Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10 9 8 7 6 5 4 3 2 1 Koricheva Julia Szpryngiel Scarlett Ospina Carvajal Valentina Sam Novais Martins Paulo Knapp Michal Aires Marino Romero experiment Gustavo global structural a of arthropods: role for the refuges modulate drought and variability Climate eos Gibb Heloise e unaBntn eerhCenter Research Binatang Guinea New Ecofog UMR CNRS, Sciences of Academy UFMG Czech Entomology, Janeiro of de Institute Rio do Federal Universidade Louis Missouri-St. of Sciences University Agricultural of University Swedish (UNICAMP) UNICAMP Campinas of University State im University PernambucoJimma de Rural Federal Cape Universidade Western the of University Zagreb of University London of University Holloway London Royal of University Prague Mary Sciences Queen Life of University System Czech Alabama of Sciences University Environmental London The of of Faculty University Prague Mary, Sciences Queen Life of University Czech UFMT University Trobe La Park National Forest Bavarian Ecolog´ıa AC de University Instituto State Austin F PernambucoStephen of University Rural Federal 31 10 .Kri Trzcinski Kurtis M. , 28 ebrAntoniazzi Reuber , 35 5 otc Novotny Vojtech , C´assio Pereira , ao Crist´obal-P´erez Jacob , 14 19 24 haoIzzo Thiago , 44 ae Kratina Pavel , sysMendesil Esayas , 1 haoGon¸calves-Souza Thiago , 35 41 zqilGonz´alez Ezequiel , yoTack Ayco , 7 ieaPeri´c Mirela , 6 11 32 ain Cornelissen Tatiana , 15 elyDattilo Wesley , aiaVieira Camila , oa Kadlec Tomas , 20 38 42 eec Luke Rebecca , 25 arcoQuesada Mauricio , itTeder Tiit , aolvMichalko Jaroslav , 22 36 aaPetermann Jana , ain Jausoro Mariana , 2 2 oa Roslin Tomas , 12 16 aai Westwood Natalie , 1 43 rsoBreviglieri Crasso , itraKemp Victoria , atnVidela Martin , 21 7 eoeOrivel Jerome , tfnMajnari´c Stefan , 39 26 ireRogy Pierre , naMrazova Anna , 29 3 evoRibeiro Servio , 37 oetMarquis Robert , tnsKaensin Stanis , 17 34 33 8 oiaKersch-Becker Monica , aiLi Viljur Mari-Liis , hnSui Shen , 2 ai Bernaschini Maria , niaBusse Annika , 22 32 oi Maritz Robin , in Srivastava Diane , 27 Samuel , 30 9 Gustavo , Katerina , 4 9 Nicholas , Fabiola , 13 43 , and , 23 34 , , 40 18 , , Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. nldn ln rhtcua opeiy rebr,dwe od ers eflte,adla hles–re- (Scheffers – extremes shelters – and leaf components variability and climate structural litter, to habitat leaf inhabitants of debris, their variety al woody of a downed exposure by bark, the provided tree duce complexity, refuges architectural Microclimatic plant 2019). including Casas, and (V´azquezbourde niche or elevational climatic (Scheffers and their refuges ranges track geographic they the as in Trisos species shifts of to unprece- led distributions at already vertical intensifying have all Donat and are 2014; mismatches, variability Change, species–environment Climate temperature (IPCC and rates events dented precipitation extreme warming, Climate INTRODUCTION predators, variability for climatic persistence. especially in species refuges, on biotic increase herbivores of change Projected importance by climate the climates. of usage enhance wetter impact to shelter in likely the contrast, therefore and mitigating is In plants in increased regions same shelters geographic elevation. certain within those and in biomass on drought latitude overall predators and of shelters and of diversity leaf leaf regardless abundance variability, Predator of revealed with analyses temperature importance arthropods. increased Our the and of diversity how drought climate. the explore local underlying for to with and elements gradient arthro- elevation to facilitative latitudinal by latitude, key sensitive km built with be shelters 11,790 species changes to leaf an However, arthropods as across terrestrial such experiment for refuges, worldwide. distributed shelters microclimatic biodiversity a in conducted eroding persist We might and variability pods. interactions climate and biotic temperatures disrupting high drought, is change climate Current Abstract 2021 29, January 44 43 42 41 40 39 M´exico.Aut´onoma de 38 37 36 35 34 33 32 31 30 29 Morelia Unidad 28 27 26 nvriyo London of University Tartu of University University History Stockholm Natural of Museum Swedish Columbia British of Mexico University de Autonoma Nacional Nacional Universidad Universidad Sustentabilidad, y Ecosistemas en Investigaciones de Instituto Chilecito Education de of Nacional 5 Faculty Universidad Landau Koblenz of University Caldas de Universidad Biolog´ıa Science Vegetal de of Faculty Multidisciplinario Columbia Instituto British of University Columbia The Entomology British of of Institute University and The Bohemia South of Preto University Ouro de Federal Universidade Salzburg Universitat Superiores Estudios de Nacional M´exico Aut´onoma Escuela de Sciences Nacional of Universidad Academy Czech Nitra Centre in Biology Agriculture of University Slovak ,21;PneoreadCss 09 Pinsky 2019; Casas, and Pincebourde 2018; ., tal et ,22) sohrmast vi des odtos ersra pce a ekmicroclimatic seek may species terrestrial conditions, adverse avoid to means other As 2020). ., tal et ,21;Suggitt 2014; ., tal et ,21)adee aiuaeteronmcolmt (Pince- microclimate own their manipulate even and 2018) ., tal et tal. et 2 ,21) uhrfgscnb atclryimportant particularly be can refuges Such 2019). ., 06 Bathiany 2016; , tal et tal et ,21) uhcagscause changes Such 2018). ., ,21;Essl 2017; ., tal et ,21;Suggitt 2014; ., tal et ,2019; ., et Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. hne nrfg s ypoetn ffc ie oftr 27)ciai odtosa h td locations. study future the predict at conditions and climatic use (2070) provide refuge future we to of Finally, sizes patterns effect precipitation). the projecting and on by (absolute use (temperature scenarios distance refuge change climate geographic in climate both changes of of including of categories measures different size, interpretations each direct of effect geographical for effects d and leaves the Hedges’ control elevation) investigated the and we latitude, on rolled colonization, d) variables) in shelter (Hedges’ (predictor size influences difference climate moderators body colonizing mean how and arthropods standardized test biomass of the To diversity richness, site. calculated and species and abundance abundance, leaves, recorded arthropod (unrolled) we between control rolls, km 11,790 leaf and an of rolled across effects manually sites beneficial 52 the at experiment measure To coordinated global, 45.8 arthropods. a climatically (from dependencies conducted by gradient and we latitudinal climatic use predictions, arid refuge current above in of the expect test patterns especially we To geographic refuges change, in climate in changes continuing dominant future with require be reflect Finally, typically to to 1). organisms predators (Fig. Larger-bodied of larger regions (Petchey requirements size. variable expect prey metabolic we body their the Therefore, larger than influence 2012). for conditions to selects climatic known favourable (predators) with We are levels more vary position. which will trophic of trophic climate higher with to both and at increase responses size, size will species’ body rolls (Daufresne body that and ectotherms leaf leaf expect by within position also of affected arthropods We trophic by usage drought. is of their provided the and biomass usage mechanisms variability affects and shelter climate facilitation richness climate increasing how on species projected abundance, scenarios and and that climatic arthropods, current hypothesize future terrestrial how and by compare current We rolls scale. of engineers. global effects ecosystem at the leaf-rolling interactions facilitative test on we climates, Here, future of impacts (He the conditions predicted stressful or under or increase latitude competition to and expected of al predation is effects as organisms such indirect among interactions, interactions the antagonistic from on focused replicated climate (LaManna have standardized of studies using effects macroecological (LaManna investigated these direct interactions recently the biotic been on differentiate elevation have can signatures which climatic experiments, in patterns Large-scale interactions. facilitative the such underlying in driver pronounced (Romero 2013). common particularly regions Romero, seem arid and al (Schemske in engineers (Vieira et latitudes also ecosystem time lower but invertebrate of at tropics, and their stronger period wet by vertebrate are long abandoned of which a are interactions, effects assemblages for shelters biotic facilitative arthropod such arthropods other influencing After many other thus 2013). to plant, for Romero, Similar available whole and the become (Vieira to they scales extend creators, and microspatial seasons, and dry temporal and in at rolls stronger (Wang leaf even as organisms be such plant-dwelling can shelters, other galls, leaf many build produce Cornelissen to to thrips) 2013; physiology refuge and plant providing mites manipulate refuges aphids, thus gallers caterpillars, structural ties, galleries, (e.g., as trunk arthropods and used various stem be make and can borers are Hastings that stem al. which structures and et physical beetles engineers, (Jones creating ecosystem organisms by by decline. other habitats provided insect by their typically to modify is leading that factors conditions organisms environmental climatic negative harsh various against the Protection mitigating factor (Garc´ıa-Robledo one change be global may by use, threatened most taxa al., the et amongst are and regulate (Pinsky ectotherms to ,21) oeprclsuist aehv xmndteipcso urn lmt n lmt variability, climate and climate current of impacts the examined have date to studies empirical no 2015), ., ,21)adtmoa atrs(iiaadRmr,21)sgetta lmtccniin a ea be may conditions climatic that suggest 2013) Romero, and (Vieira patterns temporal and 2015) ., 08 anr 00 Wagner 2020; Wagner, 2018; tal et ,21;Roslin 2017; ., tal et tal et ,21;Priest 2015; ., tal., et ,21) uha rhoos cohrsdpn netra nryt thermo- to energy external on depend Ectotherms arthropods. as such 2019), ., 09 n hi aaiyt ispt et(Rubalcaba heat dissipate to capacity their and 2009) ° tal et o60.2 to S tal et ,21;Romero 2017; ., tal et ,22) h viaiiyo hs eue,adteripc ninsect on impact their and refuges, these of availability The 2021). ., ° ,Fg )adeeainsann rm5t 90maoesalevel. sea above m 2900 to 5 from spanning elevation and 1) Fig. N, tal et ,22) h oeo efrligegnesa idvriyamplifiers biodiversity as engineers leaf-rolling of role The 2021). ., tal et tal et ,20;Romero 2007; ., ,21;Roslin 2017; ., ,21) hrfr,teelttdnl einl(Romero regional latitudinal, these Therefore, 2015). ., 3 tal et ,21) lhuhteiprac ffacilitative of importance the Although 2018). ., tal., et tal et tal et ,21) ntretileoytm,bark ecosystems, terrestrial In 2015). ., ,21;Romero 2017; ., 99 Voigt 1999; tal., et 02 iiaadRomero, and Vieira 2012; tal., et tal., et tal et tal et tal et 09.Competition 2019). 03 Brose 2003; ,21;Romero 2013; ., ,21) otof Most 2018). ., tal., et ,21;Warren 2016; ., 09,the 2009), tal., et et Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. eprtr bo,bo,bo,bo) orvralsfrpeiiain(i1,bo4 i1,Aiiy,tovaria- two Aridity), bio15, bio14, topographic (bio12, and precipitation for for variables data variables four Climate four and chose bio7), (http://www.worldclim.org/) we 1). (ii) bio4, site 2 (Fig. and bio2, each version For (bio1, scale, usage (https://envirem.github.io/). 2017) temperature experiment 2017) refuge climate Hijmans, Bemmels, the that on and and at variability (Fick prediction (Title WorldClim ENVIREM the climate arthropods from test by future extracted to shelters of were (i) of data effects purposes: use the two the predict for influence to data would climate drought current and of variability sets different extracted plant. We per data leaves body productivity sampled individual and all of of topography number al- sum the Climate, taxon-specific the abundance, by represented published weighted biomass arthropod through were Arthropod analyses: variables mass size. 70 for All dry body at masses. used mean the (dried were and calculating mass biomass variables richness, then dry dependent species the and possible detritivore, Four (except from (H´odar, length herbivore, level 1996). estimated eye equations parasitoid, body taxonomic was naked lometric lowest predator, total mass) the measuring (i.e., the body by guild to to (dry immediate- or invertebrates feeding visible size or and the invertebrates body sorting identified morphospecies Individual the later We omnivore). into all for ethanol. them collected frozen in classified We either them and invertebrates. were stored replicate leaves the per and The grouped collect mites) them bags. to stored plastic and sorted leaves, zip-lock control ly small and rolled in collected control shelters treatment we size. both leaf experiment, and leaf of the that width of of showed Maximum days estimate bioassays 10 2013). an previous After Romero, as as and measured (Vieira sufficient were was hours) deemed leaves leaves 24 control The was rolled and (within 2013). This and Rolled quickly Romero, field. hairpin. very and metal colonized the Vieira a be and in with can S1 days marked leaves Fig. 10 unrolled (see for 5-10 hairpin had exposed metal pair a the with 17 of least secured plant at control were from rolls caterpillars The by (Fitzgerald 2007). built to Marquis, Erebidae shelters axis Lasiocampidae, Geometridae, mimic leaf Oecophoridae, Tortricidae, to Gelechiidae, the Gracillariidae, S1) Nymphalidae, the to (Fig. Pyralidae, Hesperiidae, on transversally diameter including leaves surface families, in the butterfly abaxial cm and Then, the 0.6 moth leaves. roughly to from of adaxial hand cylinder the by a from removed form rolled were leaf-rolling damage. manually apparent present the were without for arthropods plant leaves used any treatment expanded was fully plant experiment, second 5-10 the the selected to we whereas Prior plant, control defence each a On indirect as same treatment. chosen of addition) randomly the type (shelter was 2011). to apparent pair Koricheva, each any belonged and in bear always plant (Romero not One pair density, trichomes) did each relative glandular that other. low within nectaries, species each plants and extra-floral plant from diversity the domatia, native common apart species but broadleaf (e.g., most m high used blocks, the 6 the only typically among We least (blocks). to species, species species. at due plant herbs plant was native forests, or different single block tropical shrubs used a Each some we trees, using other. in paired conducted each However, a 20 was locally. from followed experiment to species apart the experiment 10 m sites, Each chose 2 most protocol. randomly least In we standardized at were site, a plants each following Paired for conducted design; were block sites randomized 52 is all features, at plant Experiments and samples 500 environment arthropod least including and at sites, experiment were the Global them of of description 94% detailed S1. as Table A in independent, shrubland other. provided xeric spatially each forests, sites from temperate our apart and considered tropical km sites We including 52 biomes, etc. in of savannas (45.8 types 2017-2019 (caatingas), gradient diverse in comprising replicated latitudinal level, was ecosystem-engineering sea km experiment above of The 11,790 presence plants. an on the caterpillars across simulating leaf-rolling by experiment built geographically-coordinated structures a performed We sites Study METHODS AND MATERIAL ° o60.2 to S 4 ° )adeeainsann rm5t 90m 2900 to 5 from spanning elevation and N) tal et ,19;Fki 01 iland Lill 2001; Fukui, 1991; ., o o 24h) for C Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. rcptto bo2,peiiaino retmnh(i1) ne ftedge fwtrdfii eo water below deficit water of degree the PC1 of S2-S3). index (Figs. (bio14), scales month local driest and of 1) precipitation (Fig. (bio12), global precipitation the at Conversely gradient, (PC1 variability. wet) variables temperature to precipitation lower (dry for and PCA warming of to axis variability first temperature higher from spanning (Romero (PC1 multivari- (PCA) variables in analysis variables temperature component temperature for and principal precipitation using separately space herbivores) herbivores. projected ate we for of gradients, leaves (abundance climate as experimental the prey on well create To potential predators as available of elevation, included abundance during and temperature moderators and average predators local latitude and for Additional size absolute leaf experiments. including topoWET), (TRI, the moderators, topography (annualPET), of productivity categories climate, different investigated We for (i.e., respectively large (predictors) guilds sufficiently Moderators sizes, feeding herbivores. only different was effect and size into predators large sample arthropods of the and surveyed analyses parasitoids), the separate medium and classified omnivores small, detritivores, initially herbivores, considered we predators, Although are control leaves. 0.8 the control to and to relative size 0.5 body 0.2, mean or (Rosenberg around biomass richness, Values abundance, species leaves. arthropod higher had rolls leaf and respectively, where saudne pce ihes ims n oysz)wscluae ttest ee n=5 ie)using sites) 52 = (n level site (measured the structure at community calculated Hedges arthropod Rosenberg was difference; on size) mean effect body in (standardized and shelter researchers sampling d biomass leaf several in Hedges’ richness, biases by the for species conducted of control abundance, was to magnitude as approach sampling The (meta-regression) the meta-analysis 2013). a protocol, used (Gurevitch, standardized we Thus, a biomes. used different have we though Even the over average calculations size the Effect calculated 2.5 then x we 2.5 data analyses of which Statistical resolution we from spatial data, experiment, a such the has (Kemp of missing database (Kemp database days sites RNCEP RNCEP 10 For The the the experiment. from all experiment. temperature for 10-day daily done the was near-surface during mean temperature the database. (RCP8.5) local extracted r MIROC5 average correlation, on (Pearson measured only models We analyses climatic our predictive focused two we these 0.98), between [?] correlated strongly very were and variables (RCP8.5) MIROC5 CO using of 2 pathways version al concentration WorldClim representative from resolutions. as extracted arc-second was (RCP8.5) 10 30 scale CCSM4 (Pearson and local on resolutions at arc-second data for two database only 30 climate only the Future used at available between we extracted were correlated data 0.98), were strongly ENVIREM only [?] AnnualPET were from use r variables and topoWET to correlation, these Aridity and decided since variables strongly we TRI resolutions; The very variables 0.97), arc-minute were The resolutions. variables [?] at arc-second resolution. above r extracted 30 arc-minute the correlation, were 10 variables (Pearson Since WorldClim a resolutions resolutions. S2. on different arc-minute Table these 10 the (AnnualPET). among productivity and in site presented correlated arc-minute denoting is 2.5 variable variables arc-second, one the 30 and of topoWET) description and detailed (TRI A topography to related bles ,21) uuebolmtcvralsaentaalbei NIE aaae.Snetebioclimatic the Since databases. ENVIREM in available not are variables bioclimatic Future 2018). ., Xe tal., et 0 d and = tal., et 2012). Xc SDpooled Xe SDpooled r h encmuiymaue o lnsta a hi evsrle n oto plants, control and rolled leaves their had that plants for measures community mean the are − 03.Lre ffc ievle enlre ieecsi cuaino oldcompared rolled of occupation in differences larger mean values size effect Larger 2013). Xc , stepoe tnaddvaino h w rus oiieeet niaethat indicate effects Positive groups. two the of deviation standard pooled the is temperature hc xlie 8 ftettlvrac)rpeetdagradient a represented variance) total the of 78% explained which , precipitation tal et 5 ,21)floigteformula: the following 2013) ., 4 ftettlvrac)rpeetdarainfall a represented variance) total the of 64% : tal et 2 ,21) crso h rtai fPCA of axis first the of Scores 2018). ., msinpoetdfr27 (Romero 2070 for projected emission o n eprlrslto f6h 6 of resolution temporal a and precipitation nlddma annual mean included tal., et 02.This 2012). et Posted on Authorea 29 Jan 2021 | The copyright holder is the author/funder. All rights reserved. No reuse without permission. | https://doi.org/10.22541/au.161193241.11465772/v1 | This a preprint and has not been peer reviewed. Data may be preliminary. etvralsadteHde’deetsz audne ihesadboas n=5) eprtr (bio1, Temperature 52). = (n biomass) and richness (abundance, (Gaussian size models effect linear d using Hedges’ tested the associa- function were and their on the variables definitions) and dent used variable engineers site We for each ecosystem S1 at distribution). of Table observed error (see communities effect variables arthropod scale. the climatic on experiment with of effects the tion engineering magnitude at the the obtained of magnitudes d) on The (Hedges’ change sizes climate effect future the of using arthropods effect the mapped We maps variations: temporal and spatial Predicting regression predictive the treatments generate to of models effects LME and these environment. the predators used evaluated then using by We and graphically use effect. 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(PC1 with components overlap climatic ( not underlying models did intervals equation confidence structural 95% Piecewise their model with if Q-test significant Cochran’s statistically using and considered estimated experimental), were were or effects (control through moderator treatment The a fitting within treatments. replicates (REML) among models likelihood among variation maximum variation fixed mixed-effects restricted random a through a using examined obtained assuming was were were estimator, size models plots moderators effect mixed-effect Overall Such diagnostic the (Q– outliers. and quantile–quantile through 2010). and (Viechtbauer, normal residuals models, outliers of using random-effects terms inspected (linear residuals in through we the behaviour approach obtained appropriate of analyses, presented meta-regression normality models the our using inspected plots; to visually sizes Q) also Prior We effect distance). d 2019). (Cook’s Team, Hedges’ Core on Development moderators (R the using of conducted effects models), the tested We moderators modelling intercept. no equation the i.e., structural plus effects, and (intercept-only moderators Criteria Meta-analysis through containing models Information obtained models Akaike Null then smallest with ([?]AICc>2). were the contrasted AICc present were models sizes, to included) linear sample those best-fitting variance small were The retained their for models on corrected 2. best based < the VIF models selection; had the backward from moderators removed remaining thus All were (Zuur alPET. and diurnal (VIF) S4), or factor (Fig. bio7) inflation S3). collinear bio4, were Fig. (e.g., moderators 1, intra-annual Some higher (Fig. as by temperature defined characterized in temperature Sites variation those from values). bio2) were gradient (positive (e.g., variable variability a more (bio7). temperature producing lower thermally thus range and being trends, warming annual precipitation to temperature values) with (negative matching and variability interpretation, (bio4) of PC1 seasonality ease suitable original the temperature more of climatically (bio2), signs being range as The 1, defined diurnal Sites (Fig. mean PC1 S2). variance higher (bio1), Fig. PCA1 by 1, characterized the (Fig. those to values) were PC1 much (positive The conditions the contribute wet S1). not Therefore, to (Table values) (bio15) did year S2). (bio15) the variability along Fig. rainfall variation) denoting of variable (coefficient seasonality only precipitation and (aridity) need mods ruett goeteitretand intercept the ignore to argument visreg metafor temperature tal., et BeeyadBrht,21)and 2017) Burchett, and (Breheny precipitation akg vrin210 Vehbur 00 mlmne nRenvironment R in implemented 2010) (Viechtbauer, 2.1-0) (version package 00.Termvdmdrtr eeLttd,tpWTadAnnu- and topoWET Latitude, were moderators removed The 2010). crswr utpidb 1t hnetergahcpeetto for presentation graphic their change to -1 by multiplied were scores temperature pSEM precipitation nlme ersne analgain,wihvre rmdy(negative dry from varied which gradient, rainfall a represented predict.lm eeue ots ietadidrc ffcso aiueand latitude of effects indirect and direct test to used were ) akg nR(Pinheiro R in package n PCI and crs PC1 scores. 6 ooti h rdce eainhpbtenindepen- between relationship predicted the obtain to precipitation rma piecewiseSEM temperature ucin(ictae,21) ffc sizes Effect 2010). 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Other variable S7). increasingly (Fig. with climates richness wetter and species under climate abundance, shelter Predator drier use to a S7). likely in more (Fig. were refuges predators herbivores within and contrast, herbivores increased by contrasting biomass revealed usage data and shelter raw with (Fig. leaf modelling size effects of mixed body patterns linear predator using average performed visualization than data strongly smaller Likewise, 41% was was leaves P size control t=12.4, body on 0.88, herbivore = and S6). Average r shelters P=0.38). (shelter: abundance leaf t=0.88, P individuals higher within 0.13, the t=6.7, a of biomass 0.74, with size Herbivore plants = effect body on r the mean 3). also of usage, the magnitude but with shelter the sites, Fig. correlated Thus, on wet positively shelters 1, in leaf P=0.33). (Table stronger of t=0.98, predators effects were positive 0.14, biomass of predators, = and higher. on abundance r rolls is herbivore leaf variability control: of on temperature effect 0.11; where the = to latitudes, predator contrast P higher In with toward 1.6, correlated increased = positively predators, t larger was 0.23, P by leaves = mostly t=9.6, control effect r 0.82, indirect on = positive (shelter: (PC1 and r a abundance shelters shelters had leaf (shelter: leaf but in size within arthropods, biomass body by predator biomass usage in Predator refuge rolls increase (Table influence leaf the regions S5c). were of directly on warmer metrics effects gradient not in community positive did temperature decreased species both the Latitude via but predator variability, on contrast, 3). temperature with In shelters greater Fig. leaf correlated 3). with 1, of positively increased Fig. effects biomass 1, strongly the (PC1 (Table predator was gradients thus drought on plants precipitation and with by on P<0.001), increased influenced t=12.2, abundance richness similarly 0.86, of Predator species = magnitude and (r The 3). abundance richness predator biomass). Fig. on precipitation richness, of 1, rolls (abundance, effects Table the leaf moderator metrics but any of climate, predator by by effect on solely influenced the mediated depending significantly were varied not predators on were temperature rolls assemblages leaf and significantly arthropod of effects were pooled The on not size 1). intervals rolls (Table confidence body leaf 95% of mean on with effects values, and The rolls d biomass Hedges’ leaf positive richness, of (i.e., leaves species presence 0). control abundance, spanning to the compared Arthropod by rolled in influenced (predators, higher 2). strongly guilds feeding were (Fig. all omnivores) including plants assemblages parasitoids, arthropod detritivores, pooled herbivores, the and herbivores predators, Overall, interactions Current RESULTS a using tested was maps performance distribution (TallavaaraModel modelling were patterns to variables ecological approach response global this the of 2020). used through of interpretation ., have residuals outliers geographical studies and a and Previous provide influence), homoscedasticity to necessary. and normality, when checked d, We transformed Cook’s fit influence log qq-plots, model model. the (e.g., the circulation improved eliminated inspections that general graphical first those a therefore and as We variables (MIROC5) significant (Zuur S3). R the (VIF) to higher and only factor (i.e., us retaining S2 inflation allowed selection, variance modelling backward Figs. the predictive through (see the using collinear models in collinearity different variables were of in climatic variables biased the most be the using models may of addition, among estimation reliability structure In parameter prin- the correlations thus the inspect 1998). the and than Ling, because change, rather and used variables climate were (Hadi climatic with variables meta-analyses Separate continuous shift our independent may definitions). in variables as variable identified for included scores were S1 component Table bio15) cipal (see bio14, models (bio12, separate precipitation in and bio7) bio4, bio2, 2 aus.W oeldtecretciaeseai n hs rjce o27 sn rcp8.5 using 2070 to projected those and scenario climate current the modelled We values). < .0) u o ihteraudne(hle:r=00,t03,P07;cnrl = r control: P=0.71; t=0.37, 0.05, = r (shelter: abundance their with not but 0.001), predict.lm < k ucin oee,frbt eprtr n rcptto climatic precipitation and temperature both for However, function. .0;cnrl .4 1.,P =16.6, t 0.94, = r control: 0.001; fl rs-aiainapoc seApni S1). Appendix (see approach cross-validation -fold 7 tal., et 00.W hnrdcdtemodel the reduced then We 2010). < .0) u o ihpredator with not but 0.001), tal et temperature ,21,Gusmao 2018, ., < , .0;control: 0.001; β .8 Fig. 0.28, = precipitation tal et , Posted on Authorea 29 Jan 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161193241.11465772/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. gis ogdogt.Hwvr efudrfg sg ob ihrfrlre rdtr.A agrpredators larger (K As ones predators. small refuges larger to for than microclimatic 2012); sensitive higher stable ratio (Edney, be species offer surface-to-volume to desiccation by and usage smaller transpiration, refuge to occupied may a found leaf show susceptibility we be drought via However, varying humidity to that droughts. with have tend long likely them against taxa refuges provide is the could arthropod It shelters as predatory leaf predatory shelters, Indeed, temperature. by leaf and especially within desiccation. use, precipitation composition were refuge of biomass) species of richness, components shape species determinants by (abundance, key communities influenced the predator of differentially be metrics to different However, appear climatic arthropods. drought harsh and against protection variability for Climate than 2020) Marquis, and (Baer pressure enemies predation natural dependent conditions. against under where refuge are predators 2012), a (Romero Novotny, which vertebrate as high and more against haemolymph), often (Tvardikova refuges is prey conditions herbivores seek (green larger climatic (e.g., might on favourable food herbivores sources from and larger water water Conversely, stable of stochastic more source rates. more predictable capture more on prey a on rely access Predators (see and 2019) (Petchey arthropods Casas, leaves). adversities risk predatory and climatic (Pincebourde where than to plants variability, smaller vulnerable using less temperature often are are and likely insects Brose drought thus and Herbivorous increasing S6), with conditions. Fig. and sites harsh herbivores (Wetzel drought that in in increasing higher suggests microhabitats refuges with This is and of more trends. predation herbivores refuges use opposite by of used showed four use the all herbivores predators refuge avoid on by while of so refuges might diversity, do pattern of differed: they the abundance, use climate and variability, However, local plants, to temperature on conducted. the response levels was increase trophic in experiment different rolls predators from the leaf which arthropods by of on provided size continents body refuges mean structural and that biomass indicate results Our larger fact increasing on the of shelters from scenario of stemming effect a likely the S9). under DISCUSSION (6-17%) 6.7%), effect However, decrease (Fig. to current to Europe up future. predicted versus in of the is future especially (e.g., warming increase in predators, in general variables an little a differences and temperature change temperature, 2.3% smaller annual key will pre- to mean as itself than reflecting up variability manifested variables time of temperature is over decrease bioclimatic that effect variable a key over weak from more variability Indeed, (ranging This be temperature sizes 4b). to to S2). predicted response (Appendix (Fig. are in bio7) precipitation bio14) changes tem- to weaker (e.g., the [bio1]). responses revealed on cipitation temperature to depended biomass annual predators predator compared mean larger for time, versus by [bio7] effects usage variability of refuge temperature Maps on (i.e., 34%, temperature evaluated and of component 25% influence to perature the up by the of decrease Borneo, scenarios to and Future expected S8). USA, are Europe, Guiana Fig. abundance French in 4a, and (e.g., sites richness increase decline (Fig. for species to to respectively predator predicted predicted expected on precipitation is is refuge increasing precipitation predators of with where of effects contrast, areas abundance In in and respectively, richness Mexico). 38%, species and and on 33% refuge to of up effect by the of magnitude The influence directly not interactions 1). did predicted (Table temperature Future arthropods average on local shelters and of effects topography the productivity, ecosystem elevation, as ieymdae ylrepeaosbigsbett h togrpyilgclcntansipsdb tempe- is by effect imposed constraints This variability. physiological stronger temperature the by to influenced subject most being were predators dominant predators large by competitively larger mediated that be likely reveal can also predators results refuges. larger Our available desiccation, using to in ones vulnerable smaller less over being Despite desiccation. tal., et 02.I diin ncnrs omn rdtr,hrioe a ul hi w refuges own their build can herbivores predators, many to contrast in addition, In 2012). tal., et 06.Sc rdto ikmgtdiehrioe osuboptimal to herbivores drive might risk predation Such 2016). tal et ,21) ti hrfr ieyta ebvrsuela shelters leaf use herbivores that likely therefore is It 2018). ., 8 uhsel ¨ tal et ,21) hyaelkl espoeto prone less likely are they 2017), ., tal., et 99 Voigt 1999; tal., et 2003; Posted on Authorea 29 Jan 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161193241.11465772/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. s n hi xesosit h uueflgaptnileoytmtasto.Ipraty hyietf a identify they Importantly, functioning transition. ecosystem on ecosystem research refuge future potential on for avenue ecology. a impacts important community climatic flag an of and as future (Clark patterns research current microclimatic the predators in seen, into and predator- scale be minuscule extensions herbivores in to their remains mismatches larger scenarios and and of these prey, use of extinctions and realism asymmetric predator the by cause While 2020). use could increasing refuge (Barton Likewise, rates, microclimates on control. encounter suboptimal reliance biological prey to Asymmetric herbivores as services displace 2009). ecosystem such could and Schmitz, important change 2016) variability climate and perform Schmitz, with climatic to predators and thus Increasing by physiological and Rosenblatt usage change. feed refuge their 2009; community to performance, Schmitz, to overall predators arthropod their such and according on on of (Barton ability effect climate space strong the of niche a decrease mitigate have imprint may of can still drought overall may refuges use the refuges such The on their from changes. and forage adjust global results to to ability likely to The dominance predators sensitivity tolerances. This allowing refuges. higher using variation, and/or organisms dominance climatic dominant pressure the competitive as predation higher predators intraguild from larger predators strengthen to by point even shelters results and of Our herbivores, usage for refuge on future especially Increased of risk importance, decrease. pattern in predation plants. to increase global upon alter expected to the strongly is expected change rainfall then are where could might engineers sites conditions ecosystem in patterns future, climatic and the the from predators in engineers In projection changes ecosystem arthropods. direct by future by a provided As use that drought. refuges predict and components, that variability we structural evidence climate habitat observed, clear in as increases present such predicted we Efficient mitigate conditions, Here, strategies. may local adversities. mitigation preserving cli- and climatic and conservation with buffer improved identifying deal to for at arthropods pivotal how aim thus Understanding should is use. strategies extremes microhabitat arthropod on and terrestrial influence oscillations droughts, (Pinsky major mate extreme microclimate a of suitable having most frequency on position in the dependent trophic and among more variability also climate be are in should Klink Be- increase they communities (van energy, pollination. With changes global webs. 2021). external and ongoing al. food on cycling to et intricate due dependent nutrient Earth in closely control, on elements and organisms biological key threatened ectothermic as are are such and arthropods services, species, to cause ecosystem terrestrial response valuable all evolutionary provide of an Romero, They two-thirds be and over might comprise Omena shelters leaf Arthropods (de actively using environments to of variability. known fire-prone behaviour well climatic the and are and that did spiders) drought conditions suggest days) (e.g., we harsh 10 Predators that Therefore, over fact predators. in 2010). temperature the on microhabitats (average by shelters itself supported sheltered leaf over is experiment of select year hypothesis the influence the This of the selection. of duration moderate availability. by part the favoured not shelter some spanning be to least variable may at response climatic general changes during predator in the variability not with use climatic and shelter correlated higher climate, then positively characterizing experience in time, was thus variability predators variable, variances, increasing of climate and that communities the means found If of We annual mean site. both each the of of in consist conditions climatic considered overall we less the that being large variables ability, despite oscillations, competitive climatic temperature higher The extreme their against to (K refuges Due desiccation on climates. to rely variable (Horne thus vulnerable thermally ratios may more area-to-volume particular in surface in smaller threat predators their a of causing because 2017), organisms ., larger in reduced is dissipation) (Voigt predators (Caillon as waves (Stireman heat levels (Vasseur temperature and trophic warming fact, oscillations higher In ture than from conditions. arthropods species climatic for to such especially risks under ability greater competitive imposes strongest variability the having and/or rature tal., et 03 Brose 2003; uhsel ¨ tal., et tal., et 2017). tal., et 02 Horne 2012; 04,wihaehrflepcal olre raim,such organisms, larger to especially harmful are which 2014), 9 tal et tal et ,20) eue ih mloaetempera- ameliorate might Refuges 2005). ., ,21) ti nw htha rnfr(or transfer heat that known is It 2017). ., tal et tal et tal et ,22;Wge,22,Wagner 2020, Wagner, 2020; ., ,21;Colinet 2014; ., ,21) ihbd ieand size body with 2019), ., tal., et 2015), tal., et tal et Posted on Authorea 29 Jan 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161193241.11465772/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. rhn,P n ucet .(07.Vsaiaino ersinmdl sn visreg. using models regression of Visualization (2017). W. 56-71. Burchett, temperature a and increasing & in predict P. Breheny, models effects Climate predator (2018). countries. T.M. multiple poor &Lenton, in transforms M. variability Scheffer, warming V., Dakos, Experimental S., Bathiany, (2009). O.J. web. food Schmitz, grassland & B.T shelter Barton, among community. interactions caterpillar complex shelter-building parasitoids: a and in 2198. predators parasitism Between and (2020). predation R.J. traits, Marquis, & C.S. Baer, Council Research Estonian no. of the University (grant REFERENCES by Czech Council supported Sciences, Research was Environmental M-LV Estonian of PRG741). 42900/1312/3166). Faculty the no. no. the by (grant (grant of supported Prague Agency was Grant Sciences TT Internal Life (grant field. the Prague by the and in Sciences PRG741), help Life M´exico thank Aut´onoma (PA- for MQ de of Nacional (2019-LN299033); Sanchez LANASE Universidad (CAPES) University Gumersindo to from field CONACYT-UNAM Czech N´ıvel Superior grants 291333, in SADER-CONACYT by the IV200418), de supported assistance PIIT by was Pessoal for supported MQ de 42900/1312/3166). Aperfei¸coamento Courcelles was Coordena¸c˜ao Croatian no. de EG Danielle the by acknowledges 88882.314749/2019-01. to supported access MKT Proc. was allowing (306572/2019-2). CV for CNPq Croatia work. Medvednica thank of Park SPR Nature Republic the sites. the thank of SM 04-048/2019/P Energy and n. and MSP GAJU Environment study 001). the was of code acknowledges JM Ministry (financial AM Shimales. and Tamiru CAPES VEGA2/0100/18. acknowledges acknowledges grant EM Dunedin for CCP national fieldwork. Dale the grant. Slovak the Esther acknowledges and the of PMM Jacks stages by field. Penelope different supported the thank during and in support collection, help valuable data for their allowing conduct Alexander for to administration Graham permission Garden’s and for Botanic Kalahari Life Maritz Tswalu of Bryan acknowledges University and RM Czech fieldwork 42900/1312/3166). the the grant no. by from by (grant funding supported supported Prague acknowledges was Future was VK Sciences MK Council 42900/1312/3166). TK Council. Research no. Research 309552/2018-4). Australian (grant Environment Prague (grants an NE/K016148/1/Natural Sciences by CNPQ Life by supported of Ecolog´ıa was University supported de Czech HG (Instituto is Mancha possible. TJI La fieldwork the costeras (FT130100821). the Investigaciones Fellowship of de made staff Centro have the and who all Niebla A.C.) de and of Bosque staff Coates del the Rosamond 2014-2020, Santuario M´exico), Aut´onoma besides to the PO-FEDER de Nacional grateful the (Universidad by are Tuxtlas and RA grant Los Biolog´ıaEstaci´on ANR-10-LABX-25-01) Tropical and d’Avenir de supported (CEBA, WD Investissement is Recherche GY0007194). an which la (307210/2016-2) (BiNG, by Guiana, de R´egion Guyane CNPq French provided Nationale in acknowledges Agence support station the TC financial research of and Paracou financial #2013/0877), grants. (https://paracou.cirad.fr/) the for (PNPD/CAPES 669609 to CIRAD program grant access ERC by (STARS) acknowledges research and Scientists JO Khaja 20-10205S post-doctoral Research CAPES. RJM Zhaid as and CSF a Park, Teachers the Sciences. for State and acknowledges River Agricultural CAPES Students VN Cuivre senior acknowledges of the to a NACM University and access and Swedish field, for 322266) support. the the Resources (grant in of Natural Finland help of Chancellor of for Department Vice Academy Missouri the the the by from thanks Fellowship supported Advanced grant Newton was Society, support TR Royal career the NAF/R2/180791). from (FAPESP: funding Foundation grants acknowledge no. research Research gratefully through (grant PK Paulo CNPq-Brazil and and Sao GQR 2016/01209-9) the GQR. Proc. to by BPE provided and research 2018/12225-0, 2017/09052-4, for grants support financial acknowledge We ACKNOWLEDGMENTS clg Letters Ecology cec Advances Science 2 1317-1325. 12, , ,eaar5809. 4, , 10 ucinlEcology Functional h Journal R The 4 2186- 34, , 9, , Posted on Authorea 29 Jan 2021 — The copyright holder is the author/funder. All rights reserved. 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Data may be preliminary. experiment variability-and-drought-modulate-the-role-of-structural-refuges-for-arthropods-a-global- Figures.pdf of end the at section file Information Hosted Supporting the in online found article. be the statistical may common information avoid supporting to Additional exploration data for protocol INFORMATION SUPPORTING A (2010). York. C.S. New Elphick, Springer-Verlag problems. E.N., wasp analysis. gall-forming Ieno, data a A.F., for by graphics Zuur, engineering elegant Ecosystem trees. ggplot2: (2016). oak (2016). L.H. on H. Yang, herbivores Wickham, & of density . degC. and . . diversity 2 on I. suppresses than effect Li, indirectly rather projected R.M., degC Screen, The 1.5 W.C., Wetzel, to (2018). warming J. VanDerWal, global & limiting N. of 791–795. 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