View metadata,citationandsimilarpapersatcore.ac.uk 8 7 Germany 6 Tucumán, de Nacional Universidad 5 the Netherlands Amsterdam, GE 4 3 Germany Halle, 2 1 Saavedra IngolfKühn M.A.Bender Irene matchingshapesplant trait Morphological ifrne bten hs eso ad h Vrin f eod Pes c Please Record. of Version [10.1111/ the and to version lead this may between which process, differences proofreading and pagination been typesetting, not copyediting, has but the review through peer full undergone and publication for accepted been has article This date: Decision Jena author Corresponding Z New 8140, Christchurch 11 10 9 c 27, c. Cota, Cota Universitario

AcceptedAlexan ResearchMuseum Zoological Campus Andrés, San de Mayor Universidad InstitutoEcología, de (LPB), Bolivia de Nacional Herbario Research Environmental for Centre Helmholtz Fra 60439 Biologicum, & Evolution Diversity, Ecology, for Institute Frankfurt, University Goethe ArticleyTécni Científicas Investigaciones de Nacional Consejo Regional, Ecología de Instituto Amsterdam, of University (IBED), Dynamics Ecosystem and Biodiversity for Institute Centre Research Climate and Biodiversity Senckenberg Halle Luther University Martin Garden, Botanical and Geobotany InstituteofBiology, for Centre German Centre for Integrative Ecology, School of Biological Sciences, University of Canterbury, Private Bag 4800, Bag Private Canterbury, of University Sciences, Biological of School Ecology, Integrative for Centre Research Environmental for Centre Helmholtz - Leipzig, 04103 Leipzig, Germany. Email: Leipzig,Email: Germany. Leipzig,04103 8

, Vinicio Santillán Vinicio , ecog 7,2,1 , Marcia C.Muñoz Marcia , 19

.0 - 3396 1,2,3 Feb Integrative Biodiversity Research (iDiv) Halle (iDiv) Research Biodiversity Integrative , W. Daniel W. Kissling , - 2018 ]. : ‘This article is protected by copyright. All rights reserved Irene M.A. Bender Irene

ealand 3,6

, Till Töpfer Till , asilla 10077, LaBolivia Paz, asilla10077,

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CC 34, 4107 Argentina Tucumán, 4107 CC34, , Eike Lena Neuschulz Lena Eike , der Koenig, 5 Koenig, der 9 4 , Thorsten Wiegand , Thorsten , Pedro G. G. Blendinger Pedro ,

,

- German Centre for Integrative Biodiversity Research (iDiv) Halle Research(iDiv) Biodiversity Integrative for GermanCentre frugivore networks across the Andes the across networks frugivore [email protected] - - UFZ, UFZ,

UFZ, Dept. Ecological Mo Ecological Dept. UFZ, 3113 Germany Bonn, 3113 Dept. Communit Dept. ,

60325 FrankfurtGermany (Main), 60325 3 , Larissa , Nowak

1,10 5 , D. Matthias D. Dehling , , Katrin Böhning Katrin , - Jena

- y Ecology, 06120 Halle, Germany Halle, y06120 Ecology, Leipzig, 04103 Leipzig, Germany Leipzig,04103

3,6 delling, 04318 Leipzig, Germany delling,04318 , Marta Quitián Marta , - Gaese

t ti atce s doi: as article this ite - 3,11 ’ POSTBUS 94248, 109 94248, POSTBUS W . cas (CONICET) (CONICET) cas

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Matthias Schleuning Matthias

, Isabell , 3,6 , Francisco Francisco , nkfurt (Main),

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CORE into the generality of interaction rules are the base for predictions of species interactions in ecological networks, networks, ecological in interactions species predictionsoffor thebase rules are interaction generalityof intothe The specialization. of functional degree the approximate usedto cantraitsbe bird and specificplant in generalphenomenon a is birds frugivorous plantsand birds. wingin ofthe pointedness mass in plants crop to related positively was partners interaction specific on specialization functional h (plant that found plant bird and ofplant specialization traits morphological whichwe ofthe test Inaddition, networks. inthese species matchingbetween trait determine traits animal and plant morphological of combinations the tropica across sampled birds frugivorous plantsand fleshy between networks eight interaction on we data Here yetcombine not understood. are partners the and interactionrules trait generalityof the but partners, interacting oftraitsthe functional by the influenced are networks these in interactions Species networks. in ecological organized are species consumer and resource Interactionsbetween Abstract Accepted speciali fulfill that species animalidentify to plant and applied canbe future, and in the communities novel in instance for Article zed functional roles in ecological communities. ecological in roles functional zed e ight

- wingshape - animal trait pairs related to size matching size to related pairs trait animal

‘This article is protected by copyright. All rights reserved relationship between functional traits and a species' specialization on specific interaction specificoninteraction specialization species' a and traits functional between relationship

and

crop mass crop

species

Our findings show that morph showthat Ourfindings

. In a metaIna . - body mass) mass) body - analysis across network across analysis were positively related in these networks. The degree of degree The networks. in these related werepositively

l and subtropical Andean range. We test whichtest We Andeanrange. subtropical l and plant

(fruit size(fruit - frugivore networks frugivore ological trait matching between fleshy matchingbetween trait ological - beak size) beak - specific fourth specific

and avian foraging behavior behavior foraging avian and influence influence

across the Andes and that Andes and the across ’

. resource and consumer consumer and resource -

corner analyses, we analyses, corner functiona l se insightsse -

fruited fruited and to the to and - fruited fruited - based based species that interact with different sets of interaction partners fulfill different functional roles in ecological inecological roles functional fulfill different partners interaction setsof withdifferent thatinteract species nic Eltonian the of concept from the follows different localities at sampled networks interaction across matchi on based rules interaction general identify is to suchprojections firsttowards step The 2016). (Morales communities potential future or incurrent likely interact are to makepr usedto canbe species consumer and resource between interactions govern the that matchingrules trait about Knowledge 1987). (Levey systems manyinteraction in matching (Gravel partners potential al. (Cohen small to constrained are usually predators whereassmall species, with prey large interact Maglianesi Stang 2007, Warren Woodwardand 1985, (Wheelwright partners potential between matching oftrait degree the by often influenced are communities ecological in interactions Species Introduction Accepted localities. from different networks across specialization functional of high a degree with associated are that identifytraits matching to is therefore networks ecological to roles functional specialized the of understanding general more get a to Onepossibility traits. morphological own fromtheir directly species animal and ofplant specialization functional of degree approximatethe to no are interactions species on Since data 2017). (Mello ecosystem the entire networkand thefor interaction especiallyimportant c specialists the co to in respect specialized functionally interact are Olesen e.g. roles, functional intheir (Dehling partners theirinteraction of traits functional istherefore ofspecies role functional the networks, Ininteraction communities. Article

2016) Matching traits also define a species' functional role in interaction networks (Schleuning (Schleuning networks in interaction role functional species' a define also Matchingtraits et al. et et al. et .

It should therefore be possible to infer interaction rules from the traits influencing the matching of of matching the influencing traits from the rules infer to interaction possible be shouldtherefore It et al. et an provide ecological functions that no or few other species can provide and may be considered considered be may and canprovide species few other or no that functions ecological anprovide

2011a, Bender Bender 2011a, 1993, 1993,

2015). For instance, size matching describes the instance, describes matching For size 2015). Woodward and Warren 2007, notable exceptions include large marine mammals, Tucker mammals,Tucker marine large exceptions notable include 2007, Warren and Woodward ‘This article is protected by copyright. All rights reserved et al. et , et al.

in their degree of specialization on specific partners (Waser and Olle and (Waser partners specific on specialization of theirdegree in

2013) since the likelihood of interactions increases with the degree of trait oftrait degree withthe increases ofinteractions since the likelihood 2013)

2017 ), species that interact with a set of species with which few other species species fewwhich other with species withsetof a thatinteract species ), he (Elton 1927, Chase and Leibold 2003), which that assumes 2003), Leibold and Chase 1927, he (Elton toriously difficult to record (Jordano 2016), it would be desirable woulddesirable it 2016), be (Jordano record to difficult toriously et al. et - occurring species (Dehling species occurring

(Kissling and Schleuning 2015) Schleuning and (Kissling

2016 , Bender , Bender phenomenon - Castilla Castilla et al. et

et al. et ojections about which pairs of species whichofspecies pairs about ojections

2017 that large predators tend to to tend predators large that

most accurately described by the mostdescribed accurately types of species that contribute thatcontribute species of types

2009, De 2009, et al. et et al. et

et al. et ). Since species differ widely differ Since species ).

2015, Sebastián 2015, 2015, Bartomeus 2015, ’

. - 2016). These functional functional These 2016).

sized prey species sized . hling

et al. et et al. et

2015). This 2015). rton 2006 rton ng traits

2014, 2014, - González González

et al. et , et

Coux speciali (Sebastián networks in roles functional (Dehling withinnetworks specialization mayalso sametraits the networks, interaction in species ofmatching the influence traits, ofthese the combination and Since different traits, 1993). Davies and (Krebs resources Petchey (Belovsky1997, energy require species large foraging, optimal theoryof with the consistent Additionally, ofpla the location and shape, wing avian mobility,suchas with animal associated traits to related instance, for are, consumersand Sebastián 2015, Stang 1985, (Wheelwright size or bill as proboscis such organs, animal respective the and sizefruit or flower matching between a to instance, for correspond, and matching) (size sizeanimal oftr consists group first The groups. intotwo categorized Stang 1987, Jordano 1985, (Wheelwright Accepted 1985, (Wheelwright ofinteractions likelihood the on constraints direct matchingimposes size because pairs, si to related pairs trait that(i) expect We networks. across animals and ofplants specialization functional the influence traits in plant specialization functional determine that traits functional identify Andes ofthe range geographical wide frugivorous and birds Davies 2008) and (Jansson fleshy both for hotspot diversity global a is mountain range Andean networks.The frugivore countries from fivedifferent networks eight interaction wethis end, combined To networks. these in species animal and ofplant specialization functional the matching and oftrait the degree both for traits matching of ofdifferent pairs Article Matching traits hav Matchingtraits Specifically, we ask (i) which trait pairs influence the matching between plants and animals, and (ii) which (ii) and animals, and plants matchingbetween the influence pairs trait which we(i) ask Specifically, we Here, et al. et zation in mutualistic networks has focused on specific localities and on the consumer perspective (e.g. perspective consumer the on and localities specific on focused has networks mutualistic in zation

stretching the tropical and subtropical Andean range to identify general interaction rules in plant rules in interaction general identify to Andeanrange subtropical and tropical the stretching

2016, 2016, ze matching will be more important in shaping plant moreshaping important in will be matching ze conduct a large a conduct - González González Dehling nt resources, such as forest stratum (Moermond and Denslow 1985, Schleuning 1985, Denslow and stratum(Moermond forest such as nt resources, e been previously identified in studies of mutualisticplant of studies in identified previously been e ‘This article is protected by copyright. All rights reserved et al. et et al. et et al. et - scale analysis across mutualistic seed mutualistic analysisacross scale

2008). Therefore, animals with a large body size tend to prefer energy prefer to tend size bodylarge a animals with Therefore, 2008).

2016, Bender Bender 2016, 2017). Second, matching traits can be related to the foraging behavior of the foragingto behavior related canmatchingbe Second, traits 2017). ,

- makes it an ideal region to test the generality of trait matching and to to matchingand trait generalityof testthe to region makesideal itan González 2017). Most previous work on trait matching and functional functional and matching work trait onMost previous 2017). González et al. et

et al. et

etal. 2016) and may be used to identify species with important withimportant identifyspecies usedto be may and 2016)

2009, Dehling 2009, (Kissling (Kissling

2017)

. aits which relate to the matching between plant and plant and matchingbetween the whichto relate aits

et al. et et al. et -

animal interactions than foraging than interactions animal 2009). This diversity, in combination withthe combination diversity, in This 2009). -

dispersal networks to compa to networks dispersal 2014, Maglianesi 2014,

influence the degree of functional degree the influence - frug ivore networks. networks. ivore - animal interaction networks networks interaction animal et al. et

et al. et

’ 2009, 2009, . -

rich resources resources rich

2015) and can be canbe and 2015) - fruited plants fruited

Garibaldi Garibaldi re the importance re - et al. et related trait trait related - rich

2011). 2011). et al. et -

,

communities (Saavedra (Saavedra communities impoverished inweakened disturbed, may be traits species of importance the because in the analyses considered in near collected 4.7 networks cover The (Fig.1). Argentina from network one and from Bolivia network one Peru, twofrom networks from Ecuador, networks two fromColombia, two networks included eight networks The Andes. the across locations eight different fleshy between networks interaction fromcombined data We networks Interaction Methods 1988). (Wheelwright communities respective are and Wheelwright 1985) 1985, Denslow and (Moermond other eachon specialized reciprocally are species these because networks across specialization functional of level highest showthe Sebastián Accepted (Oksanen estimator Chaorichness theon based link and richness bird asymptotic curve Appendix1. in is given network each of ofthe sampling description (genus were thetanagers oflargestbirds group The was genus plant most rich species The interaction events. 11,578 inspecies bird 180 with interacting plant species we227 Overall, analysed (Fig.1). species bird 61 plant and networkcompr smallest thewas events: 241 interaction (rangeofobserved networks among varied interactionevents ofobserved but thenumber 224), = deviation standard mean606, = carrying or swallowing fruit trueseed only included We networks). (weighted plant species Article °N to 26.5°S, covering a large portion of the tropical and subtropical Andean range. All networks were Allnetworks Andeanrange. subtropical and ofthe tropical portion coveringlarge a 26.5°S, °N to To assess the sampling completeness completeness assesssampling the To offr visits frequencyof theon information comprise networks All s

of the observed bird richness and the number of interaction pairs (link richness) and calculated the calculated and richness) (link pairs ofinteraction number the and richness bird ofthe observed

- González González - natural forest, forest, natural et al. et

et al. et ‘This article is protected by copyright. All rights reserved

2017). In addition, w Inaddition, 2017). - away. away.

an elevation range from 1000 to 3000 m a.s.l. (hereafter “m”) and extend from extend and “m”) m to (hereafter 3000 a.s.l. 1000 from range anelevation 2014). 2014). i.e ising 19 plant and 22 bird species. The largest network from Peru comprised 52 52 comprised Peru from largest network The species. bird 22 plant and ising19 ., networks from highly fragmented fragmented highlynetworks from., Sampling Sampling

of each network (Chacoff network(Chacoff ofeach -

4988, mean = 1447, SD = 1539). SD = mean 1447, = 4988,

effort was effort e expect that (ii) large (ii) that expect e Tangara high in high all

, 19 species, family Thraupidae). family species, Thraupidae). 19 , - fruited plants and frugivorous and fruitedplants - dispersal events in the analyses that correspond to to that correspond analyses the in events dispersal Miconia

networks (range of sampling hours: sampling (rangeof networks et al. et

- fruited plants and large and fruitedplants ugivorous bird species on fleshy on species ugivorousbird or intensively disturbed forests werenot forests disturbed intensively or

2012), we generated accumulation accumulation we generated 2012),

(Melastomataceae) with species. 39 (Melastomataceae)

The network sampled in Bolivia in sampled Bolivia network The et al

. 2015). All interaction 2015). . relativelythe in rare ’ .

birds collected at collected birds - billed bird species species bird billed A more detailed moreAdetailed -

fruited 300 -

960, 960, maneuverability in dense vegetation (Rayner 1988). Bird traits were measuredon weretraits Bird 1988). (Rayner vegetation dense in maneuverability high for suited especiallywings are that rounded indicates index Kipp’s low a whereas distances, Winkler1991 wing folded measuredthe on wingtip, the and first secondary ofthe the tip between distance pointe ofthe measure as a index Kipp’s (i.e., shapewing (mm), length width(mm),bill bill included traits Bird availability th from to collected one species fruits per 20 least measuredat on were mass mass). fruit meanindividual by plant multiplied fruitsper of number (g, (Dehling in plant traits matching as key wereidentified that traits selected Morphologica traits Morphological A1). (Fig. networks all in richness the asymptotic richness approached Bird A2). A1, (Figs. curves accumulation in the trends saturating the by as indicated sampled, well weresimilarly networks Accepted matrix interaction thespecies Dehling (Legendre variables environmental and traits species between relationship fourth the version of plant shape combinations identifywhich to trait Inorder Trait matching werenot log index Kipp’s heightPlant and data. oftrait normaldistribution a approximate to the analysis prior mass) body lengthand methodologies. the using same werecollected networks specimens Article We log We dness of the wing) and body mass (g). Kipp’s index was calculated by dividing the Kipp’s distance (the distance Kipp’s the by dividing calculated was index massKipp’s (g). body and wing) ofthe dness et al. et et al. et , - . Plant height and number of fruits were measured on all individuals of a species present in a plot. inpresent a species ofa individuals all on measured were fruits of number and height Plant . transformed plant traits (fruit diameter, fruit length and crop mass) and bird traits (bill width, bill (bill traits mass) bird and crop and length fruit diameter, (fruit plant traits transformed wasmass body

) by wing length. A high Kipp’s in A Kipp’s wingbyhigh ) length. (2014) modified this approach to test the relationship between plant traits and and plant traits between testthe relationship to thismodified approach (2014) l traits were collected for all plant and bird species observed in the interaction networks. We We networks. interaction in the observed species bird plant and all for werecollected l traits

2014). Plant traits included fruit included traits Plant 2014). -

corner analysis. The fourth The analysis. corner ‘This article is protected by copyright. All rights reserved - transf collected L

between plants and birds (plant species*bird species, unit: interaction strength, strength, interaction unit: species, species*bird (plant birds and plants between ormed since these traits did not deviate from a normaldistribution. from a not deviate did traits these since ormed

fromthe literature literature

diameter (mm), fruit length (mm), plant height (m) and crop mass crop and (m) height plant (mm), fruit length (mm), diameter - dex indicates a wing shape which enables birds to fly to long over birds whichenables wing shape a indicates dex corner analysi corner

(Dunning 2007 (Dunning -

animal matching across sites, we used a modified useda we sites, across matching animal s was initially proposed to measure and test the testthe to and measure wasproposed s initially - frugivore networks in the Peruvian Andes Peruvian the in networks frugivore )

. Trait data for all eight all for interaction data . Trait Fruit length, fruit diameter and fruit fruit and fruit diameter Fruit length, et al. et ree individuals ree

1997, Dray and Legendre 2008). 2008). Legendre Dray and 1997,

at least four least at ’ .

, depending on on depending , bird traits. used They bird , Leisler Leisler ,

museum

and

Braak Braak the two of larger selects the and permuted) matrix are theinteraction of species (bird 4 modeltype and matrixpermuted) are interaction ofthe species (plant 2 type model output of L and R,Dray in ‘ade4’package in (fourthcorner model 6 type permutation matrix interaction trai bird between relationship removethe matrix interaction the in plant identities permutating by interactions observed and plant traits matrix (i.e., correlated null hypothesis testingthe for (i.e., length fruit width, the c that traits) quantifies matrix fourth a parameters of the estimate to then analysis is fourthcorner ofthe objective matrix to a compare 2007) Jordano and Bascompte species; plant each to species offrugivore visits ofa theproportion Acceptedcompleteness. sampling by we and estimator, Chao the on based link richness, asymptotic links the by ofobserved Dehling witht increases networkanalyses in correlations meta weightedthe networkand the wedefined of z Fisher's weapplied analysis, coefficients correlation the Pearson’s across A entire fourth Article r For each network, we tested four pairs of plant and bird traits (Dehling (Dehling traits bird ofplant and pairs four wetested network,each For We extracted the Pearson’s correlation coefficients coefficients thecorrelation Pearson’s extracted We

(escalc in ‘metafor’ in (escalc -

et al. et corner analyses for each of the eight networks. To generalize the individual fourth generalizethe individual networks.To eight ofeachthe for analyses corner L R ndean range, we performed a metaa we performed range, ndean et al. et

to of plant traits (plant species*plant traits) with a matrix matrix witha traits) species*plant (plant ofplant traits

both plant traits plant traits both 2012).

2014). Second, we quantified the sampling completeness of each network by dividing the number number the networkdividing by each of completeness sampling the Second, we quantified 2014). square X

≠ 0). Null model testing is complex because it involves the relationships of the interaction ofthe interaction relationships it involves the because testing is complex model ≠Null 0). L

vs.

(Dray and Legendre 2008, Ter Braak Ter 2008, Legendre and (Dray

- bill length, plant height plant length, bill root of the total number of interactions observed in each network as sample size for each for samplesize networkas in each observed ofinteractions number the of total root

‘This article is protected by copyright. All rights reserved package in R, Viechtbauer 2010) R,Viechtbauer in package orrelation between pairs of plant and bird traits. bird and ofplant pairs between orrelation

R

- and bird traits bird and transformati - analysis by this measure of sampling effort since the accuracy offourth accuracy the since effort sampling measureof this by analysis X

= 0) to test whether the traits of plants and birds weresignificantly birds and oftraitsplants whether the test to 0) = ts and the observed interactions by permutating the bird identities inthe bird the permutating by interactions thets and observed - on to the correlation coefficients coefficients correlation the to on analysis (metacor.DSL in ‘ in (metacor.DSL analysis r Q

vs. of the eight individual fourth individual the eight of . A proposed solution is to first remove the relationship between between relationship the remove first is to solution A . proposed he total number of observed interactions (see Appendix S2 in S2 Appendix (see interactions numberofobserved hetotal

Kipp’s index, crop mass crop index, Kipp’s r

for each trait pair from the matrix from the pair trait each for

. We performed this meta performedthis . We et al. et

2012). This approach has been implemented as implemented been has approach This 2012). Q

of bird traits (bird species*bird traits). The The species*bird traits). (bird traits bird of P metacor’ package in R, Laliberté 2011) 2011) R,Laliberté in metacor’package

values of the two permutation types ( types two permutation ofthe values

vs. et al. et - egendre 2008), which combines the the combines which 2008), egendre corner analyses corner to approximate a normal distribution normaldistribution a approximate to

body mass. W mass. body

2014): fruit diameter 2014): - analysis in two ways.First, two in analysis ighted the metathe ighted ’ . -

corner results for thefor results corner X X . Prior to the metato the Prior . e used a null model null useda e

(plant traits* bird traits*(plant bird L of the individual ofthe individual

and then to to then and vs. vs. - analysis analysis - corner corner bill bill Ter Ter - repeated the analogous procedure for the plant species and here defined the functional originality of plant species species ofplant originality functional thedefined here and species thefor plant procedure the analogous repeated i species bird few other only that by traits characterized species mostlywithplant interacted (Dehling centroid the and community between distance as the Euclidian measured then was originality network.Functional the respective in present species bird ofall centroids meaninteraction ofthe unweighted inplant trai species bird a of interactioncentroid weighted a in resulted This 2007). Jordano and (Bascompte strength interaction respective the by weightingthem species plant these of the coordinates PCoA averaged We network. withthe respective in interac species bird specific a that plant species all we selected network, each in species originalityofbird functional the determine to networks.Second, all across plant traits variationin thetotal represent that fouraxes Analysis(PCoA). Coordinates usedinPrincipal and a werecalculated plant species all matrix trait insingle a werecombined networks ofall plant traits First, with interact species few other only that with partners interact originality functional the wecalculated specialization, functional for measure As a specialization Functional Accepted effect to random a identity’as ‘network included models All amongtraits. comparable are modelestimates t were standardized birds and plants of traits functional to model fitting, variables. Prior as predictor traits bird or fourplant functional the and plantsand in R,Bates package models (‘lme4’ effect mixed fittedlinear we mass), body and wingshape width, length, bill (bill traits mass) bird and crop height and f diameter, plant (fruit functional four ofthe importance relative identify the To specialization. that whichensures networks, across 1 ofdeviation standard a mean and a of0 to scaled were values These ofvalues. normaldistribution a approximate andshape infour a Article We aimed at identifying which plant and bird traits were most closely associated with species’ functional functional withspecies’ weremost closelyassociated traits bird whichplant and aimed identifying at We - dimensional bird trait space based on the PCoA coordinates derived from bill width, bill length, wing length,bill width, from bill derived theon coordinates PCoA based space trait bird dimensional

body mass. Functional originality values of plant and bird species were log species bird and ofplant values originality Functional mass. body birds. In the two overall models, we thus fitted species' functional originality as the response variable variable the response as originality functional fittedwespecies' models, thus Inthe two overall birds. ‘This article is protected by copyright. All rights reserved

t space (Dehling t space o a mean of 0 and a standard deviation of 1 across networks which ensures that that whichensures networks across 1 of deviation standard a omean and a of0 et al. et modelestimat et al. et et al. et

2016). A high functional originality value indicates that a bird bird a that valueindicates originality A functional high 2016).

2016). Third, we calculated the community centroid as the centroid wethe community calculated Third, 2016). 2015) across all species in the eight networks, separately for separately networks, eight inacrossthe species all 2015) es were comparable between plant and bird species. bird and plant between comparable were

(Dehling (Dehling

the interaction centroid of a specific bird species species specificbird ofa centroid theinteraction . P et al. et airwise Mahalanobis distances between distances between Mahalanobis airwise

2016) for all plant and bird species. species. all for bird plant and 2016) , the extent to which species to the extent , - transformed to to transformed

This analysis resulted in resulted analysis This ’ .

nteracted with. We with.We nteracted ruit length, plant ruit length,

ted ted - and weighted meanweighted and variable) predictor respective the including models ofall weights Akaike the summed (i.e., importance variable theobtained We 2002). Anderson Burnhamand (AICc, sizesmall for sample corrected informationcriterion mass( body lengthbill and ( length bill width and bill for wascase this species the bird for ( fruit length and fruit diameter plant species, For werecombined. |0.5| than higher coefficient withcorrelation a wheretraits models exclude av models.To ofthe global 2016) network. in the respective observed species each for originality functional of estimates we weighted networks, among effort sampling correct in differences for to Inorder parsimony. of interest in models the finalfrom the terms wethese omitted results identical inresulted qualitatively higher also initiallyincluded We network. moreone inthan occur could species a because random as a effect was included ‘species identity’ werederiv originality functional of thatestimates the for fact account AcceptedInnetwork A1).the individual (Table results qualitativelyidentical showed completeness ( mass mass body vs. crop and highest width( vs. bill diameter differencethe major co in no was there expectation, our Contrary to 2). (Fig. traits frugivore ofplant and pairs all for matching signal oftrait meta The Trait matching Results 2016). Barton R, in package ‘MuMIn’ model.avgin from derived modelestimates (full value AICc lowest with modelthe Article To identify the best model, we first fitted all possible submodels (dredge in ‘ (dredge submodels wefittedpossible all first model, identify thebest To

correlations - analysis across the eight Andean interaction networks, weighted by sampling effort, showed a strong showedstrong a effort, weightedsampling by networks, Andeaninteraction the eight across analysis

model coefficients across all models with a ΔAICc value smaller than 2 relative to the best the best to relative 2 than smaller value withΔAICca models all across modelcoefficients across networks across r ‘This article is protected by copyright. All rights reserved

= 0.32, = r

- = 0.84). We compared the resulting set of sub setof compared the resulting We 0.84). = rrelation strength between traits related to size matching and foraging strategy. Fruit Fruit strategy. foraging matchingand size to related traits between strength rrelation level taxonomic information (i.e., genus and family). However, because thi because family).However, genus and (i.e., information taxonomic level r r

= 0.21, =

= 0.74) and crop mass and fru mass crop and and 0.74) = P oid collinearity among predictors, we constrained the resulting set of models to to models setof wethe resulting constrained predictors, collinearity among oid < 0.001) and plant height vs. wing shape ( shapewing plant height and vs. < 0.001)

. The correlations be correlations The .

by sampling effort, i.e., the square i.e., effort, sampling by

P = 0.02) were slightly weaker. The meta weaker.The were 0.02) slightly = ( vs. length bill tween length fruit it diameter ( it diameter r

= 0.90), bill width and body mass ( masswidth body bill and 0.90), = ed from different networks. In addition, Inaddition, networks. from different ed - root of the total number of interactions ofinteractions number the of total root - r models acc

= 0.54) were highly correlated, while were highlycorrelated, 0.54) = r

= 0.33, = MuMIn’ package in R, Barton R,Barton in package MuMIn’ - analysis weighted by sampling sampling by weighted analysis

ording to the Akaike the to ording P ’ . < 0.001) showed the< 0.001)

s, correlation correlation s, r

= 0.26, =

P r

= 0.85) and and 0.85) = < 0.001) 0.001) < s Tityridae, Argentina). Argentina). Tityridae, and Blue 1), Peru Ramphastidae, ( Ruddy( Pigeon 2), Colombia Trogonidae, Quetzal ( headed Argentina). (Verbenaceae, 1), sp Schefflera grandiflora Alchornea species in the plant werefound specialization functional of the degrees highest networks, analyzed Inthe eight specialization Functional m signal oftrait mass thefor crop meta in than the variable morewere matching oftraits pairs the between coefficients 0.03, = (b 0.10, = (b length ofbill thewhereas effects ofbirds, originality functional of thedegree Inthe averaged 1.84). = (ΔAICc the and second 0) = lengthbill (ΔAICc wingand shape included a had modelstwobirds, For 3a). (Fig. was 0.57) not significant = originali functional of thedegree 0.69) = heightmass (ΔAICc plant crop and included a w models two we plants, identified For (Fig.3). species bird amongplant and ofspecies, originality functional i birds most common usually are the these although Passeriformes

Acceptedprasinus Aulacorhynchus Article ΔAICc value smaller than 2; the best model included only crop mass (ΔAICc = 0) and the second the and mass 0) = (ΔAICc crop only modelincluded thebest 2; than smaller ΔAICcvalue Clusia elliptica Clusia We identified specific plant and bird traits that explained the differences in functional specialization, i.e. specialization, functional in differences traitsthe that bird explained plant and identifiedspecific We - black Tanager ( Tanager black

P = 0.62) were not significant (Fig. 3b). were (Fig. 0.62) not= significant . (Araliaceae, Ecuador 1), 1), Ecuador (Araliaceae, . atching across networks. across atching Pharomachrus auriceps Pharomachrus - body mass relationship, trait correlations were always positive, consistent with the overall withthe overall consistent positive, werealways correlations trait massrelationship, body

(Clusiaceae, Peru 2), Peru (Clusiaceae, Tangara vassorii Tangara The majority of the functionally most specialized bird species did did species bird specialized most functionally majorityofthe The

(Euphorbiaceae, Colombia 1), Colombia 1), (Euphorbiaceae, ‘This article is protected by copyright. All rights reserved

, Ramphastidae, Ecuador 2), Chestnut 2), Ecuador Ramphastidae, ,

The most functionally specialized bird species in the networks were the Golden werethe in thespecies networks bird specialized most functionally The - banded Toucanet ( Toucanet banded ty of plants (b = 0.23, = (b ty ofplants model, the pointedness of the wing (b = 0.15, wing= (b ofthe model,the pointedness , Thraupidae, Bolivia 2), and Crested Becard ( Becard Crested and 2), Bolivia Thraupidae, , Isertia laevis Isertia

Symplocos arechea Symplocos , Trogonidae, Colombia 1), Masked Trogon ( Trogon Masked 1), Colombia Trogonidae, , Patagioenas subvinacea Patagioenas Aulacorhynchus coeruleicinctis Aulacorhynchus

. In the averaged model, crop mass was positively related to to related was positively mass model,crop In the . averaged (Rubiaceae, Ecuador 2), Ecuador (Rubiaceae, Oreopanax caricifolium Oreopanax

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< 0.01), whereas the effects of plant height (b = 0.04, = (b height whereas ofplant the 0.01), effects <

(Symplocaceae, Bolivia) and and Bolivia) (Symplocaceae, - - tipped Toucanet ( Toucanet tipped best model included wing shape and body mass wingbody and shape modelincluded best ΔAICc value smaller than 2; the best model thebest 2; than smaller valueΔAICc n seed dispersal networks. networks. dispersal nseed , Columbidae, Ecu Columbidae, , Endlicheria sp Endlicheria

(Araliaceae, Colo (Araliaceae, P , Ramphastidae, Peru 2), Blue 2), Peru Ramphastidae, , = 0.03) was positively related to to related positively 0.03) was = Aulacorhynchus derbianus Aulacorhynchus Pachyramphus validus Pachyramphus - ador 1), Emerald Toucanet Emerald Toucanet 1), ador analyses (Fig. A3). Except Except A3). (Fig. analyses Trogon personatus Trogon

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P on bird species that have a sufficiently large gape width that enables width that gape large sufficiently a that have species bird on 1 (Wheelwright networks interaction in wellecological matching studied is size to related oftraits importance The networks. in plant species animal matchingplant and of the determined foraging, animal to matchingand size metaOur Andes the across Trait matching specialization. functional ofavian the degree to related was positively wing ofthe thepointedness while ofplants, specialization offunctional the degree to related positively wasmass that plant crop we detected Nevertheless, networks. were traits animal plant and Specific Andes. shape matching( size to related traits animal thatplant and func matchingand oftrait degree the thattraits determine plant combined eight We Discussion Accepted( Trogon Masked the Golden wingsexample, were,for pointed foraging(Schaefer while fruits encounters species bird a where influences directly therefore Wingshape Denslow1985). and (Moermond canopy the foraging and in whi forest understory, move into dense wings Rounded allowbird a 1988). (Rayner maneuverability and movement capacity its wingto is related bird’s were (Olesen interactions ecological matc size that hypothesis in the has resulted This 2007). Warren and Woodward Maglianesi with short prob a pollinators bird insect and Article In contrast to our initial expectation, we found that traits related to foraging strategies of frugivorous birds, birds, frugivorous offoraging strategies to related traits wethat found expectation, initial our to Incontrast

equally important as trait pairs related to size matching in matchingsize to related pairs trait as important equally

and -

analysis across eight Andean interaction networks showed that plant and animal tr animal and that plant showed networks Andeaninteraction eight across analysis crop mass crop

et al. et

985, Stang 985,

2015), and small predators are usually unable to feed on large prey (Cohen preylargeon feed (Cohen to unable usually are small and predators 2015), Trogon personatus Trogon - body mass body ‘This article is protected by copyright. All rights reserved - frugivore interaction networks, sampled along the Andean range, to identify functional to Andeanrange, along the sampled networks, interaction frugivore et al. et et al. et

) were important in shaping plant shaping wereimportant in )

2009, Garibaldi Garibaldi 2009, , Trogonidae). These birds were also among the most functionally functionally most were the among also birds These , Trogonidae).

2011b). The results from our meta from resultsour The 2011b). le birds with pointed wings are better equipped for longfor equipped better wings are with pointed birds le oscis or bill are constrained to visit short flowers (Stang flowers short visit to are constrained bill or oscis only only - headed Quetzal ( Quetzal headed et al. et weakly associated with species' functional specialization across across specialization withfunctional species' associated weakly fruit size fruit

2015). For instance, bird instance, For 2015). et al. et

tional specialization of plants and animals. We found found animals.We and ofplants specialization tional

2002). In our Inour 2002). - beak size beak Pharomachrus auriceps Pharomachrus

shaping - frugivore

them - ) and animal foraging( animal and )

interaction networks. The shape ofa shape The networks. interaction analysis support this hypothesis. this support analysis

to swallow the fruit (Levey 1987), 1987), (Levey fruit the swallow to networks, bird species with species rather networks,bird

interaction networks across the across networks interaction - dispersed fruits strongly depend depend fruits strongly dispersed hing is a general principle in principle general is a hing , Trogonidae) and the and Trogonidae) , ’ .

aits, both related to to related both aits, et al. et plant height plant - distance flights flights distance

1993, 1993, -

frugivore frugivore et al. et

2009, 2009, - wing and the dominance of specific trait values in a community, the detectability of trait matching may differ among mayamong differ matching oftrait detectability the community, a values in trait specific of the and dominance deg Depending2).the on g (Ecuador 1180 to Bolivia) & 2, 1 g (Peru from 706 varied size maximum body instance, For pools. widelytheirspecies in differed that wecommunities compared Moreover, withthe matching trait increases detect to likelihood among completeness and effort in sampling amongnetworks al. while small place, one mass crop with large a plants mostlyon feed therefore energy on Petchey Belovsky1997, 1993, Davies and wingshape Swartz 1987, Rayner and in (Norberg bats foragingbehavior in birds specialized Acceptedplant tra singleidentify to a wewereable Nevertheless, networks. ofstudied the range wasweakacross traits overall bird plant and and specialization functional between werelationship the that found networks, across specialization that large we expected While specialization functional traitsand functional between relationship The fromspecies generalizations against Andea the across exemplified here matching, oftrait importance the overall not question variability does This species. animal plant and matchingbetween oftrait degree the in fluctuations result also in res in fluctuations flexibly to respond can species consumer small by dominated thatare communities ecological modified (Dehling networks Article

2014, Howe 2014, 2016 The relationship between crop mass and body mass is consistent with the theory of optimal foraging (Krebs foraging (Krebs optimal theoryof with the ismass consistent mass body and crop relationshipbetween The Despite the overall significant relationships between all trait pairs, the degree of trait matching differed matchingdiffered oftrait the degree pairs, trait all between relationships significant the overall Despite it (crop mass) and a single bird trait ( trait mass)single bird a and it (crop - rich resources to meet their energetic demands (Corlett and Primack 2011). Large 2011). Primack and (Corlett demands meetenergetic their to resources rich on interaction networks networks interaction on , especially in the case of the crop mass crop ofthethe case in especially , et al. et the networks. Wing shape has shape networks.Wing the , Muñoz ,

bodied

‘This article is protected by copyright. All rights reserved 2014). For instance, we would expect that the degree of trait matching is weakened inweakened matchingis oftrait degree that the expect would weinstance, For 2014). -

et al. et fruited plants and large and fruitedplants frugivores frugivores

2017 is a general phenomenon general isa - rich to species to rich ). tend to tend

et al. et the studied studied the wingshape

2008). According to this theory, large bodied bird bodied According large theory, this to 2008). visit several individuals of plants with small crops small with ofindividuals plants several visit likewise likewise - poor ecosystems. poor

number of observed interaction events (Dehling interactionevents ofobserved number - billed bird species show the highest showthespecies bird billed , which offer a high amount of resources accumulated in in accumulated resources of higha amount offer which , networks - ) that were significantly associated with functional wereassociated significantly that )

body mass relationship. Th massrelationship. body been shown to influence flight maneuverability and and maneuverability flight influence to shown been ource availability (Bender (Bender availability ource

et al. et across different types of plant of different types across - bodied species (Saavedra (Saavedra species bodied . Previously .

2003). This suggests that the suggests that This 2003).

,

it has been shown that shown been it has i s could be due to to due differences be could et al. et ’ n range, but cautions range,but cautions n et al. et .

ree of trait variation of trait ree level of functional functional of level

- 2017), which willwhich 2017), frugivore networks. frugivore - bodied frugivores frugivores bodied

2014). Moreover, Moreover, 2014). influence s need to feed to sneed

(Dehling

the the et al. et

of

2014). 2014). et

are able to fly long distances (Moermond and Denslow 1985, Dehling Denslow1985, and (Moermond fly to able distances are long forest and ofthe canopy inwings feed the usually with species pointed Bird phylogeny. species' of independent t is related specialization functional that indicating groups, different taxonomic across werescattered specialization withoffunctional degree high species a bird networks, Across community. crops. small with than plants dispersers seed moredistinct functionally potentiallyrequire, and attract, crops withlarge with species less bird g massm208 2,= body (Peru 3000 communityat Blue For instance, 1993). Davies and (Krebs demands highenergy large attract might especially crops s with functional plant associated are taxonomy, species than rather values, trait thatspecific suggests finding this network, in each specialized most functionally were the plant species oth to compared species setofbird distinct morphologically a mass attracted crop large a produced that that plants indicate results Our specialization. Accepted0.34 wasa there nor pair, trait the same to large (e.g. matching traits by wascorresponding determined birds and ofplants specialization forests. tropical for consequences functional severe havesimilarly (Galetti large thelossof that highlighted have studies previous While specialized. most functionally were asthey species types ofplant (Müller landscapes fragmented in forest patches connecting for important particularly are species whysuch reason th Article ese species can provide long canprovide species ese Since we found a high degree of higha degree found weSince wingswereide with species pointed Bird , - - banded Toucanet banded P billed bird species). However, the identified plant and bir plant and the identified However, species). bird billed

< 0.0< et al. et al. et 5

2014). According to our a Accordingour to 2014). 2013, Vidal Vidal 2013, ) or wing shape and body mass (n = 304 species, 304 = mass (n body and wing ) or shape - bodied animal species from tropical forests has sever forests fromtropical animal species bodied

distinct traits, such as the many tanager species. Hence, our findings suggest that plants thatplants suggest findings our Hence, species. manytanager suchas the traits, distinct

( Aulacorhynchus coeruleicinctis Aulacorhynchus ‘This article is protected by copyright. All rights reserved et al. et -

distance dispersal to plant communities (Böhning plant communities to dispersal distance 2013), our results indicate that the loss of bird species with pointed wingsmight with pointed species bird the lossof that results our indicate 2013), - bodied birds (Dehling birds bodied

trait matching between individual trait pairs, we expected that functional functional that weexpected pairs, trait individual matchingbetween trait high nalyses, these species also provide seed provide also species nalyses,these correlation between plant height and crop mass (n = 243 species, species, 243 (nmass= crop height and plant between correlation er plant species in the community. Since phylogenetically unrelated unrelated phylogenetically Since the community. in plant species er ntified to visit the most distinct set of plant species inthe species setofplant most distinct the visit to ntified ). In contrast, plants with small fruit crops are mostlyby visited are crops fruit small with plants In contrast, ). ), one of the largest frugivorous birds in the Peruvian bird bird in the Peruvian birds one frugivorous ), ofthe largest

et al. et Clusia elliptica Clusia d traits, crop mass and wing shape, did not belong did wingshape, mass crop traits, and d

r 2014) that rely on such resources to meet their to that suchresources rely 2014) on

= 0.22 =

e consequences for the plant community plant community the for consequences e et al. et , P < 0.0< pecialization. Plant species withlarge species Plant pecialization.

2014). Due to their Dueto flight ability, 2014).

(Clusiaceae) interacted wi interacted (Clusiaceae) o specific trait values largely values largely trait specific o - 5 dispersal functions to specific specific functionsto dispersal - ). ). This Gaese Gaese ’ , et al. et indicates that there are are indicatesthat there

. large

2006). is the This 2006). - fruited plants and plants and fruited th the r

= (Kissling and Schleuning 2015) Schleuning and (Kissling communities ecological novel in traits functional species' from interactions unknown predicting for base frugivory in birds and plants between frequencies interaction shaped foraging animal matchingand size to related traits range that geographic large founda across We Conclusion communities. seed valuable particularly likely provide are to species bird These 1985). Denslow and (Moermond fruit types of specific reach thus, to are, able and longerdistances move to able over than fruitof tracking strategy followdifferent wingsa withpointed canopy communities, Inbird 1982). Smallwood and (Howe community the in plant species other fr might benefit birds offrugivorous group functionally distinct a attract mass and crop large a into invest that species the fewIncontrast, plant 2012). mostly fruitsare their into littleenergy comparatively that invest thosespecies Inplant communities, communities. bird in plant and withspecialization functional associated are that oftraits the types in important differences AcceptedFast byFund Marsden a supported was also DMD Arts. the and Research, Education, ofHigher ‘LOEWE grant.K.B. starting (UvA) Amsterdam of University a acknowledges W.D.K. 118). (FZT Foundation Research theby German funded t of support the acknowledge gratefully We manuscript. our on feedback theirconstructive for reviewers thank two anonymous We Acknowledgements communities. identifi the for useful is traits ofthese identification The traits. morphological with specific associated but significantly weakly, is birds and plants of specialization of functional the degree that we show Furthermore, climatechange. Article

cation of plant and bird species that fulfill functionally specialized roles in current and future and incurrent roles specialized functionally fulfill thatspecies bird plant and of cation dispersed by similar types of often small ofoften types similar by dispersed —

Landes

- Offensive zur Entwicklung Wissenschaftlich Entwicklung zur Offensive

‘This article is protected by copyright. All rights reserved - G., D.M.D., E.L.N. and M.S. received support from support received M.S. and E.L.N. D.M.D., G., he German Centre for Integrative Biodiversity Research (iDiv) Halle (iDiv) Research Biodiversity Integrative for Centre German he , for instance as a result of species' migrations and local extinction local and migrations species' instance offor , result as a omlong - distance dispersal or from dispersal to different locations relative to to relative locations fromdifferent to or dispersal dispersal distance - bodied bird species (McKey 1975, Kessler 1975, bird (McKey species bodied

networks. The generality of trait matchingthe is trait generalityof networks.The - ökonomischer Exzellenz’ of Hesse’s Ministry ofHesse’sMinistry Exzellenz’ ökonomischer

species with more rounded wings as they are wingsare they as withrounded more species - dispersal functions to pla functionsto dispersal the research funding program program funding research the ’ .

- Rios and Kattan Riosand - Start Grant Grant Start - foraging birds birds foraging - s in response to to sin response nt Jena - Leipzig Leipzig

Accepted Article BO1221/20 825/1; (PAK Ecuador” South in Research and Monitoring 823 ResearchUnit the framework of in the (DFG) Foundation Colombia). de Republica Innovacióne Tecnología Scho Student by funded was Colombia instudy de Tucumán. Nacional bywas Universidad the granted San Javier de Sierra in Parque research conducting 2009 (PIP CONICET by funded deAves de Ecología Laboratorio the (UOC - 1101) administered 1101) larship “Francisco José de Caldas”, COLCIENCIAS (Departamento Administrativo de Ciencia, de Administrativo (Departamento COLCIENCIAS deCaldas”, José “Francisco larship ‘This article is protected by copyright. All rights reserved

by the Royal Society of New Zealand. New Societyof thebyRoyal - 1025 and and 2014 1025 Rufford Small Grant for Nature Conservation (# 11042 (# Conservation Nature for Small Grant Rufford

-

IER for their help in the field in Argentina. in field in help the their for IER - 592) and ANPCyT (P ANPCyT and 592)

The study in Ecuador was funded by the German Research Germanwas the by funded study in The Ecuador

- 825

Furthermore, Furthermore, “Platform for Biodiversity and Ecosystem Ecosystem and “PlatformBiodiversity for ICT 2013 ICT - 1).

we are grateful to the membersof the to grateful we are - 1280). Permission for Permission 1280). The study in Argentina was Argentina in study The ’ .

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- ‘This article is protected by copyright. All rights reserved

Ecology 92:26 Ecology Ecomorphology of b Ecomorphologyof Ecol. Evol. Ecol. Revisiting the cost of carnivory in mammals. mammals. in Revisitingcarnivory of cost the

- specific interaction patterns and size matchingplant a in size and patterns interaction specific Vertical stratification and caloric content of the standing fruit crop in a tropical intropical a fruit crop the standing content of caloric and Vertical stratification -

300. 2017. 2017. - - eating birds and bird and birds eating size, gape width, and the diets offruit width,diets the and gape size, - - - dispersal net dispersal 1526. 1469.

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Oecologia 176:837 Oecologia

traits ponents of seed of ponents - environment relationships in the fourth the in relationships environment -

- 38:380 Ecography dispersal mutualistic networks. networks. mutualistic dispersal

- a tropical plant tropical a eating birds. birds. eating – -

J. Evolution. Biol. 29 Evolution. Biol. J. 848. -

dispersal networks. networks. dispersal -

- J. Stat. Softw.36:1 Stat. J. -

– pollinator interaction web. interaction pollinator Ecology 66:808 Ecology frugivore network differ networkdiffer frugivore - - o generalization. generalization. o 392. rich rich temperate zone. zone. temperate ’ .

seed :2181:2190. - dispersal dispersal -

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Accepted Article from scaling freshwaters: in interactions predatory Body and size 2007. P. Warren, and G. Woodward, s communities. to individuals tructure and function of aquatic ecosystems. Cambridge University Press, Cambridge, UK, pages 98 pages UK, Cambridge, Press, University Cambridge ecosystems. aquatic of function and tructure ‘This article is protected by copyright. All rights reserved -

In: Hildrew, A.G., Raffaelli, D. and Edmonds and D. A.G.,Raffaelli, In: Hildrew,

- Brown, R. (ed.), Body the size: R.(ed.), Brown, ’ .

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location in South America where they weretheycollected. where America in South location interac ofobserved number the and total species plant species, ofbird number the network,each For frugivorevisits. frequency of the width indicate line and m) (2500 and m), Bolivia (3000 m), 2 (1500 m),1 Peru Peru (2000 2 m), (1000 1 Ecuador m), Ecuador (2500 2 Colombia fleshy between 1. Fig. Legends Figure Accepted Article

Overview of the eight interaction networks sampled across the Andes. the across sampled networks interaction Overviewthe eight of

Argentina (1000 m). Interactions between species are indicated by lines between boxes; box size box boxes; between lines by indicated are species m).Interactionsbetween Argentina(1000 - fruited plants (bottom boxes) and frugivorous birds (top boxes): Colombia 1 (2000 m (2000 a.s.l.), 1 boxes): Colombia (top birds frugivorous and boxes) (bottom plants fruited

‘This article is protected by copyright. All rights reserved

tion events is given. The lines connect the networks to the to networks the connect lines given. The is events tion

Networks represent interactions interactions represent Networks ’ .

*, lev the indicate Asterisks coefficients. varianceofcorrelation ofz mean the indicate fourth analysisof 2. Fig. Accepted Article

P < 0.05). <

Matching of plant and frugivore traits across eight Andean networks. Results are derived from a meta from a derived Resultsare Andeannetworks. eight across traits frugivore Matching and ofplant

- corner corr corner - transformed Pearson’s correlation coefficients coefficients correlation Pearson’s transformed ‘This article is protected by copyright. All rights reserved elations between plant elationsbetween

- frugivore trait pairs, weighted by sampling effort. Triangles Triangles effort. weighted sampling by pairs, trait frugivore

el of significance for each trait pair (***, pair trait each for ofsignificance el r . Grey lines indicate the estimated theestimated indicate lines Grey . ’ .

P < 0.001; 0.001; <

- Accepted the square to proportional Articlenetwor inspecifica species bird a plant or a represents dot each modelofthe significance of the indicate level model best setof in the werenot included model w interact i.e. originality, functional measuredby was specialization effects.Functional random as identity’ ‘network and ‘species identity’ mass),including body and shapewing width, length, bill bill birds: height plant fruit fruit length, diameter, (plants: variables predictor as traits and variable, response mixed from obtained are Results respectively. relat thefor significant plots residual partial and birds frugivorous 3. Fig. -

averaged coefficients across a subset of best models (ΔAICc < 2). Grey estimates indicate variables thatvariables indicate Grey estimates 2). < models (ΔAICc best subseta of across coefficients averaged Relationships between morphological traits and species’ functional specialization in (a) plants and (b) (b) plants and (a) in specialization functional species’ and traits morphological Relationshipsbetween ith partners characterized by traits that only few other species interact with. Shown are results from from results are with. Shown interact species few other by only that traits characterized partners ith ‘This article is protected by copyright. All rights reserved - root number of observed interact ofobserved number root s. In (a) and (b), the relative variable importance is given and asterisks asterisks and is given importance thevariable relative (b), and Ins.(a) - - averaged estimates ( ***, ( estimates averaged effects models with functional specialization of plants or birds as birds or ofplants withspecialization functional models effects

ions in the respective network. the respective in ions k. Dot size and the degree of shading are are ofshading the degree and size k.Dot ionships in (c) plants and (d) birds, birds, (d) plants and (c) in ionships

P < 0.001; *, 0.001; <

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P the extent to which species to extent the ’ < 0.05). In (c) and (d), (d), and In(c) 0.05). < .

and crop mass; crop and