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Trait-based diet selection: Prey behaviour and morphology predict vulnerability to in reef communities

Green, S. J., & Côté, I. M. (2014). Trait‐based diet selection: Prey behaviour and morphology predict vulnerability to predation in reef fish communities. Journal of Animal , 83(6), 1451-1460. doi:10.1111/1365-2656.12250

10.1111/1365-2656.12250 John Wiley & Sons, Inc.

Accepted Manuscript http://cdss.library.oregonstate.edu/sa-termsofuse BC CanadaV5A1S6; This article is protected bycopyright. reserved. All rights 10.1111/ differences between thisversion and the Version Record. of Please cite thisarticle as doi: through the copyediting, typesetting, pagination for publication accepted This articlehasbeen J.Green Stephanie vulnerability to predation in reeffish communities Trait-based diet selection: Prey Running headline: Ecology : Section Boutin Stan : Editor type Article : Standard Paper Date Accepted 08-May-2014: Revised 30-Apr-2014 Date : 06-Feb-2014Received : Date Tel: +1(541)908-3839 Email: [email protected] * b a Department ofZoology,OregonStateUn Department ofBiologicalSciences,SimonFraser AcceptedCorresponding author: Article 1365-2656.12250 1365-2656.12250 Prey traits determine Prey predation vulnerability a,b*

and Isabelle M. Côté

behaviour andmorphology predict iversity, Corvallis, Oregon, 97331-2914 USA Corvallis, Oregon,97331-2914 USA iversity, and undergone full peer review buthas notbeen undergonereview and full peer a and proofreading process, which may lead to University, 8888 University Drive,Burnaby University,8888University resting on or just above reefs are themostvulne are resting justabovereefs onor important determinantsof diet including positioninthe body size, body shape, 4. large. comparisons ofprey inlionfishstomach contents toavailabilityreefsin invaded at 1) of diet selection by lionfishwithin of dietselectionby models that account for relatedness among prey taxa prey among relatedness models thataccountfor species that could facilitate ordeter Constructinggeneralised predation by lionfish. mixed- linear aboutmorphologi hypotheses firstgenerate 3. We Atlantic. fishcommunitiesan reef preys uponspecies-rich ( Indo-Pacific by lionfish preyexamining selection vulnerability topredation independent ofspecies identity. apply We thistrait-based approach to and general morphological based onidentifying This article is protected Allrights bycopyright. reserved. 2. interactions novelpredator-prey for predicting spatially and temporallyvariable.a result, species-specific As selectivity data are oflimited use often is predator a to available ofprey species suite the However, anddynamics. structure 1. Summary Understanding how predators select theirprey select howpredators Understanding We present a method for predicting dietselecti for predicting presentamethod We Both analysesreveal that a number predicted oftraits toaffect vulnerability topredation, Acceptedin situ Article visual observationsand of prey availability consumption for individual lionfish,and (2) selection by lionfish. Small,sh twoindependentdatasetscollect because they are assemblage-specific. assemblage-specific. they are because rable. that exhibit parasite cleaning parasitecleaning Fishes thatexhibit rable. and aggregation behaviour, are behaviour, are aggregation water columnand behavioural traits of prey that confer confer ofprey that traits behavioural d is rapidly spreading across the Western theWestern across israpidly spreading d can provideimportantinsightscan intocommunity cal and behavioural traits recurring across fish across recurring traits cal andbehavioural on that is applicable across prey assemblages, prey assemblages, on thatisapplicableacross Pterois volitans , we test whether these , wetestwhetherthese

allow-bodied solitaryfound fishes ), an invasive predator that ), aninvasivepredator ed at different spatial scales: scales: spatial different at ed traits predictpatterns predators should select prey items predators shouldselect 2004) and Webster etal.2001, Almany 1995, Juanes (Gilinsky 1984, and functionofcommunities structure the into key insights provide therefore can prey their select predators which process by Limaet al et al.1996, 1995,Arcese and Hixon ofspec rates thedemographic Predation affects Introduction volitans/miles and availability, selective predation, stomach contentsanalysis, Key-words oncommunity dynamics. interactions predator-prey ofchanging outcomes the predict to and species predator useacross beof therefore availability,gatheredare predator typically which species. could more for Atrait-based approach yieldobservations ofselection thatarerobust interactions predator-prey This article is protected Allrights bycopyright. reserved. 5. 200. factorofnearly a by the riskofpredation heighten traits vulnerable Together, risk. greater significantly at are active arenocturnally that riskof lower asignificantly behaviour experience

Accepted Article atrait-ba that reveals Ourstudy behaviour, predator-prey inter predator-prey foraging behaviour,

similar results to broad-scale comparisons of prey useand comparisonsof similar resultstobroad-scale in a manner that maximizes en maximizes inamannerthat sed approach to studying diet selection yields insightsstudying into dietselection to approach sed across prey assemblages. Importantly,assemblages. preyacross Holt 1984, Kotler 1984, Karanth andSunquist 1984, KotlerKaranth Holt . 2001, Creel et al. 2007). Understanding the Creeletal.2007).Understanding . 2001, ies in mostecosystems ies predation than non-cleaning fishes, and fishes fishes, predation thannon-cleaning actions, prey characteristics, use resourceuse characteristics, prey actions, . predicts that predicts theory . Optimalforaging

in situ ergetic gain whileminimizingergetic gain (Caley etal. 1993; Carr observations, in situ Pterois additions (i.e., through invasion; e.g. [Caut etal e.g.additions (i.e.,through [Caut invasion; characteristics that affect thei thataffect characteristics va types predation isthought when prey tooccur ofpredator-pre nature Theheterogeneous predicting selection diet can changingapplied that be across community Selectivecompositions. findings forpredicting selection limiting andSoulé1999]), theu e.g. [Crooks and,in Phillips etal.2009, et al.2010]), Jaeger rang thepredator's across prey assemblage ow variable, andtemporally predator isspatially th Isaacet al2013).However, et al.2012; 1999; Wen et al. 2006), and a smaller number et al.2006),andasmaller 1995,Einfalt&Wahl 1997;Nilsson& Wahl Br se in experimental choose species predators areselectively that species prey Identifying onwhich focussed studies of number large a with research, ecology foraging ofmuch aim the al. 2007;Nunnetal,2012). 1995,Almany et Hambright 1991,Hernández Osborne 1981,Sco and predation; e.g.Green some prey inquantitiesdi across space and time (e.g., Beukers-Stewart &Beukers-Stewart J andtime(e.g., across space in the environment, sodiet composition willtrack changes community inprey composition shoul thenpredators profitable, equally types are This article is protected Allrights bycopyright. reserved. Accepted Article the energyexpended inlocating, and processing handling item (Pyke the sproportionate to their availability (i sproportionate totheiravailability r encounter rate and handling time (Pyke et al. 1997). It al. 1997). therefore handling et time(Pyke and rate r encounter from varying sets of prey. of prey. from varying sets focussed on species selection onspecies focussed e [e.g., Holt 1984,Steele1989,Boste [e.g., etal.2002, ttingsStein 1988,1989, & (e.g. 1985,Wahl Main tt and Murdoch 1983,Doná tt and tility of experimental and assemblage-specific and assemblage-specific tilityof experimental creasingly, anthropogeni creasingly, . 2008]) andremovals (e.g . 2008]) onmark 2000; Heinonen & Auster2013;Provost 2000;Heinonen onmark ing both to natural processses (e.g., gradientsing bothtonatural (e.g., processses in ry inoneormoremor ry d consume prey in proportion to their abundance totheirabundance d consumeprey inproportion ones 2004). However, pr ones 2004).However, y necessitates interactions approach an to consumed (or, conversely, avoided) has been hasbeen avoided) (or,conversely, consumed e assemblage of prey available to a given available toa e assemblage ofprey

.e., dietary preference or selective orselective preference .e., dietary in situ phological or behavioural orbehavioural phological (e.g. Cunningham etal. (e.g. cally mediated callyspecies mediated zar and Ceballos1989, zar et al. et . through exploitation; edators oftenconsume edators 1977). If 1977). allprey items inlionfishstomach contentstopr individuallionfish hunting.consumed by Atalarg observationswesimultane visual during underwater collected data from occurs variants trait specific could facilitateassessed predation ordeter by lionfish,and selective whether predation on with novelfishassemblages. Todothis,we areas are mostsusceptible topredation willaid inforecasting prey declines as theinvasion spreads to Given that reef fishassemblagesacross large differelucidating region, this trait prey that variants 2008; Green (AlbinsandHixon in thenorthernCaribbean Predation by lionfishhasbeenimplicated invasive Indo-Pacific lionfish( to predationby the Toillustrate a trait-based approach dietselection, tostudying wewhether evaluate char and behavioural morphological of studies that have tested thisidea in pred community more ecology a meanstomake identity. fromspecies independently Afunctional, typesby predicted could identifying be traits associatedto predation, withvulnerability species assemblages. the strength ofinter Thus, This article is protected Allrights bycopyright. reserved. Acceptedand theinvasioncontinuesto thewest across reefs coral become establishedon Article morphol by predation isconferred follows thatdietselectionshouldbepredicta spread southward (Betancur-R (Betancur-R spread southward ogical and/or behavioural tra and/or behavioural ogical acteristics of Caribbean coral r Caribbean coral of acteristics the context of predator–prey interactions. interactions. of predator–prey the context ey abundance on invaded reefs at large. at large. reefs on invaded ey abundance Pterois volitans ble across prey assemblages vulnerability prey if to across ble ictive (McGill et al. 2006), but we are not aware aware butwearenot (McGilletal.2006), ictive actions when a predator is exposed tonewprey isexposed whenapredator actions at two spatial scales. At the smallest scale, scale, smallest the At scales. spatial two at generated hypotheses about prey that about hypotheses traits generated ern Atlantic, Caribbean er scale, we compared the abundance of prey ofprey abundance compared the we er scale, in the significant ofreef decline fishbiomass or trait-based approach approach or trait-based ously recorded prey available toandprey available prey ously recorded and

et al. et et al. et P. miles its of prey that recur across across thatrecur its ofprey 2012; Green etal. 2012;Green 2011; Côté etal.2013). 2011; eef fishes affect vulnerability fishes affect eef ). Lionfish haverapidly ). and Gulf of Mexico, GulfofMexico, and has been advocated as beenadvocated has in press ). This article is protected Allrights bycopyright. reserved. reduce recognised search We time. predator th prey, target schooling may fishspec infact hypothesized thatlionfish,whoseslowmovement aggregations dilu can because predator strategy anti- effective an is schooling behaviour that suggest many studies While accessible. least column above the water fishes occupying pelagic liv those (i.e. species benthic than that demersal< 2mfromthe (i.e. bottom) living shouldconfervariable behaviour aspects ofprey ofpr theeffects To simultaneously examine PREY TRAIT HYPOTHESES methodsMaterials and Accepted& Maljkovi the substrate(Côté by slowly over hovering thatemploy astalki Lionfish visualpredators are Behaviour Article Deloach 2002). & (Böhlke Ch species descriptions ofeach on encountered toeachfishspecies each trait fo hypotheses generated 1),and (Table predation selection, we compiled on a suite information that oftraits are likely toinfluence vulnerability to

ing on or in the ), which may be less visible,andthat benthos), whichmaying beless onorinthe aplin 1993; Deloach 1999; FAO 2002a,b;Humann& Deloach1999; FAO aplin 1993; ies because the conspicuousness ofschoolsmay theconspicuousness ies because ey morphology and beha ey morphology our study reefs (Appendix S1), basedonpublished S1), (Appendix reefs our study ree categoriesree aggregation of behaviour: solitary, te individualriskofpr r each. We then assigned a score or value for value a scoreor assigned then each.We r ng strategy, in which they approach prey inwhich they strategy, ng their fishes would be more vulnerable to predation topredation wouldbemorevulnerable fishes vulnerability tothishunting predicted We style. reefs (i.e. > 2 m above the benthos)wouldbe > 2mabovethe (i.e. reefs s afford them a close approach to potential potential to approach a close s affordthem ć 2010;Green

viour on lionfishdiet viour edation (Pitcher 1993), we (Pitcher 1993),we edation et al. et 2011).Several as the linear distance from the pelvic girdle to the dorsal ridge, as in Hambright (1991), from as in Hambright dorsalridge, (1991), tothe thepelvic girdle from as thelineardistance measured cm), (in body to TL depth offish ratio average the calculated We shapes. vulnerable, thathighervaluesindicate body depth,such qua We round)shapes. (i.e. increasingly deep-bodied selection bygape-limited lionfish,withvulnerability decreasing topredation withincreasingly thusanticipatedthatpreyBrönmark b 2000).We with increase andhandling) time forcapture cost gape-limited predators,the by lionfish.For consideredWe aspects three major morphology ofprey that may affect vulnerability topredation Morphology pr cleaners service predators ofthe predati from rates mortality lower experience cons maythroughout lessreadily be theirlifespan), cleaningthat exhibit either behaviour, facultativelyas juvenile) obligately(i.e. only or (i.e. This article is protected Allrights bycopyright. reserved. Accepted Articlelionfish (Hobson1973;Fishelson 1975;Green th because diurnal counterparts (Green times crepuscular from shelteredreef overlaps withth whoseemergence and active arenocturnally that fishes that hypothesized also We parrotfishes). that many fishspecies tendtoformloose feed asan definedshoaling shoalingWe andschooling. et al. et 2011) would be more vulnerable to 2011)wouldbemorevulnerable is strategy may result in increased encounters withhunting inincreasedencounters result is strategy may ovide (e.g. Côté 2000;Colin1975). (e.g. Côté ovide e timing of lionfish foraging behaviour at behaviour e timing oflionfishforaging less round (i.e.moreelon less round prey body depth (Hambright 1991; Nilsson& 1991; (Hambright bodyprey depth on, possibly because of recognition byfish recognition of on, possibly because ing aggregations asjuve ing et al. et s of prey consumption (in terms of energy and ofenergy s ofprey consumption(interms ody shape plays an important roleinprey plays animportant shape ody intermediate state, based ontheobservation intermediatestate,based umed by lionfish, because such species often suchspeciesoften umed by lionfish, because 2011). Finally, we anticipated thatfishes we anticipated 2011).Finally, ntified shape as the ratio of TL to maximum ofTLas theratio tomaximum ntified shape

lionfishpredationthantheir gate), and hence more andhence gate), niles (e.g. and wrasses (e.g. niles using the allometric Massa*TL(cm) function (g) scaling = convertin by g),which weobtained vulnerability topredation woulddecrease withincreasing prey size, quantified as body mass (in 1997;Nilsson&Wahl & Br by (Einfalt predators and sm predators evading at better are prey larger selectively that consume are prey gapetheir smaller maximum far than limits,likely because to tend predators gape-limited that evidence experimental ample is there However, reefs. invaded thisresultsimply whether It 2013). isunclear (much smaller than the maximum of~48%(Morris& Akins 2009;Munoz (much smallerthanthemaximum across severa oftheirlionfishpredator the size Schluter 2003;Schubert on shallow coral patch reefs off Eleuthera Isla offEleuthera reefs on shallowcoralpatch predation compared with theirunde with predation compared compound,shar ordistasteful atoxic (e.g. defense that wehypothesized et al.2012).Second, Green To quantify lionfish prey selection To quantify lionfishprey SELECTION OFPREY DIRECT OBSERVATIONS constants thatrelatelength This article is protected Allrights bycopyright. reserved. Accepted Article rangethat asize with were that for specimens enc fishspecies ofeach [www..org]) specim lateral images of3-5 et al. et to weight obtained from FishBase (www.fishbase.org). . . FishBase (www.fishbase.org). from to weight obtained ens (available in Humann & De Humann ens (availablein 2003). Finally, the average size of prey (total length) to relative (total ofprey size average the 2003).Finally, in situ g estimates of total length (TL to the nearest 1 cm) to weight 1cm)toweight lengthg (TL tothenearest estimatesoftotal fended counterparts (e.g. Hoogland Hoogland (e.g. counterparts fended , we conducted detailed visual , weconducted ountered on the study reefsimages were only used onlywere used reefsimages onthestudy ountered could be consumed by beconsumedby could nd, The Bahamas (22º22.500 N, 76º49.000 W), in N,76º49.000 W), The Bahamas (22º22.500 nd, reflects the abundance of various prey on various sizes of theabundance reflects l studies oflionfishstomachcontentsis~15% l studies fishes possessing a chemical or physical orphysical chemical possessing a fishes aller prey are less costly to capture and handle and handle capture costly to are less prey aller önmark 2000). We therefore hypothesized that hypothesized therefore önmark 2000).We p spine(s) or barb) woul p spine(s)orbarb)

b , where a and b are species-specific arespecies-specific aandb , where loach [2002] andonFishBase loach [2002] lionfish (i.e.<15cmTL; observations oflionfish observations et al. et d belessvulnerableto d et al. et 1956;Caley and 2011;Côté et al. et

gapeset by (Nilsson& size Brönmark2000). prey items,basedon than 15cmTL aspotential ledges) orunder those increvices thusassumedthatonly We sand/ prey beds. predatorsthat primarily are visual Lionfish capture. as well aswithin a the 1mradius fishes lionfishimmediately ofthe focal following each prey recorded the identitysize andofall prey items conti number offishprecluded radius ofthehunting every 5min(hen lionfish (totallengthand size to [TL] identity, abundance lionfish (total length tothe nearest [TL] 1cm). focal ofthe size the weestimated ofobservations, onset the study. At this in were included cons stalkingfishes) and prey thatactively were hunting (i.e. oriented with m (asperGreen of2-3 30 minby fromadistance aSCUBA diver (Côté & Maljkovi prey at ofstrikes success asthe as well prey, target ofthe view easy observers affords which This article is protected Allrights bycopyright. reserved. Accepted ArticleMaljkovi fac which little fearofdivers, reef 2009.Oninvaded September 2008andDecember ć 2010;Green ć 2010; Green 2010; et al. 2011).They strategy, aslow,hovering, stalking huntprey via ilitates a close approach without approach ilitates aclose umed at least 1 prey item during the 30 min. observation period item during the30min.observation umed atleast1prey nuous tabulation of prey across the 30 min period. We also the30minperiod.We nuous tabulationofprey across et al. et were available for availablefor consumption. were We 2011). Individual lionfishwere 2011). head down, pectoral spread andactively finsspread down,pectoral head consumed by lionfish during the30min.period, by lionfishduring consumed ce, six point countsperobservation).Thelarge six ce, During each observation period, we recorded the werecorded period, observation each During physiological limitsonpredator: prey size ratio the nearest 1 cm) of all fisheswithin a1m 1cm)of thenearest hunt in the open over the top of coral reefs or reefs topofcoral the hunt intheopenover within visual sight ofthe lionfish(i.e. not s, lionfish exhibit bol s, lionfishexhibit

altering behaviour (Côté& behaviour altering et al. et observed continuously for continuously for observed classifiedallfishesofless 2011).Only individuals d behaviour and have d behaviourandhave This article is protected Allrights bycopyright. reserved. Accepted fishes ofprey-sized Transectsurveys observed onour fishes prey-sized of assemblage the that assumed We site. each at collections surveyswould inourvisual and notrecorded framework reef the within preyhidden any that assumed We 1cm. nearest the to recorded were ligh using adive fishes, cryptic for and crevices a transect Along each locations. capture lionfish to available ofprey Toestimatetheabundance on6-12belt cmTL) <15 fishes(i.e. prey-sized or viaDNAbarcoding). lionfish stomachs fromwhich all ofthe fishitems could be identified tospecies (either visually (Côté analysis barcode DNA via identity species weobtained colouration, and pigments of key skin thatcouldnot could beestimated) for whichTL Articleprey itemsvisually lowest tothe fish all identified and eachspecimen from contents stomach extracted 1 cm, nearest TL the to measured then We surface. the at solution andseawater oil aclove in andeuthanized hand nets, were conductedat depths identified during ofthe site visual same surveys (24º59.072N, Island,Bahamas New Providence sitesal eight stomachs oflionfishcollectedfrom co byprey selection alsoinferredlionfish We SELECTION OFPREY INDIRECT OBSERVATIONS et al. et 2013). To minimize potentialidentification 2013).Tominimize of 10-20 m between May 2008. of10-20mbetween and July taxonomic resolutionspossible. taxonomic were conducted were conducted immediat mparing the abundance of prey items inthe ofprey mparing theabundance not be available to lionfish to consume. not beavailabletolionfishconsume. transects (30 m x 2 m) at each of the eight eight ofthe each at 2m) x (30m transects s. Lionfish collections and prey visualsurveys s. Lionfish and collections t as needed. The identity and TL of all fishes fishes all TL of and identity needed. The as t 77 º 32.207 W), to the abundance of prey totheabundanceofprey 77 º32.207W), ong a continuous system southwest continuous coralreef ong off a be identified to species because of degradation ofdegradation because species to be identified trained observer carefully searched in all holes holes all in searched carefully observer trained lionfish, we conducted lionfish,weconducted bias, we only included in our analyses inouranalyses bias, weonly included

For whole fishpreyFor (i.e.items whole Lionfish were collected using collected using were Lionfish ely prior to lionfish ely priortolionfish detailed surveys of detailed This article is protected Allrights bycopyright. reserved. Accepted (www.marinespecies.org). cross-checked from FishBase(www.fishbase.org), on speciesbased study classified region, we (Sodhi byamong fami specifying fishes prey relatedness Jennings (e.g. are thusnotstatistically independent tra morphological andbehavioural 2012). Because Team Core R 2012; al. et (Bates R software statistical forthe lme4 package the in function dir (i.e. set data foreach models created We pr and behaviouronlionfish models(G linearmixed-effects used generalised deviatedsignificantly morphology andbehaviour consump whether the lionfish predationtested Articleanalys Our totheirabundance. types inproportion ournullex selection, ofprey Inabsence the STATISTICAL ANALYSIS that suggested limitedmovementonthest and prelim Morris, unpublisheddata) basedonlab- at thesiteswithin~24hofcapture, theasse matched transect surveys et al. et 2012). In the absence of a complete molecu a complete In of absence 2012). the ey selection from our directandindi from our ey selection mblage of prey fishes availabletolionfishduringmblage fishes bouts hunting ofprey inary results from external tagging and tracking oflionfish tracking taggingand results fromexternal inary udy reefs (S.J. Green, unpublished data). Green,unpublisheddata). (S.J. reefs udy pectation wasthatlionfishshouldconsumeprey ect and indirect observa indirect and ect their most recent taxonomic groupingsmost recenttaxonomic obtained their tion by lionfish of prey types varyingtion by types lionfishofprey in LMMs) to assess the effects of prey morphology ofprey morphology LMMs) toassesstheeffects derived times to digestion for lionfish prey (J.A. derived timestodigestionforlionfishprey (J.A. ly as a nested random effect inourmodels effect a nestedrandom as ly from this null expectation. Specifically, we Specifically, from thisnullexpectation. es of direct and indi es ofdirect its can be shared among related species, and relatedspecies, among its canbeshared with the World Registry of Marine Species Registry Species with theWorld of Marine et al. et 1999), we accounted for phylogenetic forphylogenetic 1999),weaccounted lar phylogeny for the reef fishesinour the reef for lar phylogeny

rect observations ofpredation. observations rect tions) using the glmer() thetions) using rect observations of rect observations proportion of each fish prey type (species a (species fishprey type proportion ofeach stomach contents and preyavailability Providence onNew reefs, firstcalculated we the To identify the lionfishprey drivers of selec observations Indirect all levels ofall categorical variables. correctedZ tests Wald toobtain pair-wise ofthe relative comparisons selection probabilities for This article is protected Allrights bycopyright. reserved. Accepted proportion ofones(Bolker because function and cloglog-link appr Laplace using the obtained were random effect and eachfixed for estimates Parameter 1cm). nearest (TL the to lionfish ofthe body length fo the pointcountonwhichitwasobserved, nested three specified family, we Article effe (fixed facultative) or obligate (either cleaner and a defended, chemically or physically wasnocturnal, species prey the whether column, of ratio asthe (quantified shape body specific consum (proportion prey selection (i.e.proportional consumed thatwere of eachtype combination) 1 cmbins] within 1moffocal quan wefirst on Eleutherareefs, To identify the lionfishprey drivers of selecti observations Direct et al. et 2009; Zuur 2009; tified the abundance ofpotentialpr tified theabundance random effects, so that each potential prey was nestedwithin was so thateachpotentialprey effects, random ed), we included individual prey size (mass ing), individualprey species- (mass size ed), weincluded the proportion of zeros in our dataset greatly exceeded the greatly exceeded inourdataset zeros proportionof the nd length class [in 1-cm bins] combination) 1-cmbins] class[in nd length tion from our indirect observations oflionfish ourindirectobservations tion from hunting lionfish, and then calculated theproportion calculated hunting lionfish,andthen et al. et on from our direct observations oflionfishhunting ourdirectobservations on from body length to depth), position inthewater todepth),position length body cal lionfish that was being observed, and the andthe lionfishthatwasbeing observed, cal cts; Table 1). In Table1). additiontofishorderand cts; 2009). We conductedmu 2009).We response variable). As potentialpredictorsof response variable). oximation withabinomioximation

ey types (species and TL [in [in TL (speciesand ey types ltiple Bonferroni- ally distributed error ally distributederror method(Grueb using Average” the“Natural incorporated uncertainty model sample using AICc(for small sizes) model selection and al. ( i.e. those with average parameter estimates foreach trait prey variable fromitsweights inthe set oftop models dividing by and twosta themean) (subtracting centering by ourdata westandardized sizes effect relative compare In to order of models. betweencomplexity model and a model candidate (i.e., ofparameters) fitwithin the number set averaging, respectively and (Burnham Anderson support for predictions about the effect ofpr abouttheeffect support forpredictions (direct analyses For bothofour Model selectionandmu therelativeselecti of pair-wise comparisons itsabundance by and weighted Table1), effects; This article is protected Allrights bycopyright. reserved. predicte was proportional response) the where aGLMM weconstructed Next, reefs. estimat contents ofalllionfish,byits abundance dividing lionfishbyconsumed by theabunda

Accepted Article multipleBonferroni conducted 2009).Again,we Δ AICc < 4) in the MuMIn package (B 4)intheMuMIn< package AICc lti-model averaginglti-model and indirect observations), weev observations), and indirect d by its morphological and beha itsmorphological d by on probabilities for each categorical predictor. predictor. categorical for each on probabilities er et al. 2011; Burnham and Anderson 2002). 2002). 2011;Burnham andAnderson etal. er ey morphological andbe nce of each type, summed across thestomach summedacross eachtype, of nce ndard deviations(Gelma ndard e from our visual surveys of availabilityfrom ourvisualsurveys onthe of e proportion of each prey type consumed(i.e. prey type ofeach proportion in surveys of the study area (following Zuur (following surveys area ofthestudy in 2002). AICc values represent the trade-off thetrade-off represent 2002). AICc values -corrected post-hoc Wald Z teststoobtain post-hocWald -corrected arton 2013) for RStatisicalSoftware, arton 2013)

aluated relative the of strength vioural characteristics (fixed characteristics (fixed vioural havioural traitsand havioural n 2008). We calculated We n 2008). et et This article is protected Allrights bycopyright. reserved. Accepted Fig. 2; 1). (Table reefs on Eleuthera were types ofthese individualsofeach very few ph predation onnocturnal, of observed noinstances thantheirnon-cl topredation were lessvulnerable also foundsomeevidence benthos (Table2).We by lionfishthansc consumed more likely tobe (TableFig 2a 2;Fig. andb).Prey 1, livingephemeralfeeding solitarily orforming were In vulnerability line withourpredictions, measured topredation, as the of probability significconsumed, decreased being Articlecm TL; Appendix S1). fish prey availability all (i.e. individuals within1 m radius ofthe focal lionfishand less than 15 32speci Therewere fish. prey consumed asingle byconsumed lionfish asingle number ofprey ± Themaximum (4 ± TL SD). mean cm 1cm; 2-7 from prey ranged captured ± their while SD), S1).Lionfishspeciesfamilies andcm TL (Appendix 6 sizefrom 10-36 ranged (26±cm; mean 6 lionf 22hunting weobserved reefs On Eleuthera SELECTIONFROM DIRECTPREY OBSERVATIONS Results

antly with prey size and with increasingly deep body shapes body deep increasingly andwith preysize with antly during our observations was 4; twelvelionfish was during ourobservations hooling prey, as were prey living prey onorinthe as were prey,hooling observed withinthe observed vicinityof hunting lionfish ish capture a total of 32 prey fishes from12 a totalof32prey ish capture that fishes which exhibit cleaning behaviour thatfisheswhichexhibit es from 16 families recorded inpointcountsof from16familiesrecorded es eaning counterparts (Table 2; Figure 1). We 1).We 2;Figure (Table counterparts eaning ysically defended or pelagic prey however prey however orpelagic ysically defended

This article is protected Allrights bycopyright. reserved. pred behaviour found thatcleaning OnNew the Providence vulnerability reefs, again ofprey fishes tolionfishpredation We 1, Fig andd). 2; Fig. 2c (Table shapes body andincreasingly deep size withprey decreased S1). TL; Appendix all fishes<15cm visual surveys potentialprey (i.e., of r one prey fish.We stomachs containedonly a single lionfishstomachwas fishes observedin (4 ± 2-13cmTL from their fishprey ranged ± mean and (25 ± TL SD) –38cm 5cm; 10 from sizes in ranged examined Thelionfish S1). atotalof258prey specimens, wedocumented could be identified tospecies, either visually Provi collectedinNew In total637lionfishwere PREY SELECTION FROMINDIRECT OBSERVATIONS morphological and behavioural tra and behavioural morphological ofeach forms vulnerable exhibit that fishes combination, in considered are traits these surveys of ourenvironmental within prevalence no instance weobserved lionfish predation.Again lionfish (Table The physical presence 2;Fig. defense 1). of vulnerability didnot affect fish to withso their solitary counterparts, and shoaling schoolin Eleuthera reefs, As on (Table 2;Fig. 1). influenced alsostrongly nocturnal activity (Tab than non-cleaners lessfrequently relatively Accepted2). counterparts their ‘invulnerable’ Article of the same size and body shape on New Providence reefs (Fig. reefs Providence onNew body and shape samesize of the it are ~200 times more likely to be consumed by lionfishthan consumed ~200 timesmorelikely tobe it are icted consumption by lionfish, icted lionfish prey selection lionfish prey selection ecorded 103 species from 30 families duringfrom 30families the 103species ecorded or through DNA barcode analysis. Fromthese analysis. barcode orthrough DNA 2 cm; mean± numberofprey 2 cm; SD).Themaximum fishes from 33 species and 15 families (Appendix 15 families(Appendix and from33species fishes litary fishes most vulnerable to predation by topredation by litary fishesmost vulnerable le 2; Fig. Aggr le 1). New Providence reefs (Table 2; Fig. 1). When 2;Fig. When 1). (Table reefs Providence New 15; morethanhalf(57 15; dence and 108 contained whole fishprey that and 108contained dence g fishes were significantly less vulnerable than than vulnerable less significantly were g fishes s ofpredationonpelagi

on New Providence reefs reefs on NewProvidence with cleaners being selected selected being with cleaners egation behaviourand egation of 108) ofthelionfish of c prey, despitec their observations and This article is protected Allrights bycopyright. reserved. Acceptedsize and prey vigilance vulnerability on tolionfishpredation.feature that Another prey is work Further Lima& 1998). dependent (Bednekoff det and predator prey vigilance strategy hasb an anti-predator behaviour as th morevulnerable are whole, solitary species (Pitcher1993). strategy anti-predator effective the supporting reefs, vulnerability onEleuthera wefoundthat Countertoour prediction, of fishprey tolionfishpredation onNew Providence reefs, solitary and that behaviour increased predicted lionfishprey selec selected were never column >2mabovereefs to predationonEleuther benthos mostvulnerable column positionalso strongly influenced vulnerability topredation; prey occurring onand inthe Article thatdonotengage in and shallow-bodiedfishes cleaning determinants are behaviour vulnerability important of topredation, withsmall, solitary ofboth ouranalyses In particular, preyassemblages. within across species spatial scales, demonstrating that vulnerability by topredation can conferred traits be that recur different very two at collected data by confirmed were predators lionfish by selection affect that aboutthebehaviou ourpredictions The majority of Discussion ex situ stomach contents reveal that prey that reveal contents stomach tion in our large-scale analysis of New Providence reefs. Providencereefs. ofNew analysis tion inourlarge-scale ection strategy, which arethough ection strategy, een shown to depend on a combination school size, combination schoolsize, todependona een shown by lionfish inourstudy. also lionfish by an schooling species, the success of schooling ofschooling thesuccess an schooling species, While ourobservationsindicatethat,onthe While hypothesis thatschooling be can behaviour hypothesis parasite removal consum removal parasite schooling behaviourdecr schooling a reefs, and pelagic prey occupying the water water the occupying prey and pelagic reefs, a ral and morphological feat ral andmorphological needed to tease apart the effects ofschool theeffects apart neededtotease

size, body shape, aggregation and size bodysize, shape, t to be species- andcontext- t tobespecies- ed selectively. Water Water selectively. ed eased the vulnerability the eased in situ ures of reef fishes reef ures of behavioural This article is protected Allrights bycopyright. reserved. al. ofhe could impairtheprocess S1). Thus,high(Figure of 2C;Appendix rate 4-7) range: depth ratio (body shape a body are of grow vul throughherbivorous fishesclasses size vulnerability tolionfishpredationwith functional correlates role Forexample, ofprey. lionfishmay by re selective predation have time, inrelation tolionfish predationneeded are pressure, Moreover, totest thisprediction. persistence. Time-seriesdata documenting the re time, withpotentia substantial declinesover (Green can(e.g. reach Figure 2).Aslionfish that densities allow the rapid ofprey depletion showthat We mortality lionfishpredation from willbe greatest prey possessing a for andmorphological specific suiteofbehavioural lionfish’slength. only 15%ofthe relatively large- against simply ineffective be Alterna toacapture. lionfish immediately prior lionfish. During ourdirectobservati of many orbarbs)may spines (e.g. physical defenses prey use which in manner the that is explanation Onepossible defense. chemical or physical unexpectedly little influence had onvulnerability lionfishpredationa presence was to of the identity, usefulinsight ourfindings mayprovide fish communities elsewhere inthe data region; invaded onfishassemblages frombefore and

Acceptedanalyses Because 2006),inthelong our term. Article et al. et 2012), prey types selectivel thatare 2012),prey rbivory, which suppresses algal growth on coral reefs (Mumby (Mumby reefs growth oncoral suppressesalgal rbivory, which percussions oninvaded percussions lly implicationsforlocalpopulation serious be ineffectiveagainst the hunting of strategy bodied lionfish predators; prey were on average average on were lionfishpredators;prey bodied ons, prey appeared largely unaware of stalking ofstalking unaware largely ons, prey appeared lionfish-induced mortality on juveniles stages mortalitylionfish-induced on tively, the defenses of small-bodied prey may of small-bodied tively, thedefenses characteristics oninvaded characteristics nerable to lionfish predation, and many species andmany tolionfishpredation, nerable yconsumed may postmorerapidand focus on prey characteristics andnotspecies characteristics focus onprey lative change in biomass of prey types over over inbiomassofprey types lative change that is vulnerable to predation by lionfish that isvulnerabletopredation into the potential effects oflionfishpredationon the potentialeffects into

marine food websif marine food Atlantic coral reefs Atlantic coralreefs et et This article is protected Allrights bycopyright. reserved. information and onprey availability consumption be obtained, either through species share at least 1trait that ispredicted toaffect vulnerability topredation, and 3) a trait-basedapproach that: are 1)the predator's apply to requirements systems. these The only each of in structure, community for consequences predation willaid inforecastingchanges to predator-preyrelationships, ultimately and gene ofidentifying approach al. 2008).Using our [ predators ofnative diet the alter where they also inva are species Lowprey trophiclevel items. into new expanding rapidly predators thatare North (Herborg systems freshwater American Western 2013) a Atlantic (USGS of spread et al.2009),recent ( prawn Asian ( python the Burmese at occuring thatare exchange ofspecies consequence In particular, areamajor invasions exchange. biological species novelpredator Predicting of thestrength ecological patterns and processesaffected willbe by the human-mediated ongoing of process species) targeted forlionfishculling. could be (i.e. of vulnerableindividuals locations example, action. For early intheregion invasion ofa may establish be used spatial to priorities formanagement AcceptedLymantria dispar Article ] by] [ native Python molurusbivittatus Python an increasing rate (Mooney and (Mooney rate an increasing small-bodied, shallow-bodied, benthi small-bodied, shallow-bodied, where native fish communities ar communities where nativefish nd progression of snakehead fishes (Channidae) throughout throughout fishes (Channidae) snakehead of nd progression Coccyzus erythrpthalmus erythrpthalmus Coccyzus (e.g. the consumptionofinvasivegypsy (e.g. areas, where they encounter and consumenovel and encounter wherethey areas, et al et ) northward from the Florida Everglades (Rodda ) northward theFlorida from diet consistent diet ofmultiple species, 2)prey -prey is essential for understanding how forunderstanding -prey isessential Penaeus monodon Penaeus ral prey traits that ral prey traits . 2007) are just a few examples ofnon-native examples few . 2007)arejusta ding non-native ranges at an increasing rate, anincreasing at ding ranges non-native

Cleland 2001). of Cleland 2001). The expansion e composed ofahighproportion e composed and ) into the Gulf of Mexico and GulfofMexico ) intothe c or demersial, and nocturnal andnocturnal c ordemersial, confer vulnerabilityconfer to C. americanus in situ ] (Barber et (Barber ] observation is ofpredation direct observation we canbe selection diet that suggests analyses compositi wedid)orscat (as gutexamining content similartoour broad-scale gathered viamethods and species predatory formany challenging traits as important ofdiet drivers selecti analyses andbroad-scale observations ofpredation at simultaneously selection diet examine ecological communitiesHartvig (e.g., prey confer and behaviour This article is protected Allrights bycopyright. reserved. Accepted view indicates thatatrait-based study isthegeneral firstidentify predict ofprey traits that vulnerability topredation, and Article Blanchard identity 2005, (Jennings species largely ignoring size, based onbody view 1980, PolisandStrong 1996),andasize-based constructed by quantifying the interaction streng intotwoprevailing paradigms: characterized thinkingMcGill etal.2006).Current aboutpr Finally, study thatgenerating a functional shows our traits approach has for value ofprey selecti abouttheeffects abundance. environmental prey dietin broad-scale pairing of predationorby selection, also has significant influence onspecies interactions within of predator-prey interactions, interactions, ofpredator-prey notlogistically feasible. et al. et on. However, directly observing prey capture capture prey observing directly on. However, two spatial scales.Importantly,two spatial our both on on the structure of ecological communities(i.e. ecological on thestructureof 2011). It firstto 2011). the isalso,toourknowledge, information ondietcomposition iscommonlyinformation formation fromguts withsurveysformation ortissues of a species-based view, in which food webs are foodwebsare inwhich view, a species-based edator-prey interactions can be broadly be broadly can interactions edator-prey ll estimated from these broad-scale analyses when when analyses broad-scale these from estimated ll analyses, with 'snapshots' of diet obtained by ofdietobtainedby 'snapshots' with analyses, th between pairs of predators and prey (Paine andprey (Paine ofpredators between pairs th ofstomach contents the identified same prey on. The similarity identified traits of by both , which classifies predator-prey interactions interactions predator-prey , whichclassifies

in which variation inmorphologyin whichvariation et al. et in situ 2011). Our 2011).Our in situ is This article is protected Allrights bycopyright. reserved. 2004.Oddspec and M.S.Webster. Almany, G.R. Syms,C. M.McCormick, L. Peacock, Almany, G., References Council (NSERC)Cana by Funding wasprovided and DavidFerguson. donated generously Providence was ofstatisticalanal discussion Group for Research on themanuscript,an Caley comments Julian for NicholasDulvy, TedGrosholz, to MarkHixon, Foundation Environmental Education Lad SusanReigner,grateful to Akins, are We Acknowledgments available through Fig Sharepending acceptance of thestudyfor publication. D Data accessibility

Acceptedata used in analyses: speciesdata uploaded Article reef fishes. Ecology 85:2933-2937. 85:2933-2937. fishes. Ecology reef 152:751-761. Oecologia fishassemblages. reef prey incoral

da Graduate Fellowship to SJG and FellowshiptoSJG da Graduate by Stuart Cove’s Dive Bahamas and on Eleuthera by Trisha Trisha Eleutheraby andon Bahamas Dive StuartCove’s by volunteers for assistance in the fi for assistanceinthe volunteers as online supporting information (Appendix S1) and Annabelle Brooks, Leah Neal, and many Reef andmany Leah Neal, Brooks, Annabelle Jonathan Moore, John Bruno, Julia Baum and Julia Bruno, Moore,John Jonathan a Natural Science and Engineering Research Research andEngineering Science Natural a yses. Logisticalyses. thisstudy onNew supportfor d to Andrew Cooper and the Earth to Ocean EarthtoOcean and the toAndrewCooper d ies out as predators reduce diversity ofcoral- diversity reduce ies outaspredators and G. Jones. 2007. Predators target rare target rare 2007.Predators andG.Jones.

an NSERCDiscoveryIMC. to Grant eld. We are also grateful arealso We eld. This article is protected Allrights bycopyright. reserved. AcceptedLaw, andthe R. Castle, andS.Jennings. 2011.Coupledenergy pathways J., M. Blanchard, 2004. G.P.Jones. and B.Beukers-Stewart, D. P,G.Or A.Acero Betancur-R, R.,A.Hines, L.Lima. 1998.Re–exami Bednekoff, P.A.andS. 0.999999-0. version classes.Rpackage Article 2012.lm B.Bolker. and Bates, D.,M.Maechler project.org/package=lme4 http://CRAN.R- 1.9.13. R package version Multi-modelinference. Barton, K.2013.MuMIn: P.To Marquis,andW. A., R.J. N. Barber, I. N.Smith, andM. P.,J. 1996. Hatch. Arcese, risk dilutionandcollectivedetectionde 4611. of Proceedings demography. for passerine resilience of size-structured food resilience ofsize-structured Biology 299:155-184. and Ecology speciesofcoral oftwopiscivorous ecology 38:1281-1293. ofBiogeography Journal lionfishinvasion: insighReconstructing the 2026. London. of Society oftheRoyal Proceedings distribution ofnatievpred ators.. Ecology 89:2678-2683. ators.. Ecology webs. 4:289-300. Ecology 4:289-300. webs. Theoretical ri. 2008. Invasive prey impacts the abundance and Invasiveri. 2008. theabundance prey impacts tí, A. E. Wilbur, and D. W. Freshwater. 2011. andD.W. E. Wilbur, tí, A. The influence of prey abundance on the feeding feeding onthe abundance prey The influenceof pend upon predator targeting behaviour. behaviour. predatortargeting upon pend Nest predation by cowbir predationby Nest e4: Linear mixed-effects modelsusingLinear S4 mixed-effects e4: http://CRAN.R-project.org/package=lme4 http://CRAN.R-project.org/package=lme4 ning safety in numbers: interactions between ning interactionsbetween safety innumbers: the National Academy of Sciences 93:4608- ofSciences Academy National the ts into Greater Caribbean . biogeography. Caribbean into Greater ts reef fish. Journal of fish. Journal reef Series B: Biological Sciences 265:2021- Biological Sciences B: Series

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and management of the and management on hard bottom reefs of on hardbottomreefsof Pterois volitans ) in This article is protected Allrights bycopyright. reserved. Linkage, Webs: Paine, R.T.1980.Food interact andjuvenile oflarval ecology I.G. 2012.Theforaging and Cowx. L.H. Tweson Nunn, A.D., 2000.PreyNilsson, P.A.andC.Brönmark. 2003.Rapidworldw B.Myers, Worm. R.A.and Pitcher, T.1993.Fish schooling behaviour. Phillips, R. McGill,andD. R., S.Bearhop, Pyke, G. H., H. R. Pulliam, and E. L.H. R.Pulliam,andE. Pyke, G.H., Charnov. G. Chouinard Lucas,Coderre and Provost, C.,E. D. FoodPolis, G.A.andD.R.Strong. web 1996. Accepted andR.Reed. 2009.What Rodda, G.,C.Jarnevich, Article fishes. Reviews in Fish Biology and Fisheries. 22: 377-408. Fisheries.Biology 22: fishes. ReviewsinFish and impactson behavioural andmorphological Nature 423:280-283. nonbreeding period. Oecologia 160:795-806. 160:795-806. Oecologia period. nonbreeding variation inmigration strategies and preferences ina during suite ofseabirds the ofAnimalEcology 49:667-685. Journal Fishes. Croom Helm Ltd. Ltd. Fishes. CroomHelm 265-277. axyridis: The influence preyInsect mobility of species. and of prey Journal Behavior. 19: Naturalist 147:813-846. theory and tests. The Quarterly Review of Biology 52:137. Biology52:137. Reviewof theoryand tests.TheQuarterly Invasions 11:241-252. pythonssuitable forinvasivealien spreadin in vulnerability to a gape-size limitedgape-size predator: vulnerability toa Dawson. 2009. Stable isotopes reveal individual 2009. Stableisotopesreveal Dawson. T. Pitcher, editor. The Behaviour ofTeleost Behaviour Pitcher, editor.The T. complexity and community dynamics. American American dynamics. andcommunity complexity 1977. Optimal foraging: A selective review of selective review A Optimalforaging: 1977. ion strength and community infrastructure. and community strength infrastructure. ion ide depletion of predatory fishcommunities. ide depletionofpredatory partsoftheUSmain northern pikepiscivory.Oikos 88:539-546. northern g from Everglades National Park? Biological Park? National Everglades g from . Prey selection by the lady beetle Harmonia Harmonia . Prey beetle thelady by selection

land are climatically areclimatically land This article is protected Allrights bycopyright. reserved. and environm R CoreTeam.2012.R:Alanguage 67:359-367. Biology Fishes. Environmental of predatordeterrent. ‘lands 1989.Theocean Steele, J. toxicity fromcoral-dwelling ofskinsecretions gobies and their potential as a role L.,Llewellyn, Munday,L. E.2003.The P. Jones, G.P.and Schubert, M.J., Caley, M.J., W. Murdoch. 1983.Selec Scott, M.A.andW. forage 2003.Vulnerabilityof marine P.A.McGinn,andF. A.Buckel, Juanes. F. S.,J. Scharf, F. andR.A.Stein.Savino, J. 1982.Predato Accepted M.S. C.K., G.R.Almany, D.H.Williamson, Wen, tolargemouth Esocids ofthree D.H.andR.A.Stein.1989.Comparativevulnerability Wahl, ofhabitatselection D.H.1995. Effect Wahl, Article and Oceanography 28:352-366. 28:352-366. Limnologyand Oceanography Marine ofExperimental Journal fishes topiscivory: ofprey onsusceptibility behavior effects toattackcapture. and Fisheries 111:255-266. Society as influenced bysimulated, submersed vegetation. the Transactions American of Sciences 46:2095-2103. Sciences 46:2095-2103. bass ( ofFi Journal introduced fish.Canadian Statistical Computing, Austria. Vienna, predatory fishes. Marine Ecol Marine predatory fishes. effectsreserves ofmarine ondiet, prey availabilityand selection prey by juvenile Micropterus salmoides cape’. 3:185-192. EcologyLandscape 3:185-192. cape’. ) predation. Canadian Journa Canadian ) predation. ogy Progress Series. 469: 133-144. Series. ogy Progress Biology and Ecology 294:41-59. 294:41-59. Ecology Biology and r-prey interaction betwee r-prey and behavior onvulnerab and tive Predation by the Backswimmer, Notonecta. Backswimmer, the Predationby tive sheries and Aquatic sheries and ent forstatisticalcomputing. RFoundation for Pratchett and G.P. Jones. 2012.Evaluating the Pratchett andG.P.Jones.

l of Fisheries and Aquatic l ofFisheries and Sciences 52:2312-2319. Sciences n largemouth bassand n largemouth ility topredationof ility

Morpholo Behaviou

This article is protected Allrights bycopyright. reserved. 1. Table A. Walker, Ieno,N. J. N. Zuur,A. F., E.

Acceptedgy Article r

predation by invasive lionfish ( invasive lionfish predation by Models and Extensions in Ecolog Models andExtensions Morphological and behavioural tr and behavioural Morphological attribute attribute Prey Water Water Nocturnall Aggregatio Cleaning or Physical size Body shape Body column column y active n size behaviour defense chemical position

al al Categoric Binary al Categoric Binary Binary us Continuo us Continuo

Data type type Units/Levels Benthic, demersal, Benthic, demersal, Yes/no =0/1 Solitary, shoaling, Yes/no =0/1 yes/no = 0/1 Mass (g) Body length todepth Pterois volitans/miles indicate increasingly increasingly indicate ratio: Higher values pelagic pelagic schooling shapes linear y with R. Springere, New York. York. New y Springere, with R. aits of prey fish predicted prey fishpredicted aits of Saveliev, and G. M. Smith. 2009. Mixed Effects Saveliev,andG.M.Smith.2009.Mixed

) vulnerability ↑ predation Predictions vulnerability of to Pelagic < benthic < demersal + nocturnality = Solitary < shoaling < schooling = behaviour + cleaning = + defense ↑ ratio (i.e. ratio mass= to influencevulnerability to ↓ vulnerability vulnerability ↑ ↓ shallow bodied) =shallow bodied) vulnerability ↑ vulnerability ↓ vulnerability ↑

Response observati variable feeding In situ ons variable. variable. Relative variable importance isthe sumofthe weights ofall models that that contain particular models considered with withthose model of those considered, and model ofthoseconsidered, This article is protected Allrights bycopyright. reserved. Providence, Bahamas. from lionfishstomachcontentsandvisual inferred A. Table 2.

Accepted Article in situ Ra nk 11 10 8 7 6 5 4 9 3 2 1 Top generalised linear mixed-effects models linearmixed-effects Top generalised observations oflionfishpredationoncora 9 9 9 9 9 9 shape Body

Morphology Behaviour 9 9 9 9 9 9 9 9 9 9 9 Bo siz dy e

Δ AICc istheAICc difference in 9

Clea ner

w i

9 9 9 9 is the probabilitythat model isthe the best set. of The top Defe Δ nse

AICc < 4. 9 9 9 9 9 9 9 9 9 Aggreg ation

size surveys of reef fish on coral reefs off New off reefs fish oncoral surveys ofreef l reefs off Eleuthera, Bahamas, and Bahamas, offEleuthera, reefs l 9 9

Nocturn (GLMMs) of lionfish dietselectionfrom (GLMMs) oflionfish

ality

values between model model values between

9 9 posit Wat colu mn ion er

logL 71.5 72.6 69.2 68.9 68.8 69.2 69.1 74.0 70.7 69.0 69.4 ik

8 7 9 0 0 4 4 4 3 9 1 ------160. 160. 159. 161. 161. 159. 159. 160. 160. 159. 157. Cc Cc AI 00 00 90 50 30 90 60 60 60 50 90 i and thebest Cc Cc AI 2.1 2.0 2.0 3.5 3.3 1.9 1.7 2.6 2.6 1.6 0.0 Δ 2 0.07 8 8 0.07 7 2 0.07 2 0.03 2 0.04 2 0.07 2 0.08 6 0.05 6 4 0.05 9 0 0.09 0 0.19 9

B. Wei

ght d 0 1 1 1 1 1 0 1 0 1 0 1 f importanc (direct/in Stomach Stomach contents analysis variable Relative direct) e This article is protected Allrights bycopyright. reserved. Accepted Article 14 13 12 7 6 5 4 1 3 8 2

9 9 9 9 9 9 9 0.63/ 0.87

9 9 9 9 9 9 9 9 9 9 9 1/1

9 9 9 9 9 9 9 9 0.16/ 0.89

9 9

9 9 9 0.23/

0.5

9 9 9 9 9 9 9 9 9 9 0.85/1 0.18/1

9 9 9 9 9 9 9 9 9

9 9 9 9 9

0.2/0 .65

372. 373. 375. 370. 374. 69.0 68.9 369. 72.3 374. 374. 45 25 27 78 08 91 82 74 0 2 3 ------765. 764. 764. 762. 764. 161. 161. 762. 161. 765. 765. 30 90 80 00 40 70 50 30 50 90 80 3.3 2.8 2.7 0.0 2.4 3.7 3.5 0.3 3.6 3.9 3.7 3 0.06 6 0.08 9 6 0.08 7 0 0.32 4 0.10 8 2 0.03 6 0.03 3 0.27 1 0.03 8 2 0.05 8 6 0.05 8 0 1 0 1 1 1 1 1 1 1 consumed by lionfishinthedataset. consumed by noindi >100) because [SE] error (i.e., standard large with uncertainty estimated values, parameter *areaverage with denoted Numbers set. data g)as per Table tra indicates shading 1.Grey measuredasbody the ratio length of todept (cm) 'Benthic','Schooling' and and 'Solitary' arecompared against'Shoaling'. 'Body shape' is level baseline the against compared are 'Pelagic' and 'Demersal' Levels avoidance. indicate line) indicate selective onprey displaying predation trait, while negative the (tothe values left ofthe intervals. 95% confidence mean valuesboundedby (S coral reefs Providence andonNew observations') This article is protected Allrights bycopyright. reserved. Accepteddenote theof prey combinations fishsizeyielding and bodydepth ratio vulnerability low to Elacatinus genie. C) solitary such as fishes nocturnal deeper-bodied morphological and behaviour offon coralreefs Ne fishes bodied shapes)for indicat body values depth;lower g)combinations ofprey (massin size sh andbody Fig. 2 Article(GLMMs; Fig. 1 Figure legends Model-averaged coefficients from the top generalised linear mixed-effects models models mixed-effects linear generalised top the from coefficients Model-averaged Example contour plots ofpredicted contourExample mean vulnerability topredation by lionfish forvarious Δ AICc <4; Table 2)oflionfishprey Acanthurus bahianus Acanthurus Blackbands denotechange a probability Cooler in ofpredation. colours al traits: as demersal fishessuch A) shoaling e deeper-bodied shapes whereas shapeswhereas e deeper-bodied

, B) schooling pelagic fishes such as such fishes pelagic , B)schooling Apogon binotatius, Apogon binotatius, it forms that were not present in the corresponding in thecorresponding thatwerenotpresent it forms w Providence, Bahamas, displaying varying w Providence, viduals with that specific trait formwere withthatspecifictrait viduals selection on Eleuthera coral reefs (' reefs coral onEleuthera selection h (cm), and 'Body size' is measured as mass(in 'Body measured size'h (cm),and is ape as (quantified the offishlength ratio to Positivevalues(totheright oftheline) tomach contents analysis). Points represent represent Points analysis). contents tomach and D) benthic cleaning fishes suchas and D)benthiccleaning fishes

higher values denoteshallow- values higher Decapterus macarellus,Decapterus Scarus isertiScarus in situ and the This article is protected Allrights bycopyright. reserved. Accepted Article 1 Fig. Species illustrationsmodifiedwith predation, whilecolours warmer denote scenarios permission from ReefNet (2007). ReefNet(2007). permissionfrom in which vulnerabilitypredation ishigh. to

This article is protected Allrights bycopyright. reserved. Accepted Article 2 Fig.