Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ar Lopez Laura framework conceptual a multiple stressors: on environmental research into behaviour animal Integrating er....eyfw fay eea atrshv mre rmmt-nlss(Crain meta-analyses 2, from Box emerged have and patterns 1 general Figure al. any, (see et if stressors few, .very multiple . . on years. 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Data may be preliminary. oigwt tesr,tu iliglre ietcsso tesr.Atraiey nml xoe ostress to in exposed less animals and Alternatively, reproduction stressors. into of more costs invest direct to larger sigmoidal) females yielding (e.g., cause nonlinear thus immune can strongly stressors, suppressed a this with had is investment, fitness coping also parental offspring investment female if reproductive particular, of coping higher In function limited. of had were that expe- costs resources (2007) when ( lizards higher systems al. lizards that suffer et tree found French female can and periods. in vitellogenesis females non-reproductive investment during energy, reproductive than manipulated metamorphosis of periods rimentally or abundance reproductive overwintering during energy an migration, stressors sufficient requires for with accumulating (e.g., reproduction or survival when stage for Similarly, or crucial size is disappear). threshold period habitats a ephemeral time to when given of growing a signs where in horizons, reserves and time development seasonal impaired than with mortality, and temperature elevated rather heightened show stressors to Przeslawski effects can exposure multiple sublethal following larvae Przeslawski with increased depression echinoderm 2013; 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Data may be preliminary. motnl,we raim ipret e aia,te ih aeadffrn e fsrsosthat stressors of set different a face might they habitat, new a to disperse organisms change environmental when (Crowley rapid Importantly, globally human-induced and habitats unprecedented time, natural the environments. or new following shaping distance prospective now, of is in unit true that payoffs per likely especially over cost be uncertainty long-distance both might great For the on often This fitness. depend and both mobility) (that expected success). to costs in (including the about relative escape transit time uncertainty distance increases 1) on about or substantial that uncertainty (Fig. depends involve is stores space there energy can in 1-3 adaptive in dispersal, dormancy escape investment is trade-offs front-end and of dormancy handle substantial costs dispersal a or as organisms Both for well need how dispersal as the Whether environments, on and alternative mortality 2017). depends that in Dahirel which factors and environment, & payoff, regarding (Levin current Bonte fitness time insights or 2009; net provides space Pena dormancy expected in la escape and/or to de dispersal attempt & adaptive should organisms of when evolution influence the drying on temperatures, extreme Theory 2009). address Fox to & (Thomas animals Jr. supply by or Hairston 1998)), food used 2003; (Chapman limited commonly development; a is suspended instead and EIT on by conditions, based 2014). dispersal dormancy (Smirnov or of stages case tolerance resting special of (a costs diapause the 2000)), avoiding (Danks (Humphries and (Parmesan torpor exposure Europe reducing into in involves entering locations (EIT) new of time occupy risk to in population change their Escape same climate the reduce anthropogenic in by dramatically migrants unsuitable to moderate-distance made (Rivrud and shown deer short- been to red have relative by (Sawyer hunters migrations migration human long by autumn environmental harvested undertake of anthropogenic being that driver by deer primary triggered mule a be individual was varia- also snowfall), large-scale can (e.g., predictable EIS tracking ther mammals, availability. often various food EIS, by temporary (Berg and exhibited of stressors patterns form migration, weather cyclical Seasonal tempora- can in common, environment. organism, organisms a tions an new is involves some substantial, a insects, (EIS) of 1-3), and to space form birds, trade-offs relocating in some permanently, Escape on or direct demands. or dispersal sections (the long-distance metabolic rily immediately preceding through in and time reduction the locally or long-term in faces space relatively organism discussed in an stressors are stressors environmental which of escape suite effects the indirect with stressors coping and to from alternative escape stressors. an increased to temporal As the exposure or condition; of spatial maintain cost Larger-scale to required indirect 4: stressors energy large ex- Trade-off the to a favor of in responses represent can stressors) bring physiological would selection to the in predators sexual stressors) to and to other risk, ornaments due exposure or high both mortality predators but in to (e.g. food invest exposure cost abundant their to increasing have direct diverting (e.g., responses sites risks strong males physiological some greater favor a in if taking selected investment then Alternatively, in reduced sexually can stressors. result of in to selection would cost investment posure sexual the scenarios reduced Strong at Such (An- even where success. stressors. traits, ornaments cliff mating to selected or fitness no sexually size a or into relative energy little suffer large limited in can either result males possessing can cases, Williams on traits these heavily & in depends (Stuligross 1994); often future dersson success the biased, mating in male males, fitness For became reproductive ratios in sex number blue reductions offspring reduced example, further Additionally, a For of 2020). and 2020). fitness. nesting likelihood Williams adult successful the & of and increasing (Stuligross probability offspring lowered produced both a to offspring via of exposure of fitness terms adult reproductive in then in cost reduction cliff), ditive indirect fitness a large ( over a bees falls orchard in survival result offspring can if (e.g., stressors (Domis survival offspring offspring average per reduces investment their reduce sometimes tal. et si lignaria Osmia 06.Frhroe o-irtr utryseishv oe vrvs ra fhabitat of areas vast over moved have species butterfly non-migratory Furthermore, 2016). tal. et 00.Freape h ne fhmnhnig ahrta h ne fsvr wea- severe of onset the than rather hunting, human of onset the example, For 2010). xoe orsuc iiainadtepsiieiialpi uee nad- an suffered imidacloprid pesticide the and limitation resource to exposed ) tal. et 03 esr20) omny(nldn iento restivation; 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Data may be preliminary. h nerto fbhvorl hsooia n iehsoydcsoshglgtteiprac fdifferences of importance the highlight decisions history life that and reality physiological a behavioural, organismal is of of it integration understanding The that to overall believe the option our we consequently the condition-dependent, complex, and, preserve stressors. or component is to each to state response stores of responses integrated often analyses energy overall our maintain is this guide to While time usefully need time. can or the or the space include space on must in in trade-offs depends 1-3 escape balances escape 4) trade-offs organism (trade-off in of the time how balance success or on adaptive that space depends in which given the escape situation Furthermore, In to possible trade- 1-3. 2). each decision history (trade-off in the life risks fitness Further, on taking versa. expected depend capacity by organism’s vice coping acquired and capacity physiological 3) be coping and physiological (trade-off only behaviour depends adaptive organism’s offs can the 1) scale, the often In larger (trade-off on that or trade-offs. depends stressors energy term which avoid other longer requires stressors, behaviourally turn, the the in should for to that, animal exposed decisions (tolerance) an being and of much options costs emphasizes how adaptive the on 1, scale, The on stressors. 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Anne thank We study. future communities natural for Acknowledgements of avenue dynamics promising and comprehensive outcomes structure and our the distinct timely how explain help qualitatively a of could investigations offer determine 1) empirical (Figure can effects and of stressor interactions theoretical context on Further affected framework the nature, ecosystems. these in the Figure and how effects, of communities and and behavioral interactions for position 5 and species Box trophic physiological affect in can their and stressors stressors, and strength, how stressors, multiple outline consider we to Specifically, to responses communities. on scope natural individual our stressors on expand those primarily we of focused 5 have impacts 2019). we Hellmann influence while & can Finally, (Bell generation future one the Coninck transgenerational in into multi-generational (De generations stressors With others 4. stressors to Box developmental multiple responses in or discussed to behavioural effects plasticity, genetic adaption stressor multiple identify of of help mechanisms trajectory hidden could or stressors speed present, multiple or the Cambronero past an to the affect shaping in response that that, in integrated factors adaptation constraints social genetic overall learning) and of the ecological (including mix Examining a shaped plasticity stressors. of multiple influence developmental to the response within-generation examine integrated to and be would transgenerational four studies and the future balance for organisms direction how fruitful in Another variation to relate traits here. phenotypic discussed of (Reale trade-offs (Sih suites histories in tendency; differences life animal exploratory individual in and how or differences 2008) boldness individual Stamps consistent aggressiveness, & of in importance (e.g., the syndromes in al. over for behavioural interest factors, mounting or environmental multiple been personalities to has respond there organisms by decade, how used in a variation responses understanding to behavioral regard alternative With and tradeoffs. trade-offs such stressor-related navigate additional to to organisms variables illuminate relevant environmental volumes to insights multiple large likely novel collecting and simultaneously behavior generate and in animal will advances on recent efforts data of these light of in in Furthermore, behavior conservation. and animal or ecology Beppler considering three among drugs, explicitly empiricists interactions pharmaceutical that scenarios higher-order among While suggests define to interactions to occur. (e.g. focus models Beermann conceptual can stressors our (e.g. developed more interactions experiments have extend into ecology must higher-order stressors of we outside of where fields numbers stressors, stressors, larger be- multiple increasingly acquisition primary of predicting incorporating energy two accurately are effects increased of than interactive means of of more including consequence strength involving understanding, a deeper and as a nature arising develop the to stressors, possibly However, primary risk) two 2). to time (predation (trade-off exposed haviour or are stressor organisms space third where in scenarios a ecological escape with on to primarily focused ‘forced’ have we be Here, may they that, directions do time research cannot or Future they 4). space (trade-off if in scale but (avoid physiological larger 1), behaviourally necessary a trade-off compensate the at term; still mount short of cannot potentially not the might speed simply can 2) in organisms the Organisms (trade-off energy, and energy exceed enough. more more might physiological quickly get with change this to of response even risks increase, of because more mix suddenly play, taking rate a levels of into notion stressor come the using the physiological local case, higher to cope, that build if In relative and both plasticity. but physiological to reversibility, stay behaviour, time have adjust and versus organisms slowly, and escape speed increase capacity levels organisms relative stressor If should their responses? 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Cue not are tolerances when co-tolerance (negative Co-tolerance: stressor Cross-tolerance: stressor another to Cross-susceptibility: model effect: Antagonistic interaction) effect: Additive terms of German Glossary Fellowship (NC 1: the 212 Box by Postdoctoral TRR funded NSF SFB partly a the was 1557836. of by supported grant PCT part was Foundation Fellowship. supported as LKL Postdoctoral was (DFG) addition, Chancellor’s Foundation MAG In Research Colorado Fellowship, AS. of to Endeavour University 1456724 Awards and IOS Australia grant Foundation an Science by National by supported was esr nomto nteevrnetta niae h rsneo tegho atclrstressor. particular a of strength or presence the indicates that environment the in information sensory : icee otymasb hc ordc tesrepsr.Daseeg rmarsrepool. reserve a from energy Draws exposure. stressor reduce to which by means costly discrete, A n niomna atrta a eaieeeto nogns’ testruhreduced through fitness organism’s an on effect negative a has that factor environmental any rsswe pce ipa orltdtlrne omlil tesr ttecmuiylevel community the at stressors multiple to tolerances correlated display species when arises h obndeeto tesr seult h u fec tesraoe(..n statistical no (i.e. alone stressor each of sum the to equal is stressors of effect combined the hr h hsooia epnet/ffc foesrso nrae h oeac oanother to tolerance the increases stressor one of to/effect response physiological the where ET copihsecp ysbtnilyrdcn ciiy eaoimadenergy and metabolism activity, reducing substantially by escape Accomplishes (EIT) ES copihsecp ymaso ipra oanwlocation. new a to dispersal of means by escape Accomplishes (EIS) h obndeeto tesr sgetrta htwudb rdce ne an under predicted be would what than greater is stressors of effect combined the h obndeeto tesr sls hnwa ol epeitdudra additive an under predicted be would what than less is stressors of effect combined the hr h hsooia epnet/ffc foesrso nrae h susceptibility the increases stressor one of to/effect response physiological the where hr-emrsos ooeo oesrsosivligimdaeenergy immediate involving stressors more or one to response short-term A 9 ³ .ILwsspotdb ainlScience National by supported was IYL ). 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Data may be preliminary. o :Hwbhvo a hp tesreet ncommunities on effects stressor (Mikheev patches shape habitat can outside behavior from temporally How parasites 5: or trematode trout Box spatially contracting rainbow its by of example, through For risk affected moving manifest. increased to be simply an interactions By can at for animal. animal time in focal an points the locations, ( close by sufficiently distant at use between stressors patch migrating separate themselves, spatial sequence stressors or the in the range with variation in variation timescales, home temporal temporal shorter by by much not over instead determined manifest exposure but stressor also of can magnitude effects relative carryover and studied, less far Though Hope challenges. 2013; future Croft to & respond (Edenbrow (Crino syndromes foraging behavioral including of behaviors of al. acclimation. variety function et of physiological a (Zavorka maintaining result impairments that affect cognitive a suggesting in maze, to as a result exploring organisms can in stress to errors under more changes made impactful but stressors fish ( future potentially acclimated different, minnows the to common responses warm-acclimated despite on instance, understood stressor For single well suggesting a less to impacts, exposure is developmental the of of effects carryover severity The the lessened (Horwitz exposure plasticity developmental phenotypic adults, beneficial in presence performance the decreased and in temperature performance elevated improved to (Schnurr in can life experience resulting developmental in physiology, such striatus, later or Indeed, altered. stressor(s) behaviour is the in that future such of changes the plasticity in permanent behavioral) stressor or to that physiological lead with (either Experience phenotypic experience development. via of during acclimation effect to experienced the lead stressors can for stressor pronounced a the particularly with in be predators may pursuit effects to Carryover response agricultural in survival an increased to which (Ferrari active, exposed future more later generally when are mortality environments disease- risk higher to and susceptible rates (Dinh more have growth pesticide bivalves lower them Carryover example, considerably makes stressors. for suffered which waves future systems; 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infiatyrdcdislcmto pe n aeo orc oaigdcsosfloigexposure following decisions foraging correct of rate and speed locomotion its reduced significantly tal. et tal. et 2015). 06.Bhvoa aroe ffcshv lobe bevd .. aplsfo high- from tadpoles e.g., observed: been also have effects carryover Behavioral 2016). s hlest vi rdtr,hwvr hnte opt o hleste are they shelters for compete they when however, predators, avoid to shelters use ) tal. et tal. et p CO 05,sca ewr oiin(Boogert position network social 2015), 09.Dmefl avepeiul xoe ofo iiainadheat and limitation food to exposed previously larvae Damselfly 2019). 2. hl xouet hs tesr uigdvlpetsilrsle in resulted still development during stressors these to exposure While tal. et tal. et 04.Freape h esoesahr species, hare sea keystone the example, For 2014). 2020). hxnsphoxinus Phoxinus 11 tal. et tal. et tal. et 00.Dvlpetlsrs a enshown been has stress Developmental 2020). 00 htmyitrc ihteaiiyto ability the with interact may that 2020) 04 Chaby 2014; a agrban oprdt cool- to compared brains larger had ) tal. et tal. et 98 uof20) Interactive 2008). 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Data may be preliminary. ad hnrmiigseiscncmest ucinlyfrti os tesreet edt eweakened be to (Jackson tend communities effects resistant stressor more loss, to this leading for levels, functionally compensate trophic levels (Galic key can across replaced trophic when not species across is remaining situations propagate groups when can In eliminated those hand, effects or impact. of numerical reduced functionality greatest and subsequently ecosystem the behavioural are the and have (Vinebrooke pronounced if degree stressors stressors pool, stressors the to community these sensitive those upon the particularly which to contingent from are upon likely groups species level(s) or are functional groups trophic and but functional the predict species as to of well difficult as co-tolerance species are and predator communities tolerance pest on of of effects abundance stressor the Multiple on temperatures rising of (Thompson (Marquis positive-effects stress limiting environmental by of temperatures effects negative (Goulson against al. herbivores reduced ecosystems mutualism on (via et buffer the numbers effects help plant of negative can reduce have mutualisms can beneficiaries natively, turn, health indirect colony in and including and, behaviour members, member pollination) foraging single honeybee other a on for on pesticides stressors costs cultural of large impacts have negative Furthermore, can (Hegland (Hegland III). interactions mutualism Figure mutualistic 2010, a of al. deterioration of et the Schweiger to 2009, leading al. rates et co-occurrence species lower (Hegland may mutualisms stress disrupt Miller indirectly 2010; and increase, by Ross directly to also ger & instance, likely can (Shears stressors for are multiple consumers prey, rates and intermediate Single of predation on ecology responses, effects top-down antipredator feeding stronger reducing and to (Rumschlag or behavior leading abundance activity the density- foraging food change and their periphyton behaviorally- disproportionately increasing increasing top-down conditions stressors and the these if predators reducing because contrast, by their mesocosms, (snails) of host pond effects (Breitburg intermediate in mediated ecosystems the pathogens entire of other trematode of populations into collapse increased favoured energy or dramatically individual divert restructuring herbicides the and the zine both risk, to reduced lead moderate al. with can and et feed this prey however, to benefit predators, prey keystone (Francis can allowing processes by stressors physiological stressors prey, and life-history of than effects harder combined predators and (Tylianakis hit studies stressors stressor multiple If from arise, absent will often interactions is synergistic ecology when of Thompson predicting aspect for com- fundamental essential dynamics, this is predator-prey but interactions as such species interactions, mediated (Miller species behaviorally alter mutualisms communities, however, stressors Importantly, and for these ramifications 2010). how petition corresponding Schmitz by the & mediated composition and (Hawlena largely elemental stressors, are the pool environmental alters to detrital it responses the because achieve behavioural reaching her- compo- flow, such to community litter energy stress, carbohydrates plant plant and to the digestible cycling non-processed nutrient changes rich, response and for only higher in consequences not with has herbivory instance, plants but selective For sition, select This food-web. balance. to the homeostatic preference and within diet behaviour and nutritional antipredator organisms their structure or of shift foraging community may role in for bivores shifts functional involve implications the often indirect stressors change to important that responses have Adaptive 5). can (Figure stressors function to responses Behavioural tal. et 98 Rumschlag 1998; 08.Frisac,a n-pi uuls a rtc lnsfo nieteet fincreasing of effects indirect from plants protect can mutualism ant-aphid an instance, For 2018). tal. et 00.Cagsi h iigadsailsnhoiiyo pce eaiua atrsi epneto response in patterns behavioural species of synchronicity spatial and timing the in Changes 2010). tal. et tal. et 2014). 09 Schweiger 2009; 2018). tal. et 09.Freape h obnto fognpopaeisciie n tria- and insecticides organophosphate of combination the example, For 2019). tal. et tal. et 00 atn&Ie 04.Freape h eaieipc fagri- of impact negative the example, For 2014). Ives & Barton 2010; 04 Thompson 2014; tal. et 12 09 White 2009; tal. et 08.Udrtnighwsrsosimpact stressors how Understanding 2018). tal. et tal. et tal. et 08.I tesr pcfial inhibit specifically stressors If 2018). tal. et 08 Dib 2018; 2016). 05 Tosi 2015; tal. et 00.O h other the On 2020). tal. et tal. et tal. et tal. et 09 Schwei- 2009; 07.Alter- 2017). tal. et tal. et 2014). 09.In 2019). 2004), 2008; Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. o()adfclognssfo )n edak )uidrcinlfebc n )b-ietoa feedback. bi-directional C) and feedback uni-directional B) feedback, no A) from organisms focal and sor(s) 2 Survival Stressors Figure the and potentially Fitness. by potentially influenced yielding Rate while sources are jointly Growth/Reproduction Mortality Assimilation them Mechanisms, and Other Survival Energy Coping or ameliorate linkages. Predation between and these and of on trade-off Risk act Stressors and activity both a Availability by the or Food increased creates physiological by 2). Activity or trade-off triggered behavioral rectangles, other. 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Data may be preliminary. fmlil tesr htaea es atal orltdi pc n ie loigtepi fsrsosto stressors of pair scenarios. the unique or allowing effects possible behavioral spatial time; experience of fewer and the can number space with organism the instead, in left an expands correlated If, which is respectively. greatly partially in it uncorrelated least time, scenarios tolerance. organism, completely at distinct or the are and be qualitatively space that of of escape shorter in stressors subset that is of multiple a stressor exceeds bar of is costs or the space this relative matches or avoid Note: temporal time stressor the to choices. the or either on choose (i.e., of spatial can organism depend extent the the organism will temporal When of the choice spatial that stressors organism), relative optimal than organism. two less The the the The of is (B) each of to but and (IV). available that with organism choice choices overlaps than time an optimal stressor behavioral in which a the over of escape of is scales large-scale), set extent bars) tolerance is the [dark] the so it to by determine (green it and (and likely more temporal operate long space), more local, experienced and very in is is bars) very is stressor choice [light] escape as not the duration However, this (pink is (i.e., is stressor and (as time. scale the scale (III), impractical spatial in when should spatial time greater is However, escape a space in its in over space). in escape invest and occurs in to escape than stressor organism short optimal the thus, rather very when become and, (II), optimal is can stressor low, be (e.g., it duration the are space increases, stressor tolerate space in duration to the in constrained stressor optimal avoidance If highly be of is (I). generally costs space, choice should habitat), and optimal time of the in patch be co-occur small that a stressors affects multiple it of combination a or 3. 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Data may be preliminary. ytebnfisascae ihlwrpeainrs n nrae oaigopruiis ovrey the Conversely, exceeded is opportunities. cost foraging this increased consumer, and the on risk cost predation physiological effects lower a mediated with predator has trait associated the also and in A density benefits stress stressor from the physiological consumer though by high the Even indirect cause releasing predator. may thus positive the hunting, example, has of and for activity A, which reduced Stressor predator) to producer. leading keystone a on a ( benefits indirect negative (e.g. interactions. and level consumer species a functional of for single benefits disruption a and affect outcomes disproportionately community complex to 5. Figure or space in behaviour-into-research-on-multiple-environmental-stressors-a-conceptual-framework avoidance of costs with on image4.emf area depends an scenarios to these stressor move effects. of file first inactive, stressor each Hosted the of in scenarios, vs. strengths least other) two active in relative the latter at be the co-occur with these if and (e.g., right), area In effects’ time decision (top an stressor left). optimal only or The ‘multiple (bottom stressor space space one cause in (S1) effects. or can carryover 1 co-occur time has decisions right), stressor in experienced space-use (top Here, co-occur space and neither organisms. correlated movement and or mobile not animal time right), are on in (bottom that effects’ stressors only co-occur ‘co-occurring multiple time (S2) have allows 2 this nonetheless present, stressor to not and space are and/or stressors the time when in time of periods into 4. Figure nmlbhvorcnrdfie‘utpesrso’seais hneet fsrsoscryover carry stressors of effects When scenarios: stressor’ ‘multiple redefine can behaviour Animal w yohtclseaiso o tesreet a rpgt costohclvl leading levels trophic across propagate can effects stressor how of scenarios hypothetical Two vial at available https://authorea.com/users/393011/articles/506749-integrating-animal- 15 cnroA) Scenario tesrAmay A Stressor Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. rlt .. aeo,M,Ntl,D,Witnhm ..&Ra,JCA 21) hni eea wariness general is When (2012). J.C.A. Read, types? & predator M.J. multiple Whittingham, avoiding D., in Nettle, favored M., G.M. Bateson, B.O., Lovett, Brilot, C.G., In: Jones, challenges. R.W., and ideas Howarth, science. ecosystem stressors: C.A., multiple commu- Hatfield, of amphibian effects J.W., in Understanding stressors Baxter, fish. Multiple D.L., and (2007). Breitburg, bullfrogs, M.C. contamination, Doyle, chemical & of E.E. effects Little, position. nities: network R.D., Semlitsch, social predicts M.D., stress Boone, Developmental metapopulations. in (2014). K.A. dispersal Spencer, condition-dependent Letters & Biology body D.R. of Farine, N.J., Evolution history. Boogert, life (2009). in E. Biology trait Pena, Evolutionary independent la of and Journal de central & a D. Bonte, Dispersal: (2017). M. productivity? Dahirel, 472-479. life-history & to linked D. traits Bonte, personality animal Evolution Are & (2008). Ecology J.A. in F. Stamps, Kuenen, world. & B.W., changing P.A. Kooi, a Biro, M., in Heijden, persistence der species van understanding G., Driessen, disperse: E.T., Kiers, M.P., E. Vargas, Berg, stressors. C., of McDiarmid, combinations C., three-way White, in Z.Y., and interactions Mao, Mechanisms E., Tekin, the C., Understanding Beppler, for Framework 50 Integrative Vol In: An Systematics, Plasticity. and Transgenerational (2019). of J.K. Consequences Hellmann, Multigenerational salinity, & manipulating A.M. experiment J.J. Bell, mesocosm Piggott, A velocity. C.D., flow Matthaei, communities: and macroinvertebrate L., sediment fine stream Ma, ant on S., and effects predators, Karnatz, their stressor V., aphids, Elbrecht, on warming A.J., of Beermann, effects indirect and Direct (2014). mutualists. A.R. Ives, & acute B.T. of Barton, effect the fish. heat: (1994). reef the M. coral Feeling Andersson, in (2015). interactions M.I. predator-prey McCormick, on & changes P.L. Munday, temperature Affects P., CO2 Domenici, Elevated Shared B.J.M., (2013). Allan, P.L. mutualism? Munday, Performance. & apparent Altered S.A. through or Watson, Interactions M.I., competition Predator-Prey McCormick, P., Apparent Domenici, B.J.M., (1998). Allan, J.D. Roth, cycle. populations & when R.D. predation Holt, P.A., Abrams, in each reduction on Line a effects community. net to plant stressor the due space REFERENCES represent of decreased B’ symbols composition Positive/negative and ‘plant the dispersal effect. of in seed stressor node. shift fruits the increased trophic fundamental to of from a size diet both to denotes benefits primary leading disperser thickness now A’ seed their B’ ‘plant key switches ‘Plant from A A’, A’. competition A’. lead ‘plant ‘plant ‘plant may of of of availability consumers, pollination diet the and from a activity effects foraging prefers top-down decreased ( which to increased leading producers. with pollinator on combined specialist effects A synergistic stressor negative of to effects negative direct Ecology 10. , es .. .. ) pigrNwYr,N,p.416-431. pp. NY, York, New Springer G). P.M., & P M.L., (eds. 5 1479-1484. 95, , eulSelection Sexual 3 361-368. 23, , e.Ftya J,p.97-118. pp. DJ), Futuyma, (ed. c.TtlEnviron. Total Sci. 2 1242-1251. 22, , Ecology rneo nvriyPress. University Princeton . 9 201-212. 79, , h mrcnNaturalist American The cnroB) Scenario ora fteRylSceyInterface Society Royal the of Journal 16 1-1,961-971. 610-611, , lsOne Plos tesrBdsut h esr aaiiyo a of capability sensory the disrupts B Stressor clgclApplications Ecological lb hneBiol. Change Glob. 8. , ucse,lmttos n rnir in frontiers and limitations, Successes, osr.Physiol. Conserv. nulRve fEooy Evolution, Ecology, of Review Annual 7,180-195. 179, tal. et 21) noeigemergent Uncovering (2016). tal. et tal. et 7 291-301. 17, , 6 587-598. 16, , 3. , 21) Multiple- (2018). 21) dp or Adapt (2010). 3 1-11. 13, , tal. et Oikos (1998). Trends 126, , Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. i,V,Prs ... oa .. oel,RL ajla ..(00.Boiest-eitdeet on effects Biodiversity-mediated (2020). disturbances. V.F. environmental Farjalla, of intensity & and R.L. type the Bozelli, on C.C., depend mosquito. Nova, functioning vector ecosystem A.P.F., a Pires, toxicity in the V., biopesticide variationmagnifies Dib, a temperature and Daily pesticide (2019). 33-40. chemical R. 659, a , Stoks, of & consisting L. amixture Janssens, of Daphnia T.T., two Inter- Tran, of physiology V., (2013). and Delnat, C.R. life-history sensitivity. Janssen, to carbaryl carbaryl & in insecticide L. differing the Meester, and clories De parasite magna bacterial M., a Jansen, of K.A.C., effects Schamphelaere, active De D.I.M., Coninck, insects. De dormant in habitat Dehydration Predicting (2000). (2019). A. H.V. Sih, Danks, & change. W.S. environmental Cuello, development rapid S.M., after Ehlman, during choice O., finches. exposure Spiegel, zebra Corticosterone P.C., in Trimmer, human (2014). P.H., task Crowley, multiple C.W. foraging novel of Breuner, a & effects on R. cumulative performance Ton, and improves S.C., Interactive Driscoll, O.L., (2008). Crino, B.S. Halpern, systems. marine & in K. stressors Kroeker, character- C.M., dispersal: Personality-dependent Crain, (2010). populations. A. structured Sciences Sih, spatially B-Biological & Society for Royal S. consequences Fogarty, and T., ontogeny Brodin, ization, J., Resource- Clobert, importance their J., and Cote, stressors (2017). ecosystem among L. Interactions Boyero, (2016). C.J. conservation. & Brown, in review. & A.M. E.S. a Darling, Tonin, I.M., stressors. Cote, scales: multiple R.T., facing multiple Abdala-Diaz, caddisflies at in A., variation trade-offs allocation trait Basaguren, Physiology thermal F., Environmental Lizard and Correa-Araneda, in Systems B-Biochemical telomeres (2014). Physiology and Comparative S.L. stressors of Chown, between Journal & association The S. Clusella-Trullas, (2020). M. meta-analysis. Szulkin, a & vertebrates: S.M. non-human Drobniak, M., Chatelain, threat. (1998). under R.F. foraging Chapman, improves adolescence individuals prepare during stress (2018). Stress early Does L. 37-45. future? (2015). Orsini, stressful V.A. Braithwaite, & a & Daphnia. A.P. A.M. for species Hirrlinger, Beckerman, keystone M.J., the T.A., Sheriff, on on L.E., Davidson, Chaby, ocean stressors multiple L., the of of Meester, impact the acidification De of and H., Predictability warming Marshall, of M.C., impacts Cambronero, Multistressor histories. condi- (2013). life nutritional R. invertebrates’ early Przeslawski, marine of & consequences (T Lifelong M. finches Byrne, (2014). zebra E.T. in Krause, performance multiple-stressor & learning M. modify on web. Naguib, interactions tions food O., Biotic stream Kruger, experimental (2017). V., an Brust, C.D. in Matthaei, trout & brown P.E. juvenile Jones, on effects R.K., . Salis, risk. A., predation Bruder, under foraging selection: habitat and Research use patch Ecology Vigilance, (1999). J.S. Brown, rceig fteRylSceyBBooia Sciences B-Biological Society Royal the of Proceedings ,49-71. 1, , h net:SrcueadFunction, and Structure Insects: The clg Letters Ecology nertv n oprtv Biology Comparative and Integrative 6,4065-4076. 365, , h mrcnNaturalist American The clg Letters Ecology 1 1304-1315. 11, , qai Toxicology Aquatic eipgaguttata aeniopygia 17 ora fIsc Physiology Insect of Journal 3 381-398. 23, , clg n Evolution and Ecology abig nvriyPress. University Cambridge . nmlBehaviour Animal 3,149-159. 130, , 9,619-632. 193, , ). lb hneBiol. Change Glob. 283. , ea.Processes Behav. 3 582-596. 53, , hlspia rnatoso the of Transactions Philosophical cetfi Reports Scientific nmlBehaviour Animal ,5103-5110. 7, , 6 837-852. 46, , 1 27-32. 91, , Oikos c.TtlEnviron. Total Sci. 0,320-326. 103, , 3 3882-3894. 23, , 8,5-21. 184, , 2,433-443. 129, , Evolutionary ,11. 8, , 105, , Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. adn .. evs .. oya,M,Pru,M,Kn,AL oi,SC 21) hieas Thrice (2015). S.C. anu- Tobin, and & dragonflies larval A.L. in King, interactions predator-prey M., modulate Perdue, temperature and M., Copper rans. Holyoak, catch! M.K., to Predation easy Reeves, Linking Mechanism M.T., Fundamental a Hayden, as Stress Physiological Functioning. (2010). Ecosystem O.J. to Schmitz, & multiple D. to Hawlena, responses behavioral understanding and stimuli. Describing multiple (2017). and S.E. Swearer, stressors In: & J.J. Piggott, Banks. R., Egg Hale, (2009). from York. stress J.A. New combined Fox, Press by & Academic driven N.G. declines Jr., Bee Hairston (2015). flowers. E.L. of Rotheray, lack & hardiness and C. cold corn pesticides, Botias, oriental between parasites, the E., Relationships of Nicholls, larvae D., overwintering (2001). Goulson, in K. contents Suzuki, acid amino & torpor. furnacalis. free M. Ostrinia and daily Hujikura, borer, glycerol and between H., and hibernation Outa, diapause, during Y., and reduction Sekine, temperature M., Goto, body and Physiology rate of Metabolic Review disturbance Annual human (2004). of risk: F. influence predation The Geiser, and (2018). radiation J.S. Brashares, uv nocturnality. & to wildlife N.H. response Carter, on C.E., salamander Hojnowski, Larval K.M., Gaynor, (2004). use. A. microhabitat Sih, impacts and quantifying & change Color J. up: add processing. Stacy, don’t species ecosystem T.S., things to and Garcia, When metabolism communities (2018). individual V.E. avian from Forbes, stressors changes & multiple V. pollution of Grimm, Noise L.L., Sullivan, N., (2009). Galic, A. Cruz, & C.P. interactions. Ortega, M.I. change. C.D., environmental O’Connor, Francis, rapid M.L., under Logan, rescue K.J., behavioral Kroeker, for K., selec- nities Alujevic, predator D.A., and Vasseur, rate S.B., predation Fey, S.A. alter Watson, communities. temperatures K., fish sea rising reef Corkill, ( and in M.I., trout acidification tivity McCormick, ocean brown J.L., of in effects Rummer, size Interactive P.L., egg Munday, of M.C.O., effects Ferrari, Maternal quality. (1999). environmental I.A. to reaction Fleming, personality & by of S. development comment the Einum, shape the killifish effects to mangrove genetic and the reply Environmental in (2013). (a traits D.P. Croft, systems an & freshwater M. makes Edenbrow, tropical limitation 2013). in food Descy, E. conditions under & Jeppesen, wave Amado B.W., climate Sarmento, Ibelings, heat different V.L.M., a Huszar, under A.S., to dynamics Gsell, combined Exposure J.J., Elser, to experiment. L.N.D., (2016). responses Domis, laboratory R. mechanistic process a Stoks, soil lethal: & and insecticide L. agricultural biomass Janssens, plant K.V., of Dinh, review temperature. quantitative and a CO2 K.S. of Larsen, rare: manipulations M.J., Hovenden, are F., effects Hagedorn, F.A., additive Dijkstra, S., Vicca, W.I.J., Dieleman, Ecosphere urn Biology Current 6. , Science mrcnNaturalist American ora fIsc Physiology Insect of Journal 6 239-274. 66, , 9 1415-1419. 19, , rpoeismarmoratus Kryptolebias clg n Evolution and Ecology lb hneBiol. Change Glob. 6,1232-1235. 360, , rceig fteRylSceyBBooia Sciences B-Biological Society Royal the of Proceedings r swtrBiology Freshwater clgclApplications Ecological lb hneBiol. Change Glob. Science 7,537-556. 176, , 4,10. 347, , 1 1848-1855. 21, , 18 ,38-47. 7, , 8 2211-2213. 58, , 7 157-165. 47, , . nylpdao nadWaters Inland of Encyclopedia Oikos 4 1055-1064. 14, , 8 2681-2693. 18, , lb hneBiol. Change Glob. 2,667-681. 122, , lb hneBiol. Change Glob. clg Letters Ecology am trutta Salmo 5 3110-3120. 25, , tal. et tal. et tal. et 6,2095-2100. 266, , e.Lkn,GE). Likens, (ed. 2 3361-3372. 22, , 21) Opportu- (2019). 21) Plankton (2013). 1 568-577. 21, , 21) Simple (2012). tal. et :nrsof norms ): (2015). Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. ia ..(98.Srs n eiinmkn ne h iko rdto:Rcn eeomnsfrom developments Recent predation: of risk 215-290. the pp. under PJB), In: making Slater, decision perspectives. & M ecological and and Stress avail- reproductive, food behavioral, (1998). and Temperature S.L. ectotherm. (2014). Lima, an M.I. environments. in McCormick, patchy assessment & risk in M.C.O. affect Ferrari, strategies ability M.D., Dispersal Mitchell, G.D.H., (1984). Lienart, A. Hastings, on & effects Biology D. Multiple-stressor Population Cohen, (2018). stage. R.A. S.A., life Paterson, Levin, and & taxonomy J. of Brodersen, Importance object C.D., novel fish: Matthaei, freshwater a A., on Bruder, performance K., not Lange, Early- but memory (2015). spatial S.A. and MacDougall-Shackleton, learning & stressors associative test. L. Environmental affects Garcia, stress S.D.T., (2013). nutritional Aitken, P. life T.M., Domenici, Farrell, & B., Kriengwatana, D.J. behaviour. McKenzie, and and physiology N.B., rate between Metcalfe, metabolic relationships between S., alter relationship Marras, The S.S., (2016). lunch? Killen, S.J. deprivation. free Fu, food a & by as Y.X. altered thing Wang, such is Q.Y., there Wu, sociability is C., Fu, traits: S.S., life-history Killen, on Hormesis (2013). V. Ducrot, & Ecotoxicology A. stressors Barsi, multiple T., of effects Jager, mammalian Net meta-analysis. in (2016). a availability C.T. Chimimba, ecosystems: energy & freshwater of R.D. in Vinebrooke, role C.J.G., The Loewen, M.C., Jackson, (2003). D.L. approach. Kramer, cost-benefit & A D.W. hibernation: Thomas, M.M., keystone Humphries, a UK. Cambridge, in Press, (1999). rate University J.M. bridge metabolic McNamara, and & A.I. locomotion, Houston, behaviour, foraging S. Hacquart, alter R., mollusc. acidification of Beldade, marine development J.C.A., and Pistevos, the warming S.A., disrupts Watson, ocean stress T., Juvenile Norin, fish. R., cichlid (2020). Horwitz, P.L. convict in Hurd, syndrome & behavioural S.C.P. communities. exploration-boldness Renn, prey an K.L., of Fjellner, structure B.V., the Hope, and competition, apparent Biology Predation, Population (1977). review. R.D. A Holt, stressors: H.R. natural Kohler, and V., chemicals Scheil, 3746-3762. environmental A., 408, of Duschl, effects G.J., between Oostingh, actions A.M., Bindesbol, M., Holmstrup, zooplankton: in of salinization Sciences freshwater effects. influence B-Biological to cascading The Society tolerance reducing Evolved and (2013). cross-tolerance (2019). acidification. B. R.A. trade-offs, ocean Relyea, life-history Gaylord, & to & D.K. larvae warming Jones, oyster A.D. W.D., Olympia climate Russell, Hintz, of does J.D., response Hosfelt, How the T.M., on (2009). supply Hill, food O. E., Totland, Sanford, & A., A.L. Hettinger, Bjerknes, A., interactions? Lazaro, plant-pollinator A., affect Nielsen, S.J., Hegland, Behaviour 2 263-270. 22, , 5,195-218. 152, , cetfi Reports Scientific 6 165-191. 26, , 197-229. 12, , 7,10. 374, , 10. , clg Letters Ecology hsooia n iceia Zoology Biochemical and Physiological ucinlEcology Functional oeso dpiebhvor napoc ae nstate. on based approach an behaviour, adaptive of Models lb hneBiol. Change Glob. nmlBehaviour Animal 2 184-195. 12, , 19 r nsi clg Evolution & Ecology in Trends ihadFisheries and Fish tesadBehavior and Stress 0 1358-1365. 30, , 2 180-189. 22, , 9 199-204. 89, , hlspia rnatoso h Royal the of Transactions Philosophical nmlBehaviour Animal 9 974-983. 19, , Biogeosciences 6 165-179. 76, , es olr P Milinski, AP, Moller, (eds. tal. et c.TtlEnviron. Total Sci. 8 651-658. 28, , tal. et 22) Near-future (2020). 6,95-102. 161, , 0 6629-6638. 10, , 21) Inter- (2010). Theoretical Theoretical Cam- , Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. amn .. edro,S,Mle-zy . i ..&Qn ..(09.Imn epnet tem- to response Immune (2019). oyster J.G. Pacific Qin, & species: bivalve X.X. three Li, galloprovincialis P., in abiotic Miller-Ezzy, multiple stress of S., perature effects Henderson, the of M.A., meta-analysis and Rahman, of larvae. review limitation and A embryos oxygen (2015). marine the C. on Mellin, through stressors & fishes M. marine Byrne, R., affects Przeslawski, change Climate (2007). tolerance. R. thermal Knust, may pollution & anthropogenic H.O. chemical Portner, of recognition. levels predator Low impairing (2016). by I. amphibians Gomez-Mestre, fish: threaten stress & environmental in P. of Burraco, responses coincidence N., Temporal Stress Polo-Cavia, (2012). (2019). B. rates. Helmuth, L. predation & Jacquin, J. modulates Casas, events & E., P. Sanford, Laffaille, S., Pincebourde, J., Cote, processes. evolutionary A., to Gandar, warming. molecular S., regional From with Jean, associated H. Q., species Descimon, Petitjean, butterfly C.D., of Thomas, J.K., ranges Hill, geographical C., in disciplines. Stefanescu, shifts research N., across Ryrholm, goals C., common Parmesan, Sciences B-Biological and Society divisions Royal C. stressors: Mantyka-Pringle, the multiple R.L., of of ceedings Kordas, K.J., study Kroeker, unified preda- M.C., a of Jackson, Towards effects of R.D., negative Vinebrooke, traits the J.A., life-history enhances Orr, consumer. change and intermediate Climate behavior an (2014). the on G.C. on risk Trussell, tion & effects C.M. wave Matassa, Heat L.P., Miller, (2020). M. can Czarnoleski, behavior & antipredator antlions. D. sedentary predators. Innate of Stec, absence (2020). K., the J. Miler, in Taskinen, even & fish biomedicine. A.Y. in and Morozov, infection biology promote A.F., in Pasternak, allostasis V.N., of Mikheev, a concept survive The to (2003). use Behavior J.C. animals and Wingfield, strategies & different B.S. the McEwen, copepods Reviewing marine of physiology: response Starvation challenge. food. metabolic common of (2010). The absence the M.D. (2015). in McCue, M.R. acidification Viant, ocean & and K.B. warming environmental Cook, to U., Sommer, food. and D.J., predation, Mayor, nest sites, nest to relation in evolution Monographs life-history Avian B.O. (1995). Clarke, fish. T.E. Martin, T.L., in Coggan, behaviour P.J., antipredator Lewis, compromises 592-599. M.G., fluoxetine Bertram, pollutant trophic M., psychoactive multiple Saaristo, on effects J.M., cross- warming buffer Martin, on mutualisms tolerance Insect cold (2014). S.L. for Pelini, selection & levels. I. of Toro, effects Del melanogaster. M., The Drosophila Marquis, in (2009). stressors B.J. environmental other Sinclair, to & syndromes. tolerance J.P. behavioural Walsh, adaptive H.A., Sciences dependent B-Biological MacMillan, state- Society and Royal resources the Risk, of (2010). Transactions A. sophical Sih, & B. Luttbeg, Ecology 5 101-127. 65, , 3 2-15. 43, , 5 9-13. 95, , n u cockle mud and Science eairlEcology Behavioral op ice.Pyil -o.Itg.Physiol. Integr. A-Mol. Physiol. Biochem. Comp. 1,95-97. 315, , aeyi rhytiphora Katelysia clg Letters Ecology . lb hneBiol. Change Glob. lb hneBiol. Change Glob. c.TtlEnviron. Total Sci. 5 680-688. 15, , 287. , 20 . ih&Sels Immunology Shellfish & Fish qai Toxicology Aquatic rsote gigas Crassostrea eairlEcology Behavioral 0 3834-3844. 20, , 1 2122-2140. 21, , 8,371-380. 684, , 6,3977-3990. 365, , 5,1-18. 156, , netScience Insect cetfi Reports Scientific eierna mussel Mediterranean , 7,30-35. 172, , 1 267-276. 31, , nio.Pollut. Environ. Nature tal. et 6 868-874. 86, , 6 263-276. 16, , tal. et 19) Poleward (1999). 9,579-583. 399, , 5. , tal. et 21) The (2017). Hormones Ecological Mytilus (2020). Philo- 222, , Pro- Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. tesr nboi neatos o lmt hneadainseisitrc oaetpollination. affect to interact S. species Klotz, alien P.E., and change Hulme, climate T., how Reviews persistent Hickler, interactions: has R., biotic development on Bommarco, stressors embryonic J.C., zebrafish. during Biesmeijer, of Temperature O., muscle Schweiger, the (2014). in G.R. enzymes land- Scott, metabolic of & on ecology effects community Y. a Yin, space. Toward M.E., geographic (2017). Schnurr, across B. interactions Abrahms, predator-prey & multiple A.M. predicting Trainor, scapes: J.R.B., mile: Miller, interactions. extra O.J., trophic Schmitz, The and diversity risk. Predator (2016). anthropogenic (2007). to K.L. O.J. exposure Monteith, Schmitz, and & range seasonal M.J. of Kauffman, use 7. the , M.M., alters Hayes, distance migration A.D., Ungulate Middleton, biostratigraphical H., - Finland. Sawyer, change southern environmental in of lakes indicators two as type from ephippia and evidence Chydorid class pesticide (2003). to K. generalised P.J. Sarmaja-Korjonen, be Hudson, can H.J., parasites to Carrick, communities. susceptibility T.R., aquatic and in exposure Raffel, on J.T., pesticides Hoverman, of integrating Effects N.T., model Halstead, and S.L., new interactions Size-structured Rumschlag, A dynamics: - predator-prey model on cannibalism scope mutualism. of apparent reactive Impact The (2008). V.H.W. (2009). Rudolf, N.E. stress. Cyr, on and & pesticide allostasis, a M.J. homeostasis, of Dickens, effects L.M., the Romero, reduce contaminants. change to climate susceptibility Will and exposure (2011). hydroperiod. 657-666. on C. and effects Stieha, partitioning limitation, & amphibians?: T.M. food Sesterhenn, herbicide, J.R., an Rohr, of J.H. Niedzwiecki, Effects T.M., McCarthy, salamanders: P.H., Applications Crowley, and B.S., stressors Shepherd, Multiple A.A., Elskus, J.R., Rohr, before (2002). Leave population. D. ungulate (2016). hunted Roff, a A. in Mysterud, migration & autumn meta-analysis. of L.E. a triggers Loe, environmental Ecology and stoichiometry: B., anthropogenic Zimermann, prey late: E.L., on too Meisingset, it’s risk R., predation Bischof, of I.M., Rivrud, effects The cause (2020). the D. and stressors Ecology Hawlena, Multiple & (2010). S. K.A. Rinehart, Trust, & M. Holyoak, abnormalities. C.L., amphibian tempera- Dolph, of animal P., Integrating Jensen, (2007). M.K., N.J. Reeves, Dingemanse, evolution. & P.T. and McDougall, ecology D., within Sol, Personality ment level. S.M., population (2010). Reader, the P.O. D., at Montiglio, Reale, Sciences concept & B-Biological syndrome V. Society Royal pace-of-life Careau, reproduction the P., the female of Bergeron, of on emergence M.M., disruptors the Humphries, endocrine and D., of Garant, impacts D., epigenetic Reale, The (2019). J.A. generations. Flaws, across & S. Rattan, 7 1058-1068. 97, , 101. , 5 777-795. 85, , 4 1028-1040. 14, , ieHsoyEvolution. History Life ilg fReproduction of Biology Ecology clg Letters Ecology clgclMonographs Ecological 9 1650-1660. 89, , omnsadBehavior and Hormones ilgclReviews Biological xodUiest rs,Ofr,UK. Oxford, Press, University Oxford 2 962-972. 22, , 6,4051-4063. 365, , 0,635-644. 101, , Holocene 0 423-440. 80, , 21 2 291-318. 82, , 3 691-700. 13, , ora fEprmna Biology Experimental of Journal 5 375-389. 55, , Ecology 8 2415-2426. 88, , Ecology hlspia Transactions Philosophical lb hneBiol. Change Glob. tal. et 8 2281-2292. 98, , 1,1370-1380. 217, , 21) Multiple (2010). tal. et tal. et Ecosphere Ecological Biological (2019). (2004). 17, , Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. a eWa,DB icmn .(00.Mlil lblcag tesreet npyolntnnutrient phytoplankton on effects stressor change global ocean. Multiple future is a interac- (2020). toxicity in E. species Litchman, acquisition and pesticide & change D.B. on Waal, Global de warming Van (2008). of D.A. Wardle, effect ecosystems. & terrestrial The J. in Bascompte, tions (2020). R.K., pipiens. Didham, R. Culex J.M., Stoks, mosquito Tylianakis, the & nu- in L. stages and developmental Janssens, pesticides between K.D., reversed Neonicotinoid Van, (2017). T.T., P. Tran, Medrzycki, bees. & honey R. in survival Cabbri, of Sciences reduce role F., synergistically The Sgolastra, stress sculpin: J.C., tritional tidepool Nieh, the S., in Tosi, Cross-tolerance (2005). non-additive for G.K. proteins. Iwama, cause arousals shock & interactions heat of P.M. Species Schulte, cost A.E., communities. and (2018). Todgham, on R.D. budgets stressors environmental Vinebrooke, energy multiple & Winter of M.M. effects MacLennan, (1990). P.L., J.M. Thompson, Bergeron, repro- bats, & bee brown wild M. little impair hibernating Dorais, additively stressors D.W., resource Thomas, and Pesticide (2020). N.M. damselfly. duction. Williams, a in & performance C. developmental Stuligross, shape stress predator-induced and stress dispersal? Oecologia Food for (2001). substitute R. dormancy Stoks, can mechanisms: reduction risk Multiple Letters insects. in (2006). R.E. stress Snyder, immune and desiccation (2014). to N.N. Smirnov, cross- temperatures and Llow Cross-tolerance relating (2013). prey. H.A. Biology cold: on Comparative MacMillan, predators the & multiple G. in of Salehipour-shirazi, talk L.V., impacts Ferguson, syn- Emergent B.J., behavioural Sinclair, (1998). of D. implications Wooster, Ecological Evolution & & (2012). G. Ecology J. Englund, Pruitt, A., & Sih, S. Fogarty, M., overview. Evans, integrative An J., dromes. syndromes: Cote, Behavioral A., (2004). Sih, R.E. Ziemba, Biology & of J.C. Review Johnson, Quarterly A.M., Bell, A., Sih, In: overview. 203-224. ecological impair and pp. evolutionary to NH, an communities.Hanover, interact aquatic lifestyles: on mixtures prey impacts indirect and Contaminant and Predators (1987). (2018). A. Sih, K.M. Parris, and amphibian. & larval 482-488. complex a 161, S.E. have in , survival Swearer, stressors and anthropogenic R., behaviours predator-avoidance multiple Hale, M., cascades: Toxic Sievers, reefs. temperate (2010). on America P.M. effects of interactive Ross, States unanticipated & M. United the N.T. Stegman, of Shears, K.F., Sciences ectotherms. of Dilliplane, of Academy J., performance National thermoregulatory Borchert, the determines of landscape M.S., thermal Schuler, the of M.J., Configuration Angilletta, M.W., Sears, ,1106-1114. 9, , clg Letters Ecology rceig.Booia sciences Biological Proceedings. 284. , 2,222-229. 127, , hsooia n iceia Zoology Biochemical and Physiological 3 350-355. 13, , 3 545-556. 53, , h hsooyo Cladocera of physiology The 5 278-289. 15, , 9 241-277. 79, , hlspia rnatoso h oa oit -ilgclSciences B-Biological Society Royal the of Transactions Philosophical ytslucifugus Myotis clg Letters Ecology 8,20201390. 287, , es efo,W i,A.Uiest rs fNwEngland New of Press University A). Sih, & WC Kerfoot, (eds. 1 1351-1363. 11, , . ora fMammalogy of Journal cdmcPes Amsterdam. Press, Acadamic . 22 clg Letters Ecology 8 133-144. 78, , Ecosphere rceig fteRylSceyB-Biological Society Royal the of Proceedings ctxclg n niomna Safety Environmental and Ecotoxicology 3 1149-1159. 13, , 1,10595-10600. 113, , 9. , c.TtlEnviron. Total Sci. 1 475-479. 71, , rdto:direct Predation: 1,10. 717, , nertv and Integrative tal. et Proceedings rnsin Trends 7,8. 375, , Ecology (2016). Posted on Authorea 1 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161217752.23254903/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. aok,L,Kek . rsrn,TA,Sgni . rse,A iln ..(00.Rdcdexplo- Reduced (2020). S.S. Killen, & A. minnow. common Crespel, warm-acclimated in M., increase Biology Soganci, volume brain T.A., despite Armstrong, capacity ration B., Koeck, synthesis. L., global Zavorka, a F.Z. Wu, stoichiometry: Z.J., C:N:P Li, environmental terrestrial Y., abiotic Peng, on W.Q., of drivers Yang, impacts D.A., the Fornara, K., stress: Yue, of and ecology groups. warming the functional of and across processes effects factors. Ecological vary Combined (2013). and J.C. predators (2018). Wingfield, by N.E. moderated O’Connor, are & Biol. communities M.C. Change perfor- marine Emmerson, swimming on and I., nutrients development Donohue, history L., life fish. early White, pelagic the circumglobal on S.M.J. large acidification Pether, a than D.M., co-tolerance. of Parsons, effect mance species S., greater Nicol, of a D.E., has role McQueen, warming the B.J.M., Allan, functioning: S.A., S.C. Watson, ecosystem Maberly, and S.I., biodiversity Dodson, on M., stressors Oikos Scheffer, multiple J., of Norberg, Impacts K.L., Cottingham, activity. S. R.D., of Daan, Vinebrooke, J., timing Scheepe, daily J., the Akkerman, shifting S.J., by Riede, P., survival Tachinardi, levels. V., trophic re- Vinne, across abundance der interactions and Sciencesvan biotic B-Biological distribution include Society species to Royal essential Predicting is the (2010). it of M.E. why Transactions Visser, change: & climate M. to sponses Macel, W.H., Putten, der Van 0,451-457. 104, , 223. , ucinlEcology Functional 4 5853-5866. 24, , 7 37-44. 27, , lb hneBiol. Change Glob. clg Letters Ecology 23 6,2025-2034. 365, , lb hneBiol. Change Glob. 4 4368-4385. 24, , 2 2097-2102. 22, , tal. et 21) ffcso he lblchange global three of Effects (2017). tal. et 3 2450-2463. 23, , ora fExperimental of Journal tal. et 21) Maximising (2019). 21) Ocean (2018). tal. et Philosophical (2004). Glob.