving factors Arc Ecological Volcanic Zeitschrift 2014. – A. Tiniakou, Sea). K., (Aegean Kougioumoutzis, shelf continental Cyclades Ana- the D., Sakelariou, in D., challenges f Vandarakis, Biodiversity Geoarchaeological P., 2009. – Drakopoulou, C. I., archipelago. gnostou, Panagiotopoulos, continental K., a Pavlopoulos, Biogeo- V., in 2010. Kapsimalis, P. richness Dimopoulos, species P., 1225–1235. endemic Delipetrou, 19: and K., Conservation 281–295. Georghiou, and total 16: M., for Thessalon. Panitsa, determinants Res. E., graphical Biol. Bergmeier, J. richness S., – species A. (Greece). plant Kallimanis, Aegean affecting Factors South 2011. the M. Aegean in Mylonas, endemism of K., and divergence Vardinoyiannis, genetic K., evidence Triantis, the New A., on Kagiampaki, archipelagos: 241–254. flow island 121: gene continental Bot. and in Ann. colonization Diversification – fragmentation, on 2018. nunculaceae). effect of P. its roles H. and the 262-280. isolation Comes, on 29: island A., Biogeo. of Tribsch, and Islands. nature U., Eco. diverse of Jaros, The Global Encyclopedia 2019. – S. (eds), faunas. , insular D., Meiri bridge P., Clague, land Pafilis, R., J., Foufopoulos, Gillespie, Y., In Itescu, 392-396 - pp. Geology. Press., perspec- : California changing Islands of biogeography: Greek University island 5:249-257. 2009. oceanic Biogeogr. Front. of M., – Models Higgins, archipelagoes. 2013. in A. In dynamics E. – Rickart, biodiversity Archipelago. S., on 87-106. Aegean D. tives pp. the Balete, by London, R., exemplified Press, L. as Academic Heaney, floras islands. island and R of Plants Stiftung, evolution (eds) and ETH, D. origins Bramwell, Insit. The – 1979. W. Skizze. Greuter, Geo-botanische eine Valentine, 161-177. Kreta, In: pp. insel – Press, significance. Die 141-197. Academic evolutionary York: 1975. its New W. and & Crete Greuter, London of Evolution. flora and the Phytogeography Taxonomy, of 329-337. elements (ed.), 19: relict D.H. Boissiera The Crˆete. spontan´ee– Lin. 1972. de flore the W. `a of la Greuter, l’homme J. de Bot. – L’apport Greece. 1971. of W. plants Greuter, endemic the of traits 130-422. Biogeo. and 162: and Patterns Soc. Eco. 2010. P. Global Delipetrou, – K., Fern´andez-Lugo, biogeography. S., Georghiou, island L., of Nascimento, model de glacial-sensitive R., a 817-830. Otto, Towards 25: J., 2015. S. J. Norder, R. F., Whittaker, K. Rijsdijk, M., Fern´andez-Palacios, J. rGoopooi 3 169-190. 53: Geomorphologie ur ¨ 10 bl uih55: Zurich ubel, ¨ Nigella (Ra- Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. h etnclyatv eenrgo.–Gooia oit odn pca ulctos41 145-178. 411: Publications the Special in in London, archaeology Evolution Society Palaeolithic Geological (ed.), and – A. landscapes region. submerged Strid, Aegean Pleistocene active In: 2016. tectonically – N. the Galanidou, and Aegean. D., central Sakellariou, Notiser. 20-28 the pp. of Botan. botanica, phytogeography – Opera The biology. Aegean. reproductive 1971. in H. principle Runemark, neglected a M. Biogeo- drift, 1242-1254. J. 90–129. cycles: Fernandez-Palacios, Reproductive 41: P., sea-level 122: Biogeogr. S. 1969. Pleistocene J. Avila, H. by – C., driven Runemark, archipelagos. B. islands Macaronesian Emerson, volcanic the R., for of Otto, implications changes J., graphical surface-area S. Quantifying Norder, T., Springer-Verlag. 2014. Hengl, 105:1. – F., Denkschr. Aegea. K. Kl., Phytogeographia Rijsdijk, Math.-Naturwiss. 1951. F. Wien. Rechinger-Moser, Wiss. and Akad. K. – Rechinger, Statistical for Aegaea. Foundation Flora R 1943. computing. K. statistical https://www.R-project.org/. Rechinger, J. for URL environment – and Austria. regions. language Vienna A surrounding – R: and Computing. Aegean 18-32. 2018. the Team. 53: M. Core from Res. R Mylonas, taxa Evol. S., animal and of Sfenthourakis, Sys. analyses K., Quaternary Vardinoyiannis, phylogeographic Zoological of of A., – Trichas, review P., A conditions. Lymberakis, glacial 2015. P., Quaternary Kapli, average N., Geology of Poulakakis, Marine implications – geological 245-261. Europe). Some Alpine 32: 1989. Research (SE and C. seas Pleistocene Aegean S. latest and Porter, during Ionian changes the sea-level of of 145-156. morphology impacts Aegean. 196: the The the on 2003. of times N. Biodiversity Aegean Holocene Conispoliatis, and the and Biogeography 279-290. of C. (ed.), pp. biogeography Perissoratis, S. Cyprus, and Sfenthourakis, Nicosia, diversity Ltd, In: 431– Plant Publishers – Hill 36: 2018. synthesis. Broken Oecol. new K. and Acta a Kougioumoutzis, richness – A., Archipelago: species archipelago. Kagiampaki, plant Aegean M., East affecting Panista, the Factors from 2010. example S. an – Sfenthourakis, 437. (East islands I., group land-bridge Gregoris, islets’ on Leros P., endemism islets: Trigas, and 271-284. M., islands 16: Panitsa, Balcanica small configuration Phytologia on archipelago – diversity and Greece). Floristic area, 500-505. palaeo-area 2010. Aegean 27: D. island K.F. Biogeogr., Tzanoudakis, in Rijsdijk, and M., E.E., changes Ecol. Panitsa, Global Loon, of – van database Quaternary. W.D., late explicit Kissling, archipelago the spatially T., during long-lasting Hengl, global by P.A.V., A Borges, explained K.F. 2018. J.B., best Rijsdijk, Baumgartner, 184-197. is K.A., S.J., 28: endemism Norder Triantis, Biogeog. island and E.E., Maximum: Ecol. Parent, Loon, Global Glacial R., van – Otto, Last configurations. Fern´andez-Palacios, J.M., R.H., L., the B.H., Cowie, Nascimento, Beyond M.K., Warren, de 2019. Borregaard, W.D., F., Ib´a˜nez, P.A.V., Kissling, A.M., Rigal, Borges, M., Martins, M.R., C.E., Frias Alonso, de R.J., 611-617. F.B.V., Whittaker, 95: Florens, K.V., Ecology Proios, – values. S.J., P Norder of 1775-1790. defense 17: in In Change islands 2014. Env. Mediterranean P. Regional on Murtaugh, – vegetation change. and global biodiversity of plant face of Press. the vulnerability University Cambridge specific – The Centipedes. 2017. M´edail, of F. Biology The 1981. J. Lewis, 11 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. rgs . ais,M,Titi,S 03 lvtoa rdeto aclrPatSeisRcns and Richness PLOS Species – Plant System. Island Vascular Continental of a on Gradient Uplift Elevational Mountain Post-Isolation e59425. of 2013. 8: Effect ONE The S. – Tsiftsis, Crete in M., Endemism Publishers Trigas, Panitsa, Hill A., multi-taxon Broken P., Trichas, a Aegean. Trigas, the M., Sea: 279-290. of Aegean S. Biodiversity pp. the and Simaiakis, Cyprus, of Biogeography Nicosia, V., regions (Ed.), Lym- Ltd, S. Spaneli, zoogeographic Ch., Sfenthourakis, The K., Georgiadis, In: 2018. Proios, P., approach. S. N., Andriopoulos, Sfenthourakis, K. Probonas, I., Vardinoyiannis, A., Anastasiou, P., Oikonomou, A., P., Legakis, berakis, K., and Kougioumoutzis, biology 215-231. K.A., relationship: 39: Triantis, species–area island Biogeogr. The of 401-407. 2012, J. R.J. 31: – single- Whittaker, Ecography by statistics. and – revealed F. as relationship. Guilhaumon, Sea, curves species-area K.A., Triantis, (Aegean inter-provincial species-area the archipelago Evolutionary for Skyros 2008. Insights in R.J. endemics: envi- Whittaker, snails island M., richness, land Mylonas, Species 1727-1735. on K.A., 32: based Triantis, 2005. Biogeogr. study K. of Vardinoyiannis, case J. and a – K. Greece). area: Press. Lika, University and M.D., Oxford heterogeneity Weiser, – ronmental M., Mediterranean. Mylonas, the K.A., in Books. Triantis, evolution Berlin. Scientific Plant Museum Koeltz Botanical – 2005. and D. Hellinica. Garden J. Flora Botanic Thompson, 1997. Berlin: K. – Tan, and flora. A. Aegean Strid, the of (ed.). Atlas D. 155-179. Valentine, 2016. pp. In: York, A. – New Strid, Aegean. 25: Press, the Academic Biogeogr., in Evolution. problems Ecol. Romeiras, and phytogeographical Global Phytogeography and Taxonomy, J., evolutionary Topography- elevation. Some Price, with 2016, 1972. R., endemism C. Strid,A., Otto, of Beierkuhnlein, increase S., and Irl, global O.R. Nogue, T.-J., Vetaas, a C., 1097-1107. and Huang, J.-C., Kueffer, Svenning, D.E.V., speciation Borges, A.S., T., Harter, isolation, Stuessy, H.J.B., driven J., Jump, D., Fernandez- Greimler, Birks, A., R.B., Strasberg, R.G., F., Jentsch, Elias, M.M., Gillespie, M.C., Attorre, D., R.E., Duarte, C., Jeanmonod, M.M., Gereau, Sequeira, Ah-Peng, promotes S.D.H., R., de P., Gabriel, M., isolation Trigas, Sanctis, J.M., ecological De Palacios, J.-A., Elevation-driven C.-H., Chou, Grytnes, P., Cardoso, R., 2013. P.A.V., 52–56. C. Field, 47: M.J., Beierkuhnlein, Oecologica 157–170. Steinbauer, Acta H., pp. – D. UK, islands. Edinburgh S. P. Mediterranean of Davis, on Irl, Society diversification Botanical In: J., – Aberdeen: M. Asia. communities. Steinbauer, West chasmophytic – South of of Ungulates. aspects life Island geographical Plant plant Plio-Pleistocene (ed.). 241-256. and of 2: Evolutionary Extinction 1971 and Assoc. on Snogerup Evolution Quat. rise French 2005. sea-level the A. of of Geer, J. effects der Int. biogeographic Van Postulating and P. Maximum: 108-119. Sondaar, C., changes Glacial 471: Aegean Hammoud, Geographic Paleoecology Last P., Paleoclimatology, 2017. South the Paleogeography, Stocchi, E. – since the Tjorve, J.H., K.M.C., Sea islands. Boxel, of Tjorve, Aegean Van E., (Chilopoda) the S.J., Loon, van in Norder, fauna E., E.F.M., 241–307. centipede Georgopoulou, Koene, K., 51: The Kougioumoutzis, K.F., Zool. Rijsdijk, satellite of S.M., 2005. J. its Simaiakis, Israel and M. – Crete Mylonas, Mediterranean). of Eastern 367-418. A., (Chilopoda) (Greece, 50: archipelago Minelli, fauna Zool. centipede S., of The J. Simaiakis, Israel 2004. – Mediterranean) M. Eastern A Mylonas, (Greece, Islands: A., islands Mediterranean Minelli, 399–412. the S., 20: of Settlement Simaiakis, Biogeogr. Human of Initial J. the – and Approach. Vegetation Palaeoecological Mammals, 1993. W. Schule, 12 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. l- A003004N 002-.2 002-0.018 0.259 -0.022 0.252 0.368 0.059 -0.022 0.248 0.048 0.362 0.329 0.368 0.048 0.330 0.507 0.362 0.184 0.394 -0.022 0.520 0.788 0.801 0.290 0.330 0.507 0.184 0.447 0.368 0.520 0.779 0.796 0.290 0.362 0.447 NA 0.779 0.330 0.507 0.796 0.184 -0.109 0.520 0.004 -0.109 0.290 0.15*** perfor- 0.447 Model 0.134*** -0.007 bold. 0.779 in 0.003 performance 0.256 best 0.796 overall 0.176*** 0.157 R BIC, with AICc, chorotype on 0.157*** each based for assessed 0.224 models was mance GLMM 0.141* 0.256*** Selected 0.034 2: Table 0.136* 0.298*** aegean-sea-greece 0.253*** 0.204 0.319*** 0.729 lasting-sea-level-stands-structure-insular-species-diversity-of-a-continental-shelf-archipelago- 0.214*** 0.137* -0.034 0.728 0.334*** image1.emf NA -0.034 0.338*** file 0.331*** Hosted 0.245*** 0.329*** 0.367*** 0.326*** 0.353* 0.131 All-E 0.328*** 0.253* N 0.380* S Centipedes R 0.187 All-E 0.299*** 0.298* AE 0.299*** z-values PE G z-values MIEs SIEs N S endemics island endemics. Multiple Aegean is is z-values MIEs AE endemics, endemics, phytoregion island is Angiosperms single PE is endemics, SIEs island species, Greek native is is G N PR+A), species, (= all is 2017. S A. cients. K. Log 6354. Triantis, 1. 357: K., Table Science M. – Borregaard, laboratories. nature’s J., of T. Tables. view Matthews, long the M., Taking J. biogeography: Fernandez-Palacios, biogeography. Island island J., oceanic of R. theory dynamic Whittaker, general 977–994. A 35: 2008. J. Biogeogr. R. conser- J. Ladle, and – A., evolution, K. Triantis, ecology, J., R. biogeography: Whittaker, Island Press. 2007. island University M. Oxford shapes J. – change Fernandez-Palacios, vation. climate and Quaternary J. plant Late R. insular Whittaker, 2016. of 99-102. Anatomy, H. patterns 532: - Kreft, global Nature, J.S., Zoology – from Cabral, biodiversity. on insights M.J., Treatise – Steinbauer, P., isolation (ed.), Weigelt, 417-429. island A. 36: Quantifying Ecography, Minelli, – 2013. richness. In: H. species 310–325. – Kreft, pp.: P., Brill, Weigelt, Ecology. of 1. age – Volume relative Myriapoda, Chilopoda and The Distribution Biology. 11693. Taxonomy, 2011. 9: 2019. K. Rep. R. Pellens, Voigtlander, Sci. – M., Mouchet, worldwide. 37: islands R., Bot. across Govaerts, of endemism T., J. Nordic Haevermans, beta – S., and Greece. Veron, alpha islands, of Ionian Determinants the 2019. systems: P. island Trigas, e02156. Mediterranean M., in Panitsa, diversity E., plant Iliadou, vascular K., Kougioumoutzis, A.-T., Valli, ihpvle ihrta . rVFhge hn25wr eoe.Maigo oe variables: model of Meaning removed. were 2.5 than higher VIF or 0.1 than higher p-values with 10 vial at available tasomdseisae uv aaeeswt enzvle soe n orlto coeffi- correlation and (slope) z-values mean with parameters curve area species -transformed adbig sad reIlnsBt sad adbig sad reIlnsTu sad reIlnsBt Islands Both Islands True Islands True Islands True islands Land-bridge Islands Both Islands True islands Islands Land-bridge Both Islands Both Islands True islands Land-bridge islands Land-bridge Islands Both Islands True islands Land-bridge https://authorea.com/users/314450/articles/444809-extreme-and-short- 2 ,R m, 2 n oc pseudo-R loocv and c 13 2 2 nec fteemdl,variables models, these of each In . R 2 S-A = R 2 R 2 R 2 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. Gavdopoula Folegandros Megalo Fokionisi Elasa Dragonada Donoussa Dia Chios Chalki Astypalaia Antiparos Antimilos Antikythira Andros Anafi Amorgos Alonnisos (km Alimia area Current Alatonisi Aigina Efstratios Agios Agathonisi mainland. the Island to in distance S1 and Appendix area (see insular mainland past and the Current to connection is the considered preceding area 2 Kya) the calculate Appendix (1 Maximum), to Glacial step used 2017). Late time values). were or al. past the level et mainland and at sea Kougioumoutzis current the island were Median between the to at data (difference of 0 distance geographical rise = that All sea-level and (distance by areas 2014). islands caused present land-bridge grid change al. For and geographical a et Past of of Rijsdijk magnitude surface (see 10.2.2. the rise the system ArcGIS sea-level than the in in smaller glacial processed occurring Islands post dynamics lastly. the fragmentation achieved the by was nine km comparable level affecting last (˜1 sea and effects for size this present geophysical minimal km when below regional were (˜1 BP m the cell stand ka 65 2017). that 11 sea-level of assumed al. at We stand median et conditions sea-level (Simaiakis eustatic 2019). median arc-seconds relative this eustatic and al. 30 of global for of et adjustments model a resolution (Norder isostatic geophysical took a cycles hydro we with a glacial-interglacial value for grid on median accounting bathymetric based the and equations For reconstructed topographic sea-level a was generalized on archipelago uses applied Aegean that the change work, sea-level earlier this In 1 Appendix Materials Supplementary level. level; sea median level; on sea based island) true or mainland; level; the sea to distance shortest rent autocorrelation; spatial dAmed dAlgm 2 al A2.1 Table 2 eeecue rmteaayi,adilnssprtdb itnesalrta h grid the than smaller distance a by separated islands and analysis, the from excluded were ) eeageae nteploegahcrcntuto ooti osraieetmt of estimate conservative a obtain to reconstruction paleogeographic the in aggregated were ) nua rals oprdt einsalevel; sea median to compared loss area insular = nua rals oprdt G e level; sea LGM to compared loss area insular = .775 5.1109 8.5169.16 33.62 5.39 87.24 158.82 5.39 200.11 0.00 35.85 33.62 77.00 180.55 5.39 38.21 127.42 40.80 163.17 5.39 159.31 93.48 49.82 2.24 35.85 89.83 203.67 0.00 0.00 0.00 45.12 0.00 40.80 0.00 92.10 180.94 105.95 58.25 131.47 89.89 10.77 101.64 180.85 181.96 76.49 21.21 38.01 27.00 358.21 118.34 210.95 4.12 7616.97 7.07 45.12 79.20 360.01 10906.46 13.00 80.23 31.17 92.85 110.54 7616.97 7.58 59.54 39.61 54.43 952.13 182.34 140.13 105.01 12.08 6.35 360.01 105.38 16.35 53.45 7616.97 37.44 23.48 40.82 29.05 161.09 20.04 1.77 6.71 32.35 952.13 7616.97 82.98 258.77 43.15 1.59 17.89 182.34 4.04 295.05 966.62 2.87 105.01 372.58 15.98 8.97 13.66 34.00 11.88 359.54 843.92 734.17 233.17 80.45 27.20 54.13 95.87 179.61 61.69 123.85 12.47 8.81 1.83 19.68 210.65 441.78 74.41 38.52 121.33 36.39 113.08 7.47 1.83 84.22 42.02 13..44 prov htgorpia province; phytogeographical = 2 raa S (km MSL at Area ) dDmed dDlgm nraei itnet h anadcmae omedian to compared mainland the to distance in increase = nraei itnet h anadcmae oLGM to compared mainland the to distance in increase = 14 2 logApr raa G (km LGM at Area ) typemed typelgm oaih fcretarea; current of logarithm = sadtp ae nLMsea LGM on based type island = sadtp ln-rdeisland (land-bridge type island = 2 urn itnet anad(m S itnet anad(m G itnet anad(km) mainland to distance LGM (km) mainland to distance MSL (km) mainland to distance Current ) Dpr cur- = Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. Skiathos Skantzoura Sikinos Sifnos Serifos Schoinousa Samothraki Rodos Ro Rineia Pserimos Psara Pontikousa Polyaigos Paximada Patmos Paros Ofidousa Nisyros Naxos Mykonos Milos Limnos Levitha Lesvos Leipsoi Kythira Panagia Kyra Kriti Koutsomytis Kounoupoi Koufonisi Kos Kithnos Kinaros Kimolos Keros Kea Kastellorizo Kasos Karpathos Kandelioussa Kalymnos Irakleia Ios (km Ikaria area Current Gyaros Gyali Gioura Gavdos Island 73 23 23 .200 0.00 8.06 5.39 57.71 34.01 5.39 0.00 0.00 13.93 5.39 0.00 0.00 42.05 12.00 33.62 137.54 73.98 158.82 82.42 0.00 58.52 116.40 33.62 5.00 13.34 29.07 5.39 17.09 82.07 0.00 5.39 5.39 19.00 23.85 73.98 4.12 62.36 59.55 0.00 159.31 97.58 0.00 142.27 0.00 33.00 86.83 6.00 3.61 62.80 69.64 146.82 15.23 105.95 72.32 33.62 31.55 5.00 35.36 33.62 87.24 7616.97 105.95 84.63 18.03 0.00 83.44 87.24 161.28 23.85 46.40 97.58 8.06 176.48 5.39 7616.97 61.27 102.42 26.25 72.32 250.23 15.31 32.39 73.98 179.49 22.01 109.18 7.81 0.00 72.12 5.39 1982.09 7.07 48.80 47.13 0.00 7616.97 110.49 109.12 102.86 438.91 16.03 56.13 40.80 173.51 107.62 360.01 84.53 16.64 40.80 84.53 93.98 47.33 225.11 710.79 6.23 0.00 31.17 203.08 4.27 1615.73 120.25 0.00 41.74 59.51 86.05 104.40 50.46 32.53 5.39 77.38 103.37 65.28 360.01 438.91 74.09 7616.97 6.32 0.00 58.24 5.39 75.04 8.13 5.39 97.58 4.03 76.27 180.51 109.66 36.67 2.83 3.61 344.91 1407.68 13.60 7616.97 16.35 9.00 7616.97 2.60 13.45 62.37 169.16 17.03 67.88 14.06 135.00 710.79 14.63 63.78 966.62 85.15 1794.26 5.78 82.76 40.04 80.52 55.84 95.08 2.43 4.00 118.79 221.47 18.12 2007.52 966.62 127.19 144.22 1.53 82.39 7.00 646.13 178.07 37.64 34.22 134.00 344.91 18.25 8.25 38.01 238.19 879.62 19.98 360.01 101.07 3.79 10906.46 57.58 360.01 138.45 41.28 10906.46 2007.52 46.91 187.54 3.00 430.17 19.65 85.80 365.72 438.91 63.78 38.59 38.34 140.94 229.74 196.57 477.03 85.15 182.34 9208.39 710.79 9.12 182.34 7616.97 1636.73 16.49 21.04 146.01 15.87 239.21 16.56 277.23 438.91 145.01 138.82 831.28 24.76 50.70 9.21 19.10 7.27 38.95 8264.62 831.28 344.91 1.51 59.48 30.34 3.98 293.03 191.09 7616.97 4.18 166.65 15.13 288.10 7616.97 149.74 99.43 504.25 7616.97 4.56 27.02 471.05 3.28 48.66 178.07 15.09 272.57 358.21 33.21 130.38 15.13 160.75 66.71 327.96 34.24 393.83 12.69 3.28 21.64 248.26 67.60 18.12 108.45 254.68 17.55 8.42 11.05 32.72 2 raa S (km MSL at Area ) 15 2 raa G (km LGM at Area ) 2 urn itnet anad(m S itnet anad(m G itnet anad(km) mainland to distance LGM (km) mainland to distance MSL (km) mainland to distance Current ) Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. N tnsfr‘aiennedmc’ Alfr‘l neis;A o Aga neis;SE o ‘single for islands. SIEs true endemics’; endemics’. the ‘Aegean island triangles for ‘multiple green for AE MIEs in endemics’; and; ‘all islands, endemics’ for land-bridge ‘Greek EAll for for endemics’; GE In lines endemics’; non islands. regression island ‘native true and for and points islands stands land-bridge data NNE on the angiosperms dots of chorotypes blue selected for relationships Species-area 3: Appendix 84). respectively), = LGM, (n and datasets (MSL centipedes levels and sea angiosperms Maximum the Glacial both Late of and islands Median the at including type island of Distribution A2.2 Table Average Ydra Tinos Tilos Thira Thasos Syros Syrna (km Symi area Current Stroggyli Skyros Skyropoula Skopelos Island 2 2 147 131 0.00 20.62 5.39 0.00 120.33 38.47 0.00 5.39 5.00 61 26.17 30.41 2.83 0.00 20.40 10.77 130.77 0.00 41.62 57.43 1.41 8.00 26.17 74 34.13 2.83 6.08 21.02 50.93 169.92 2104 8.00 70.46 58.82 82.36 1.41 8.00 7616.97 134.76 26.17 252.44 33.62 472.28 329 7616.97 40.87 22.00 82.36 4.80 125.78 734.17 99.55 38.89 123.10 586.30 472.28 127.92 223 372.58 14.48 49.75 102.60 233.43 3.50 61.83 75.74 317.85 6.29 383.75 83.78 141.33 7.94 58.00 3.50 206.93 6.05 125.96 reiln 962 22 79 island 5 True Land-bridge 2 raa S (km MSL at Area ) S LGM level MSL Sea level Sea 16 2 raa G (km LGM at Area ) 2 urn itnet anad(m S itnet anad(m G itnet anad(km) mainland to distance LGM (km) mainland to distance MSL (km) mainland to distance Current ) Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. 17 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. 18 Posted on Authorea 23 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158765865.53573134 — This a preprint and has not been peer reviewed. Data may be preliminary. 19