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Edinburgh Research Explorer Impacts of local adaptation of forest trees on associations with herbivorous : implications for adaptive forest

management

Citation for published version: Sinclair, FH, Stone, GN, Nicholls, JA, Cavers, S, Gibbs, M, Butterill, P, Wagner, S, Ducousso, A, Gerber, S, Petit, RJ, Kremer, A & Sch??nrogge, K 2015, 'Impacts of local adaptation of forest trees on associations with herbivorous insects: implications for adaptive forest management' Evolutionary Applications, vol. 8, no. 10, pp. 972–987. DOI: 10.1111/eva.12329

Digital Object Identifier (DOI): 10.1111/eva.12329

Link: Link to publication record in Edinburgh Research Explorer

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Published In: Evolutionary Applications

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Download date: 05. Apr. 2019 Accepted Article [email protected], [email protected], Ducousso: [email protected], Emails: France. Talence, F-33400 1202, UMR BIOGECO, 5. INRA, UMR1202 BIOGECO, F-33610 Cestas, France Universityand ofBordeaux, 4. Department ofZoology,of South University Bohemia, [email protected] Gibbs: Phil KarstenMelanie [email protected], Butterill: Schönrogge: [email protected], Centre forEcology OX103. and Hydrology, Wallingford UnitedEmails 8BB, Kingdom. [email protected] 2. Centre forEcologyEH26 0QB,and Hydrology,United Edinburgh Kingdom. Email: [email protected] Emails:Kingdom. [email protected]@ed.ac.uk 1. Instituteof Univer EvolutionaryBiology, Frazer H. Sinclair H. Frazer for adaptiveforest management implications Impacts adaptationof local of forest trees on associations with herbivorous insects: Article Research Original type: Article Type:Article Original Article Accepted Date: 20-Aug-2015 Received Date:17-Aug-2015 This article This protectedis by Allcopyright. rights reserved. 10.1111/eva.12329doi: lead to differencesthis between version and the Version of Record. Please cite this article as through pagination typesetting, maywhich thecopyediting, been andproofreading process, articleThis has been publicationaccepted for and fullundergone peer review but has not Petit Gibbs 5 , Antoine Kremer 3 , Philip Butterill Philip , 1,3 , Graham N. Stone N. Graham , 5 ,Schönrogge Karsten 3,4 , Stefanie, Wagner 1 , James A. Nicholls A. James , sity ofEdinburgh,EH9 Edinburgh 3JT, United 5 3 , Alexis Ducousso . Č eské Bud 1 , Stephen Cavers Stephen , 5 , SophieGerber ě jovice, CzechRepublic. 5 2 , Rémy J. , Melanie Accepted Article Abstract Running head: email: [email protected] ph: +44 131 6507194 EH9 Edinburgh 3JT,UnitedKingdom. of Edinburgh, Corresponding author [email protected] Kremer: Antoine [email protected], Remy Petit: [email protected], This article This protectedis by Allcopyright. rights reserved. type showed the relationship between abundance and phenological mismatch predicted abundancephenologicalbetween mismatch and showedtherelationshiptype gall no but abundance, gallwasp in variation significant explained Provenance supported. were hypotheses Both oaks climate-matched between for mismatch trees. nativephenological and local hypothesis,adaptation we predicted declining gallwasp abundance with increasing Under betweendifferences a origin site. increased trial and locallatitudinal with oaksthat to distanceincreasing phenological betweenoriginand provenance site, trial differences and(ii) with growth performance showwould non-native that (i) declining provenances hypothesised sessileprovenances( of oak these impacts using datafor abundance local cascadingbecause ofpotential communities. onlocally adapted impacts explored We currently predicted future experiencing populations non-localby from populations input nativeof local tree augmentation matching– the Climate imposed by strategy One times. limitations generation mitigation longis such impacts for involving maytrees Interactions bepar forest Disruption of species interactions is a is species Disruption of a interactions Oak provenanceOak affectsherbivore on abundance : Prof. Graham Graham Stone, Institute : Prof. of Quercus petraea native gallwasp herbivores sampled from 20 from sampled herbivores gallwasp native ticularly vulnerable due to evolutionary rate rate due to evolutionary ticularly vulnerable ) planted in a common garden trial. Wecommon in a garden trial. ) planted key issue in climate change biology. biology. change climate in issue key climates. This strategy is controversial

Evolutionary Biology, University Accepted Article Introduction Introduction involves the following steps: (i) identify climate variables that predict performance for the for performance that predict variables identify climate (i) following steps: the involves McEwan , fungi andmicrobes (Kennedy andSouthwood 1984;Lindenmayer are (and foundation also keystone)often species associatedfor communities of plants, (Lindner adaptation of low rates result in trees particularly sensitive to imposedpressures by and Yohe 2003;(Parmesan Walther responsesdriving population whose influenceeffects can community structure and function across Ecosystems the experien world are levels (Frelich levels matching, adaptive forest management, provenance trials, Keywords and variable impacts on oak gall communities. by a local hypothesis.adaptation Our showresults that matchingclimate would have complex This article This protectedis by Allcopyright. rights reserved. Savolainen arestands natural forest locally to adapted local current (Albertoclimates al. 2005;but see Cavers andCottrell 2014). Clim commercial forestryfor potential where predicted futurepredicted (Dawson climates to adaptation forest hasstrategy facilitating emerged onepotential as climate matching, Broadmeadow face of changeclimate (Aitken management strategies topres forest adaptive et al. et : local interactions,adaptation,: local climate plant- non-independence, population

et al. 2013;Savolainen et al.

et al. 2005, Spittlehouse and Stewart 2003). Assisted oftheform 2003). in and Stewart migration, 2005,Spittlehouseet al. 2011), effects on arethem to cascade expectedthrough associated trophic et al. 2012). Consequently, there is c there is Consequently, al. 2012). et

et al. 2008; Bolte 2008; al. et

et al. 2007), and for a given species and planting locationandgiven planting a al.species and for et 2007), et al. 2011; Hoegh-Guldberg

et al et in situ . 2002). Forest ecosystems are expected tobe cing profound changeclimate 2013), (IPCC, erve ecosystem productivityand servicesthe ecosystem in erve

climate because the long generation times of et al. 2010). Furthermore, because forest trees forest because Furthermore, 2010). al. et adaptation may be limited (Broadmeadow limited be may adaptation ate matching isbased on thepremise that

et al. 2009; Bower and Aitken 2008;al. 2009;BowerandAitken et

onsiderable interest indeveloping Quercus petraea,

et al. 2008), particularly in et al. 2008), gallwasp. gallwasp.

et al. et al. 2013a; 2014;

et Accepted Article responses tosuch variation. phenotypic traitsrelevant toherbivore success, and the of magnitude herbivore population 2007; Visser, The 2008). issues key are the host towhich extent plant provenances vary in vanVisser phenology 2013; to andKarban Aschand plant herbivore adaptation (Pearse provenancestree (Bernhardsson native studiespred suchchanges.some Whileof species (Lefevre matching drive abundancestrategy intheidentity, associated changes andgeneticsmay of aclimatepart of genotypesas tree theintroduction Roslin new 2010).As of and such, Tack (Bernhardsson shownadaptation have studies of range diversity of the host diversity ofthe trees (Dungey thathas shown theorganisms associated with standsforest reflect the genotypes genetic and resilience,but also impactsits onlocal associated biodiversity. Community genetic research be considered notonly intermsof its effect strategyother that thegenetic,modifies physical, 2014). Forests are biologically diverse ecosystems subjects remain of vigorous debate (Dawson adaptation (Broadmeadow localaccelerates subsequent that adaptivevariation andcontribute climates, tofuture matched well be will trees thatresulting expectation the forestry) in in (termed provenances locations thepredictedexperience (iv) tree seedfrom conditions; populations plant identified in the proposed range (iii) currently plantingat identifythe species the within that site; locations re use (ii) range; its across species tree target This article This protectedis by Allcopyright. rights reserved. As remediation strategies, climate matching and other forms of assisted migration migration assisted of forms other and matching climate strategies, remediation As

et al. 2013; Egan andOtt 2007; Hanks and Denno 1994; Sork

et al. 2014). However, it is difficult to forecast the direction and magnitude and direction the toforecast isdifficult 2014). it However, al. et

et2005). al.

et al. 2000; Whitham al. et

et al. 2013),others suggest et al. opposite the through local

ict increased ofabundance herbivores on non- of of plants to localherbivores and et al. 2011; Lunt al. 2011; et gional climate models to predict future climates future predict to climate models gional iveness promoting in forest adaptation and or spatial structure of tree populations, must structure tree populations, of or spatial , ofvalue and the, matching, or climate any

et al. 2012; 2012; et al. Wimp

et al. 2013; Cavers and Cottrell Cavers and 2013; al. et

et al. 2005), and a and 2005), al. et

et al. 1993; al. et vice versa

Accepted Article latitude andlongitude asthemismatch in provenance trial as between andlatitude well climate the CharnieLa the Petite provenance trial, selected the availableto maximise range insource measures at test 20provenances hypothesis planted performance for this We using multiple beshould least matched well to conditionscurrent and least perform in so well the present. provenances predictsthat selecthypothesis also lived trees are already somewhat mismatched grow. toin whichthey The the conditions populations of long- that mean may change climate recentrapid – overand time decline will Lefevre they prevailingto which to are adapted (Broadmeadowconditions climatic climate matching. gallwasps taand make sentinel appropriate an forest and are exposed to native herbivores exposed tonative andare forest Northwestwhere France,provenances 96 of anestablishe exploits Ourapproach herbivores. oak, European widespread andimportant phenology) an of and (growth performance the phenotype on(i) provenance of Mopper 2005; Mopper Mopper associated local tospecific populations host adaptation with Ott 2007; plant (Egan and aspects of gallwasp These biology, withare lifecycle, coupled apartly parthenogenetic phenology tohostplant 1988). andAkhteruzzaman highly sensitive (Askew1962;Crawley tissuesoak atthe correct developmental stage (Harper hosts (Harper Cynipidae),(; specialised herbivores larvae whose their oak in develop on This article This protectedis by Allcopyright. rights reserved. We expectWe theperformance of oak provenances to correlate positively with extent the Here we the communityexplore of effects climate matching by theimpact quantifying

et al. 2014).et al. This hypothesis locally predicts of thatnative provenances performance

et al. 2004; Stone and Schönrogge 2003). Ga Schönrogge2003). 2004; and al. Stone et

et al. 2000; Tack 2000; et al. Quercus petraea

et al. 2010)et al. 2007), andOtt andindividuals(Egan . Our study focuses on oakstudy cynipid on gallwasps. Our focuses , and, theabundance of (ii) associated insect xon for exploration of of the effects potential xon exploration for Q. petraea d INRA forestry trial at La Petite Charnie in Charnie Petite La at trial d INRA forestry ed to match conditions furthest in the future inthe conditionsfurthest match ed to

et 2004), al. andoakcynipids are grow surrounded by natural oak by natural growsurrounded ll induction requires access to hostinduction access requires to ll

et al. 2005; Accepted Article reproductive success in herbivores and associatedsuccessand inherbivores reproductive to is central leaf-eating herbivores) for (budburst availability plantresource with high between trees andassociatedforest organisms. synchronization of Forexample, breeding types. Here we test bothhypotheses using abundance compounds. defensive concentrationsof reduced and content nutrient through elevated mean) organs plant with(those high orlarge growth rates size relative to ultimate the population increased1991) predicts herbivore performance 2009).In 1974; White, the contrast, tissues an and associated resource increase in quality (Whiteherbivores for 1969;White mobilised tohigher instressed of plantphysiologically nitrogen stressed levels plants, due hypothesis The inperformance among toobservedabundance variation provenances. oak provenance the origin.then We test twocont This article This protectedis by Allcopyright. rights reserved. enemies(Kerstesde Jong2011; Thomson natural and of fitness with associated impactscues, onthe might occurby to local of the adaptation disrupting photoperiodany and thermal combination latitudes lower latitudes.from As a result, phenological mismatches with localherbivores matchingclimate because, ina warming world,it often involvesintroducing tohigher plants (Laube temperature and between photoperiod interaction by (Aitkenoften determined cues photoperiodic onsetwhose is budburst, clines timingof inphenological latitudinal showevents, as the such 1996;van al. et Asch plant host to show local organismsto adaptation (Alberto phenology Phenological Phenological matching key of biological events iscrucial manyfor interactions predicts that predicts herbivores should show

et al. 2012;van Aschandet al. Visser plant vigourhypothesis rasting hypotheses linking herbivorous insect herbivores and trophically linked taxa such taxa aslinked herbivores and trophically data across provenances for 20 cynipid gallprovenances 20 cynipid data for across

and abundance on more vigorous plants or or on more vigorousplants abundance and improved performance and abundance on abundance and performance improved et al. 2008; Savolainen 2008; al. et trophic trophic levels. We thus expect associated 2007; Yukawa, 2000). Many 2007;Yukawa, plants 2000).

et al. 2010; van Asch 2010; al. et

et al. 2014). This is relevant tois relevant 2014). et al. This (Cornelissen

et al.2013b; Ducoussoet

et al. 2007)orthe

et et al. 2008; Price

plant stress plant stress et al. 2012; al. et

Accepted Article per provenance,givingper ofa total 200 study 24of trees distributed acrossfive soil zones (see FileWe S1). selected 2 perplots soilzone Lakm from Petite Charnie). Each sampled provenance isplanted replicate in multiple plots Materials and Methods 2010). generalised linear in a among populations (Hadfield mixed model Nakagawa,framework and by comparingresults the of thatanalyses incorporate the variance-covariance expected (Stone replicates statistically unlikelyindependenthence to represent and are flow, gene ancestry and phenology our into analyses. provenances (for example, variation in chemical defences) by incorporating provenance potentialdistinguishprovenances.phenological other differences effects We from between decline with increasing differencephenological introducedbetween and locally native oak we the test Specifically, impact and of phenological mismatch oncynipid abundance. gall the assess 20 provenances, phenotypic wequantify the 2008).Here Visser, This article This protectedis by Allcopyright. rights reserved. Bercé, (Forêt de site thetrial to provenance theclosest included Table and S1), 1, (Fig. variation environmental separation sourcegeographic greatestand available the incorporating 34 degrees whoseprovenances longitude,latitude and of origins15 degrees span of S1).We selected 20 File supporting are information in the trial provided of layout andthe planting 0.168° detailson west; (Fig. France 48.086°Northwest north, 1; in Charnie Provenance effects were usingexplored alarge experimental oftrial Provenance trialdesign Our analytical Our recognisesanalytical that approach tree populations byare linked shared common

et al. 2011). Here we investigateet al. 2011).Herewe the of potentialimpact non-independence local adaptation hypothesis adaptation local plots containing 4800 trees. Previousplots containing 4800trees. INRA mismatch in budburst date between the samethe datebetween budburst mismatch in , i.e. that herbivore abundance will

Q. petraea Q. at La Petite at La Petite ca. 50 Accepted Article survey date indicates earlier budburst. Although oak phenology is not known to show strong phenology to isnotknown Althoughoak indicates budburst. survey earlier date oakstudies (Crawley on and Akhteruzzaman 1988; Tack other in used closely that parallels (this spring 1995 in bud, 5 isafully open and bud dormant of of phenological matching ( provenances in(discussed moredetail below), providing basisthe examinationfor of impacts score phenotypic highly.dominance meas These apicalstraight branching even and strong with healthy stems, trees while low score, receive a dominance apical poor bushy, and growth irregular with trees Stressed 2001-2002. winter (iii) vigour.plant A higher during 2001-2002. value is winter associ (ii) orcontrarymisleading results. unlikely 20provenances produce as regard 1995data for Wethus to using S4). all the File significantlyin 1995and 2009 were positiv and tree same the for 8 provenances. Phenology scores subset of a in trees 582 2009 for scoringin et across (Alberto G×E years interactions like like (i) of character FileS1 illustrations states): for information supportingsurveys the dataforeachtree provided (see for following traits phenotypic This article This protectedis by Allcopyright. rights reserved. sampling Herbivore 2). 1,Fig. (Table Budburst Diameter at breast height at Diameter Q. petraea Q. Form The speciescynipid The surveyed in this study have all broad geographic distributions; – a measure based on tree ontree shape, record measure based – a – spring budburstphenology was on scored and closely related oaks, they span the Western Palaearctic from the the Iberian from they Palaearctic Western and span oaks, related closely

Budburst ( DBH ) and performance ( ) – tree diameter at a height of 1.3 meters, measured meters, of 1.3 height at a tree diameter ) – al. 2011), wenevertheless 2011), al. repeatedphenology ely correlated (seesupportingely correlated information ures showedsubstantialamong variation ated with more ated withrapid more growth andhigher ed on an ordinal scale 1-10during from ed onanordinal DBH, Form DBH,

et al. 2010).A higher scoreonthe a 0-5 ordinal a 0-5ordinal wherescale, 0 is a ) on herbivore abundance) onherbivore Accepted Article Statistical Analysis Stone Shirley extensive 2002) andour field this with own groupexperience (Csóka and (Redfern key morphological field using speciesin the a to identified andgeneration (a module of woody growth theprevious from year carrying new the leaves). Galls were catkins, acorns; there are no cynipid galls on fema ring onthe scars bark. For twig each wesurveyed thegalls on parts (buds, all leaves, stem, selectedtree, two 10twigs eachconsisting years’ the last growth oneach identified using of plot. within giving survey, reach using weeach trees pruner Duringeach us~4m a pole edge minimise surveyed To effects, 2400 trees. each of the 200study plots,we sampled galls from the24 12 of available trees, a total of were andboth year, generation(asexual) gall each 20selected provenancesthe 2009.Gallwasps 2008 in have and and a spring(sexual) autumn eastwardsPeninsula toIran andthe Caucas This article This protectedis by Allcopyright. rights reserved. (i)Controllingstatistical for non-independence of provenances populations such as populations are available, an alternative istouse a meas data genotypic limited where more here), and (as populations of numbers larger for However, small for numbers of populations using population genomic (e.g.data Hearn non-independence of migrationarisingeach and through generatedcommon ancestry be can of structure a generalised linearmixed model vari one ormore flow) as gene ancestry and/or to is species expectedsimilar incorporate

et al. 2002). et al. One way toaddress potential statistical F ST

(Weir andCockerham 1984) (Weir ities among populations (due to shared commonshared to (due amongpopulations ities us. us. Cynipidgall abundance was sampled across ure genetic pairwise differentiationof between ance matrices inance covariance random the effect (Stone et al. 2011). Covariance (Stone etal. 2011). matrices for trees thetwointernal six rowscomprised of non-independence of populations within sampled, giving four surveys in total. For sampled, giving total. in surveys four le flowers in Europe) of its terminal shoot terminal its of inEurope) flowers le

as anestimateas of their expected

et al. 2014).et al. et al. 2005; al. et Accepted Article were then estimated as either 1- GeoF the betweenhighly the significant relationship a pseudo- the in test (Mantel provenances related to related differentiation of genetic measures using impact on of 1984). Forcomments the alternative provenances where covariance is estimated as 1- are given in supporting information, File S2) File information, supporting given in are pairwise pairwise Guichoux geographic separation between populations. be function distance, as variancecan of estimated strongly with covariancea geographic covariance (Stone This article This protectedis by Allcopyright. rights reserved. estimated as 1- as estimated covariance between all (ii)the samepopulations); (ie. 0 independent entirely statistically provenances to be assumes identity This model matrix. the provenance as using were an identity covariance matrix available, data which genotypic subset of five dataset/covariance matrix following (i)the combinations: 17 provenances for non-independence, provenance theconsequencesofanalyses of the usingexplore all described belowwere To conducted usingthe between provenances withlowest differentiation. matrices, are provided are inmatrices, information thesupporting File S2. approach toaddressing non-independence, population theand values used in covariance ST ; (iv) the full set of 20 provenances using an identity model; and (v) the full set of 20 set full the and(v) usinganidentity model; 20provenances set of (iv) full ; the We used data from 10 putatively neutral microsatellite markers (listed in File S2; inFile (listed markers microsatellite neutral 10 putatively data used from We F

F F ST et et al. 2011)available for 17 of our 20 study 1) (Table to estimateprovenances ST ST as implemented in FSTAT version 2.8.3.2 (Goudet 1995; Weir 1995; andCockerham, (Goudet implemented in FSTAT2.8.3.2 version as for the full set of 20 provenances, referred to hereafter as as hereafter to referred 20provenances, of set full the for , File S2.To see allowustoanalyse 20 study data all for provenances, we used F ST

; (iii) the same the 17provenances,; (iii) covariance matrix using a estimated 1- as et al. 2011). Where genetic differentiation between populations populations correlates between differentiation Where genetic al. 2011). et adegenet F ST or1- GeoF R package, ST . Variance-covariances between provenances F , such that covarianceexpected wasgreatest 17 provenances, using 17 provenances, a covariance matrix ST andgeographic distance the17for alternative approaches to control to control for approachesalternative GeoF p = 0.01 with 999 iterations) to 999 estimate =0.01 with iterations)

ST .

A detailed discussion of this GeoF ST

(full details(full Accepted Article phenotypic traits across both years. The proportion of variance explained variance phenotypicproportion traits The of bythe both years. across between years, whileabundance estimating the was of distribution Ourincorporation applied. responsethe was variable data usuacount and 50,000500,000 iterations andburn-in of with a parameter iterations. samplingevery As 450 prior inverse-gamma was used with shape andscale to0.001. equal Models were run for (with scale1000) were used (Gelman2006) exce specified matrix variance-covariance (1- co-varianceprovenance represented was by provenances, incorporated between the relationship that for eachprovenance.In models level of was provenance effect representedby asi relationshipbetweenthe the provenances, effects). Whenapplyingfor anidentity model a fixed effect with two levels), and provenance, soil zone, plot, and tree (all randomas gall of variance the partition to (GLMMs) models linear mixedthe two andusedgeneralised surveyyears thefor data gall wecombined type, (ii) Statistical modelling of provenance effects on gall abundance This article This protectedis by Allcopyright. rights reserved. a mean (µ) of 0 and a variance (V) of 10 of (V) variance a 0and of (µ) mean a pr effects, Forfixed version 2.15.1. R 2010) in (Hadfield version 2.16 package MCMCglmm the using framework (MCMC) Carlo Monte Chain Markov in a Bayesian were the Modelsfitted effect. of levels between correlation through response that variable in the exceeds expected 1963),inwhichvariation (Matheron whileindependence,the secondincorporated an Schielzeth (2013) (R code for deriving RsqM from MCMCglmm objects in is provided RsqM fromMCMCglmm deriving (2013)(Rcode for Schielzeth effectwas estimated marginal as R the To investigate the influence of tree provenance on the of abundance each individual 8 2 F . For random effect variances, scaled F_{1,1}priors ST of (RsqM),of the model following and Nakagawa or 1- ngle random effect with a unique independent effect withaunique ngle random ior settings assumed a normal distribution with normal distribution a ior assumed settings year two random effects. One incorporated the relationship between gall abundance and tree identity matrix to capture the ‘nugget effect’ effect’ ‘nugget to the capture matrix identity lly manyzeros, contained a Poissonerror counts per shoot into the effects of year (as (as year of the effects into shoot per counts pt for the pt residual the variance which for for an GeoF as afixed effect allowed forin variation ST

) to account for provenance non- year fixed Accepted Article abundance, GLMMsabundance, used tomodel were countsindividualgall of types with oftreeEffect phenotype (Nakagawa 2004). pow more is thisapproach so rate, error wise stringent (2001).Thefalse is and Yekutieli discoverycondition aless the family- rate than and Hochberg andBenjamini (1995) following Benjamini - hypotheses rejected the amongst discoveriesof false expectedproportion the discoveryon - multiple tests false rates of effects not variance explainedbyproportion of the fixe bythe multiplying each and then overdispersion for – account package to MCMCglmm the in models distributed Poisson automatically for fitted is unit that data each for level a distinct with effect random - a term UNIT parameters, including residual variance the by effectcalculated all estimated variance by dividing parameter sumof therandom its the S3). TheFile variancesupporting information component of each random was effect This article This protectedis by Allcopyright. rights reserved. linearity between theselinearity variables was assessedbetween by inflation variance (VIFs)factors their using The fixedThe effect lower gall abundance on provenances that show budbursteitheror before oaks. local after herbivores thatthe of for influence hypothesis phenologypredicts becurvilinear, with will growthtree and health on abundancegall are expected to be linear, the local adaptation Form Form, DBH, Budburst year account potentialfor differences in abundance levels between the two studyyears a factor wasfittedas afixed effect. (tree health), or To investigate whether tree phenotypic traits were good predictors of variation in gall in gall variation of were predictors good traits phenotypic tree whether investigate To For each gall type, gall with per shootwere initiallycounts For each modelled Budburst , or both Budburst was therefore fitted as both untransformed and squared terms. To terms. and squared both untransformed wastherefore asfitted Budburst (spring phenology) asfixed effects. While the influences of and and Budburst erful than alternatives as such Bonferroni d effect (i.e. 1-RsqM). We addressed the Weaddressed 1-RsqM). (i.e. d effect 2 as fixed effects. The severity of co- of severity The effects. fixed as

year DBH and either and (growth), (growth), Accepted Article site. in differences to correlate with oaks native have toburstyet their buds 0 (scores 1). and We thus expect phenological mismatch with some southern provenance trees have open leaves 4(scores and 5), northern provenances time budburst:the byshowing southerly earlier (Fig.with 2c), provenances more latitude adapta local with consistent usingvigour, measured markedly Telavi lower Georgia, showed a in from divergent in more (Fig.provenances 2a). Only the most genetically divergent provenance, vigour using (measured did notdid include zero(pMCMC) wasused to assessthe predictive capacity ofthe fixed effects. coefficientsfixed effect the of the distribution posterior probability that The above. described fitted MCMCglmm the with package in the randomeffect structures prior specificationsand Budburst treebetween phenotypic model traits, containinga fixed effects the main genetically genetically divergent provenances toshow higher stress (measured using increases trialsite and the origins provenance among variation Phenotypic provenances Results the Rpackagethe This article This protectedis by Allcopyright. rights reserved. If provenancesIf are locally adapted, and divergence in selection history between , and their three pairwise interactions was fitted for each gall type. All models were models All type. gall each for was fitted interactions three pairwise and their , HH version 2.3-23 (Heiberger 2013). To assess the importance of interactions importance the To assess 2013). (Heiberger 2.3-23 version DBH DBH ). ). tion. Budburst tion. Budburst phenology shows , does decrease, with increasing divergencegenetic (Fig. 2b), Form Form shows relationship, butincreases nolinear variance in latitude between provenance origins and the trial trial provenance between latitude andthe origins with genetic distance, thenmore we expect distance, genetic with Form score than other score than provenances.Plant

a strong relationship with relationship strong a Form Form, DBH, )

and lower Accepted Article Incorporation of provenance usingIncorporation covariance 1- the models). and datasets alternative the across results of comparison S2for information gavethe higher estimates of variance provenance (see for supportingcomponent effects A unlikelyappropriate. is be to model identity and was >10% for 7 of the 14 gall types (Fig provenance dataset (which assume provenances to be fully independent statistically), the statistically),fullyassume independent dataset be provenances (which to provenance substantially100%) varied the between statistical between provenances (expressed theas mean variance component, ranging between 0 and Oak provenance effects ongallabundance on less thanof 1% shoots in bothsurveyyears and were excludedfrom further analysis. and withoutanybetween consistent yearpattern gall-typesall except generation magnitude, generation just of from 4 (asexual of counts were galls recorded.No acorn Total 2). the foursurvey during seasons,repr generation) Herbivore abundance This article This protectedis by Allcopyright. rights reserved. generation sexualvariance(in of percentage ingall abundance from3% explained byprovenance ranged provenance effects generation 95% (asexual 20 datasetprovenance between attributes 38% (sexual generation We recordedWe The percentage of variance in gall gall abundance in bydifferences shootexplained variance percentage of per The albipes

(Fig. 3). The identity models should thus be viewed as providingviewed as be thus should models identity The 3). (Fig. A. solitarius A. ca. 725,000 20gall types galls generation of (7 sexual and 13 asexual Neuroterus quercusbaccarum ) to 33% (in asexual generation (in asexualgeneration ) to33% , with typesfive exhibiting differences>10-fold (Table 2) ). Gall Gall ). abundance years significantly between differed for ll incorporatingmodels provenance covariance Cynips longiventris s or seasons. Six 20 of the gall types occurred esenting 15different gallwasp species (Table individual gall-types varied over five orders over orders five varied gall-types individual . 3). Because provenances are related, the models used. In identity models ofthe used. models 20 Inidentity models GeoF ) of variance) in to abundance gall

ST

covariance matrix for the same the matrix covariance for Neuroterus quercusbaccarum ) to over 330,000 (asexual 330,000(asexual ) toover Neuroterus albipes and ) Accepted Article models for the 20-provenance dataset, identifying where they differ from the identitymodels identifying the dataset, from 20-provenance they where differ the for models we andhere results), on thetherefore focus results of the1- (seesupporti structures co-variance provenance 3).(Table The signs and strengths relationshipsof stablewere theacross various datasets and 14types of the gall tested three but identifiedall were phenotypic for provenance traits traitsTree phenotypic as predictors of gall inabundancevariation choice of variance-covariance model. byournot influenced effects was follows, analysis fixed what of for Importantlyestimate. re of variance-covariance form the of knowledge as 95%of variation the in abundance of 7 of the 14 gall-types tested. Without more predictor of theimportant abundance ofmanyexplaining gallers, atleast 10%,andas much between provenances (TableS2.2, S2.1). Fig. On basis, this host is tree provenance an differences to abundance attributable gall in variation the of low estimates artificially This article This protectedis by Allcopyright. rights reserved. 5of the for relationships 14gall typesthree (twosexualand generations; Table3, asexual by measured gallssupported more per as shoot, predicted by consider eachtrait in isolation. galls of generation ( term only yielded one significant our analyses interaction confounded, withlowvariance inflation factor variables phenotypic 3). The (Table samedataset the for gall type (asexual generation gall type (asexual Tree health,Tree as measuredby the of least and at one shoot per abundance relationships between gall Significant DBH , was a betteroverall gall predictor of abundance per shoot, with significant N. numismalisN. C. quercusfoliiC. ;

mean coefficient = 0.031, pMCMC = 0.014), hereafter we hereafter pMCMCcoefficient =0.014), mean =0.031, Form ,

significantly predicted the abundance of only one ; Table 3, Table Fig. 4a)for; trees whichhealthier s (1.05, 1.02, and respectively). 1.02 Because the plantvigourthe hypothesis. Tree vigour, as lationship it is not possible to give a correct correct give a to is notpossible it lationship ng information S2 for full comparison of comparison full S2for nginformation Form, DBH,

GeoFst GeoFst Budburst and co-variance matrix Budburst : Form for sexual for were not not were Accepted Article hypothesis). and while vigour plant by (as onlarger hypothesis), the predicted trees abundant varied thethree Relationships for asexual types:autumn gall generation on abundant more larger trees (Fig. 4b, aspredicted by theplant vigour hypothesis). spring ( For galls generation the sexual 4b). Fig. This article This protectedis by Allcopyright. rights reserved. involves introduction of non-native genotypes that take decades to establish and live for for and live to establish decades take that genotypes ofnon-native involves introduction climat to particular these aspects in apply of strategies to to develop, take time remediation consequenceslikely ofandremediation; and non-intervention impacts wherethe both of challe pests. The potential providersor service abundancedistribution and ofassociated taxa diversity,the on foundation influence species are likely impacts Climatechange to strategies. of impactscommunity global both environmental change andany associated remediation Discussion showing the latest budburst(low budburst score, 4c).Fig. ontrees galls generation the abundant weremost asexual summer/autumn while score), were generally onprovenances mostabundant stage, expectedthelocal spring as adaptation hypothesis. Instead, sexualgalls generationfor term.However,squared nogall type showedpe types showedsignificantly relationships, non-linear incorporating asignificant with abundance/shoot, significant correlations for ei N. quercusbaccarum Phenology, asmeasured byPhenology, Predicting future change in ecosystem functioning requires an the of understanding requires an functioning change inecosystem future Predicting showed the opposite pattern (as predicted by plant stressthe Budburst e matching of long-lived foundation trees, whichtrees, foundation long-lived of e matching understand their trajectories through time. Allunderstand trajectoriestheir through time. showing budburst (high theearliest budburst ak abundance at an intermediate phenological intermediate ak at abundance an A. testaceipes these– whether are beneficial ecosystem , wasthe overall predictor best ofgall nge for a given system is then to estimate estimate to then is system given a for nge ght gall types 3, (Table Fig.Four gall 4c).

and N. numismalis N. albipes N. numismalis N. ), galls were were galls ), were more more were Budburst

Accepted Article differentiation ( differentiation genetic of measure a on based specified is covariance which in models to relative densities consis be true, to we knownot which entities, as statistically provenances treat independentthat modelsthat indicate Ourresults origin data. provenance and genetic on population based estimate covariance models that to independence variance-covariance among provenances, identity from models oak assuming total these of relationships isrobustacross form the shoot) oak of density(abundance per population the in variation significant with associated is turn in this that herbivores, and associated influence Our resultsshow that oak provenances vary covariance ofoakprovenanceInfluences onthegallwas Petanidou S2.3), their absolute values are small, providing little resolution of the relationship of genetic of of relationship the resolution little providingare small, values absolute their S2.3), change at guild and community levels (e.g. Forkner levels(e.g. guild andcommunity at change same guild,the addressing arecognised ne the consequences of host plant variation in performance and phenology specieson multiple in toprovenance our study trialquantify of a use keyA of isits in performance. aspect decline so climateas changes, provenances becomenative while increasingly andmismatched becomeincreasinglythough initiallytree provenances that, climate-matched, introducing not Our analysescenturies. shed light onthelikely native herbivoresfor consequences of This article This protectedis by Allcopyright. rights reserved. pairwise pairwise by assuming would beinferred provenances than ongall of tree influence the provenance then F

etal. 2014, van Asch ST and related measures are consistentlysign are measures related and F ST or GeoF ST ). If these models accurately describe provenance co-variance,

et al. 2012, Vatka abundance is substantially greater (38% - 95%) (38%-95%) issubstantiallyabundance greater tently underestimate provenance effects ongall effects provenance tentlyunderestimate ed to quantify the impacts environmental to of the ed quantify models that assume patternsof verymodels that different p guild, andincorporationp guild, of provenance significantly in phenotypic traits likely inphenotypictraits significantly to

et al. 2014). tobestatistically While independent. multiple gallwasp species.multiple showWe that ificantly larger than 0 (Table S2.1 and S2.1 (Table 0 than larger ificantly

et al. 2008, Ohlberger al. 2008, et

et al. 2014, al. et Accepted Article (Bangert and Whitham 2007;Wimp and Whitham (Bangert toassociatedgenotypic levels variation trophic genetics worklinking plant host community power given the non-independence of populations (Felsenstein 2002, Stone explanatory its provenance and of value theimpact for a “true” to estimate difficult therefore is It discussion). comprehensive a S2for (see distance geographic and differentiation This article This protectedis by Allcopyright. rights reserved. (Vakkari each study. AlthoughFinnishoakpopulations showintraspecific substantial genetic diversity adaptive contrasting of levels to reflect likely strongprovenance andthe between this presentthePetite Charnie hostplant trial at effects is of generations sexual th of tree any of genotype noeffect for found reciprocalgallwasps. In transplant expe genotypic effects in structuring a community robur factors (Hersch-Green herb for effects hostgenetic of importance the show strong genotype plant effects with thestudies of genetically diverse hybrid (e.g.populations Dungey a diversity shared population, feature a from wouldbeexpected natural genetic single than Charnie Petite (Petit phenotypic diversity) the than much wider geographic array provenancesof Lapresent at the geneticadaptive diversitythe population a tree within presentsite, at woulda andwe expect host effects to genetic of relative importance othe )

The strong show we effects provenance The are growingwith the consistent body of in Finland, Tack in Finland,

et al. 2006), theyet al.2006), diversity less genetic showsubstantially also weexpect (and

et al. 2002). Our provenance trial study involves a muchgreater of array a studyinvolves trial Ourprovenance 2002). al. et Neuroterus quercusbaccarum

et al. 2011). Working near the northern limit of pedunculate oak( pedunculate of 2011).limit et northern Working al. nearthe et al . (2010)foundspatial tobe effects more important than tree on associated herbivores (Tack

et al. 2005). A key challenge in riments at local and regional scales, Tack genetic and phenotypic and di genetic ivore communities relative to ecological relative other ivore communities e three and types gall their three studyin (asexual e of herbivorous insects that included insectsof herbivorous that several r factors is likely to correlate positively is likely r factors with to correlate are shared with ourstudy). Thecontrast shared are

et al. 2010). Generally, theet Generally, 2010). al. this fieldthis tounderstandis

et al. 2000) that often 2000)that al. et versity of the oaks in et al . 2010). et alet . (2010) . Q. Accepted Article oak-gallwasp interactions (Egan and Ott 2007), and Ott (Egan interactionsoak-gallwasp plasticity phenotypic local and adaptation. the been to which theirextent natural ofherbivores enemies result responses the and are of Van Asch et al. 2012, Vatka et 2014) al. insectivorous birds–particularlyof phenologybudburst on abundance of associated le has levels. This trophic beenshown, example, impact inoak for changes inworkonthe of ofspeciesmajor responsescomponent toclimatecascade that change can through associated spring budburstphenology. in Phenologicalvariation changes have long beenrecognised as a Impacts of between-provenance variationin phenology are implemented. migrationof assisted forms where matching orother climate insituations increased to be this This article This protectedis by Allcopyright. rights reserved. this the that sp contrast between hypothesise be presentcan trial.Weto the wider phenological extended inaproveance variation 1984) and (1988) Akhteruzzaman Crawley(1962, and U.K. innative phenological variation local response to relationships tot Similar trees. species ( on abundant were more early budbursting trees, gallwasp lifecycle inthe spring generations can beverygalls Spring andautumn different. ofthe responses the We found hypothesis. also adaptation local provenances, rejectingour withabundance increasing mismatch nativebetween phenological andplanted oaks The strongest predictor of between-provenance variation in gallwasp in variation is between-provenance density predictor strongestof The It is striking althoughthat adaptationto individualtrees has been demonstrated in N. numismalisN. and N. quercusbaccarum hose we observed were recordedfor Parus major . A key inresearchquestion on these systems has ring andautumngenerations may exist because ) were more abundant ) weremore budburstingon late none of our gall types showed decliningshowedour gall types none of (Visser 2008,VanAsch andVisser2007, while same the autumn generations of af-feeding moths and breeding responses and breeding moths af-feeding ; our results show ourresults thethat ; same trends

Q. robur Neuroterus populations by Askewpopulations gallwasps in gallwasps Accepted Article generation gall types. Support gall types. generation wasthusevenly springpositive gall generation for types,sexual Sign taxa. and generations gall among varied vigourdensitytree between gall and withof correlations phenology, As thesign impact. little contrast,density. tree commonly In shape - used as indicator physiologicalan of stress – had Impacts of between-provenance variationin tree vigour. and/or plant hosts (including many aphids; Harrington generations between involve lifecycles, alternation particularlythese complex where more a may impacts bottom-up andcomplex understood, poorly such of The are complexlifecycles dynamics Parmesan, 2010). (Singer and generation either in limiting perhaps adaptation phenological generations, lifecycle local two on the induction.for gall Annual gallwasp phenology may phenologically of young suitable tissues higher abundance have may budbursting trees later This article This protectedis by Allcopyright. rights reserved. poorly understood. Mechanis galls. generation autumn for defended) galls andworst generation thenbeboth best (=best for spring Highlyvigorouscould trees 1970). defence inthesummer (Feeny, metabolic springthe to growth in from investment generation gallsautumn couldbedriven by 1974; White, 2009). Onepossibility is that apparent contrast the in patterns springfor and (Cornelissen Tree vigour between-provenance in explained variation significant Tree also gall oak

et al. 2008; Price plant stress1991) andthe hypothesiset al. (White 1969;White the long-known seasonal shift in oak resourceoak in shift long-known seasonal the while two of three were asexualwere negative for three twoof while split between the plant vigour hypothesis hypothesis vigour plant the between split with vigour correlations tree were ificant ms of oak defence oak defence against remain gallwasps ms of pply to othergroups multivoltine orwith thus be the result of contrasting selection

et al. 2007).et al.

Accepted Article 2005; Ducousso climatematched trees, andsignificantly dissimilar gallwasp assemblages. emission highscenarioto couldleaddifferences phenological major between native and under >10°of on oak impact a required phenology,the latitude little probably have shifts of shif latitudinal the small while suggest that respectively south, 0-2° (Broadmeadow further and5-12° Broadmeadow native oak populations native beshared oak species. Thewill by plant towhichany extent other such (Aitken more southerlyintroducing provenances 2009;Jenkins Coppola In populations. climateEurope, predic models Predicting the impact of plantingclimate-matched onassociated herbivores This article This protectedis by Allcopyright. rights reserved. ‘window opportunity’ (Durant of narrow phenological is strong herbivore which in fitness interactions diversity. hypothesised We thatgenetic impacts herbivore interactions, reflecting herbivoreavailable phenotypic plasticity andadaptive differences influence associated herbivores is likelytovary according to specific plant- on) young tissues, they show no strong preference for host trees at a single trees host phenologicala at theyfor youngno strongpreference show on) tissues, no evidenceof such awindow gallwasps:while for oviposit they to survive prefer better (or on newly leavesflushed (Forkner includes many feedand Visser,2007).This 2010;Asch that van Schopf insects Netherer and The widespread existence of latitude-dep widespread existence of The Climate matching necessarily requires introduction of phenotypically divergent tree phenotypically of introduction requires matchingnecessarilyClimate

et al. 2008) suggestsphenologicalthat the et al. et al et

et al. 1996). Under low and high emissions scenarios for 2050, scenarios for emissions high low and Under 1996). al. et . (2005) identified climate matched sites for five UK locations thatwerelocations UK five for sites matched climate (2005) identified .

et al. 2009),and et al. matchingthus alwaysclimate involvewill

et al. 2008; vanet al. Asch andVisser 2007). However, we found that show budburst(Broadmeadow that earlier ts required under a lowemission would scenario undera ts required ly correlated with with ly correlated ofsuccessful exploitation a t warmer summers and winters (Giorgi and (Giorgi and winters summers t warmer of phenologicalbe mismatch for greatest will endent photoperiod cues plant phenology in contrasts between contrasts climate-matched and

et et 2007;Forkner al.

et al. 2005). (Fig. al.Our 2005). et results 2c)

et al. 2008; al. et

et al. Accepted Article provenances, whichlocal increases resource herbivores heterogeneity for (Hubert and local and matched both of planting mixed advocate UKguidelines forestry in Current 2005). strategy and itssuccess at maintaining fo become established before native provenances sapling andyoung before tree the conditions jus to aidentify to survive as is isable managers that climate-matched provenance a forest whether climate isbut also provenance matching all. feasible that The challengeat with for will dictate provenance specific a vigourfor variablesslope betweentherelationship range. environmental and of The adapted locally in the performanceexpected ofnativeprovenances decline asclimates move awayfrom any climatesofcurrent provenancesintroduced ma to relevant two stages in a climate matching strategy: expected the poorperformance under complex. be may stage. As discussed above, impacts onherbivores with multivoltine ormulti-host lifecycles This article This protectedis by Allcopyright. rights reserved. and Wessman(Buma 2013; Lefevre resilience-targetedthan rather strategies may Theoretical nevertheless Dumroese such 2013). studies adaptive-,suggestthat and (Williams ranges their current with outside risks moving associated plants and algorithms, in matching strategy lackoperating include of procedures, uncertaintiesover climate which models touse forestry matching an adaptive as ofclimate slowingthe implementation cited as down settingsconservation (Broadmeadow Assisted migration is increasingly seen (Lefevretrees on host selection accelerate both environmental that conditions creates and 2007), Cottrell The we relationships The found between hostplant vigour andherbivore abundanceare

et al. 2014), and host-plant mediated selection on associated organisms.

et al. 2014). would the communitythat add impactsWe

et al. 2005; Dawson rest productivity is untested (Broadmeadow not only theimpact on associated levels,trophic fail. Climate matching is a recently developed recently is a Climate matching fail. be the best way to safeguard forest systemsforest to safeguard way best be the tched to predicted andthetchedfuture conditions, topredicted as a viable strategy viable as a for commercial and tifying have been reached, selection its and

et al. 2011; Stone 2010). Issues

et al. al. et Accepted Article 2014; Thomson drafts of the drafts manuscript, andKatJu Bradley, feedback on Tansey Christine Tom for Oliver Ally advice, and Phillimore, statistical for strategies on interactplant-herbivore-enemy and change management forest biodiversity: of predicting of theimpacts climate matching research ‘Climate CLIMAFO project NERC-funded under the workresearch comprises This Acknowledgements biodiversity.forest-associated decisions managers take concerningcan informed climate impacts matching likely of on (Bidar enemies natural herbivores by their on inflicted including ecological processes, mortality phenotypes the and other plant both of andherbivore numerical understanding responses adaptive then requires impacts inturn the spanning trials enviarray of different in an effectsGiven on strong traits plant includingenvironmental phenology,of the work repetition so to effects provenance wefound we expectthe a significant contain genetic component. haveanalysed beeninbe shown highly sampled the to provenances2012), heritable (Sinclair we phenotypic traits the oak single trial, ona based was study While our al.2010). et genetic to provenance to separate environmental contributions (Tack and required phenotypes al. 2007; Sork have only been mappedin onlyhave tiny species of been a minority (Robakowski (Tack ecologicaland other factorsto genotype plant such as forest This trees. requires better unde of assisted needmigration to be considered, carefully particularly species sofor foundation This article This protectedis by Allcopyright. rights reserved.

et al.2010). Geographical patterns localof adaptation and treephenotypic variation

etal.2013), andstudies that targetmu et al. 2010;van Asch al. et

et al. 2012). Much more needs to be done before forest before be done to moreneeds Much 2012). al. et t-Bouzat and Imeh-Nathaniel 2008; Imeh-Nathaniel Rasmann and t-Bouzat ions. (NE/H000038/1). Wethank JarrodHadfield ronments Understandingis clearly desirable. ronments lja Ernst, Matt Ernst, lja and Curtis, Wainhouse, Stefan rstanding of the relative contributions of host relative contributions the rstanding of the tree phenotypes exposed to herbivores to treephenotypes the exposed ltiple provenance trial locations provenance are trial ltiple

et al.2012; Savolainen

et al.

et

Accepted Article Aitken SN, Yeaman S, Holliday JA, Wang T, Curtis-Mclane S (2008) Adaptation, migration Adaptation, Curtis-Mclane S(2008) T, Wang JA, Holliday YeamanS, SN, Aitken cited Literature http://doi.org/10/647Data Centre: available Data this study for for from are download theNERCEnvironmental Information Data archivingstatement the developmentof paper. this in role their Review for Axios thank field. theWe help in invaluable for Rushton Katherine This article This protectedis by Allcopyright. rights reserved. Alberto FJ, Aitken FJ, R SN, Alberto Alia Alberto FJ, L,Alberto Bouffier Louvet J-M, Lamy Bangert RK, Whitham TG (2007) rulesand assembly Genetic community phenotypes. TheAskew RR (1984)biologygallwasps. In of of Galls Askew RR(1962)TheDistribution of Frigerio JM Boury C, FJ,DeroryJ, Alberto Applications, climatechange populations. for outcomes tree or extirpation: change evidence from tree populations. Journal of Animal Ecology, Evolutionary Ecology, Evolutionary Anathakrishnan, pp. 223 -71. Oxford and IBH Publishing Co. New Dehli Quercus petraea. Grad SelectionAlongEnvironmental Natural gradient.altitudinal seedresponses for and leaf phenology inna

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19 , 1645-1661. , ed. TN , ed. Evolutionary The The Accepted Article Benjamini Y, Hochberg Y (1995).Controlling Y,(1995).Controlling Hochberg Y Benjamini This article This protectedis by Allcopyright. rights reserved. Benjamini Y, of Yekutieli control D the (2001)The discoveryfalse in multiple rate testing Bower AD, Aitken SN (2008) Ecological geneti Ammer A, Bolte M Löf C, A(2008) Imeh-Nathaniel MG, Bidart-Bouzat Buma WessmanB, climatechange,Buma Forest CA resilience, (2013) andfor opportunities for and future change CJA the (2005)Climate Ray Samuel D, MSJ, Broadmeadow Cavers S, Cottrell JE (2014) The basis S, JE(2014) ofThe resilienceCaversCottrell treespecies in itsuse forest in and Janss IN, KM, Robinson Abreu C, Bernhardsson B testing. to multiple approach powerful under dependency. albicaulis (Pinaceae). 306 adaptation: Aspecific case of problem. general a broadleaved tree species inBritain. Forest Research, Climate change impacts, strategies and integrative concept. 1354. Defenses against Insect Herbivores. adaptive forest management in Britain. Britain. in management forest adaptive genes. defence plant in structure genetic with communityco-occurs herbivore in metabolomeand Geographic structure

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Accepted Article Rasmann S, Pellissier L, Defossez E,Jactel (1991)Price PW ThePlant Vigor Petit RJ, Csaikl UM, Bordacs S T, KallimanisPetanidou AS,Sgardelis SP, Pearse IS, Karban R (2013) Leaf drop affects herbivory in oaks. globally coherent A (2003) G Yohe C, Parmesan Ohlberger J,ThackerayOhlberger IJ, Winfield Mabe SJ, Nakagawa S (2004) A farewell Bonferroni to farewell S(2004)A Nakagawa This article This protectedis by Allcopyright. rights reserved. Netherer S, Schopf A (2010) Potential Schopf effects A(2010)Potential S, Netherer Nakagawa S, Schielzeth H (2013) A general and simple method obtaining and R Schielzeth S, general for simple method Nakagawa H(2013)A natural systems. systems. natural plant-insect interactions along elevation gradients.elevation interactions alongplant-insect populations. from diversityoaks -Phylogeography of over patterns basedondata and 2600 111. mismatch.phenological suggest useinacommunity future flowering pollinator phenologyVariable and Proceedings of the Royal Society B-Biological Sciences, age-sizematters: truncationpopulation alters response to mismatch.trophic example. asspecificthe processionary forests-General moth aspects and pine European mixed-effects models. linear generalized publication bias. 142. Forest EcologyandManagement, Forest Ecology and Management, Nature, Behavioural Ecology

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4 2 , 104- from , 133- Accepted Article Savolainen O, Lascoux M, Merila J (2013) Ecological genomics of local adaptation. of genomics Ecological local J (2013) Lascoux Merila adaptation. O, Savolainen M, Robakowski P, Li Y, Reich PB(2012)Local ec M, P(2002) Shirley Redfern This article This protectedis by Allcopyright. rights reserved. Singer Parmesan MC, C (2010)Phenological SchonroggeK, HarperLJ, Lichtenstein CP(2000) trees. adaptation local in and (2007) Geneflow Knürr T, Pyhäjärvi T O, Savolainen Spittlehouse DL,Spittlehouse Stewart RB (2003) Adaptation to climate change in forest management. adaptation local closelyfor in adjacent Evidence KA, C(1993) Hochwender Stowe VL, Sork Sork VL,Aitken SN, Dyer AJ RJ, Eckert Ecology, intrees. photosyntheticbroadleaved optima deciduous temperature influence Preston Montford,Field Studies Council. Reviews Genetics, Review ofEcologyReview Evolution and Systematics, Journal of Ecosystems and Management, leaf herbivores. herbivores. leaf subpopulations of Northern Red Oak ( 911. responsesand population tochanging climate. the of approaches into local genomics trees: landscape for adaptation understanding TransactionsSciences,Society oftheB-Biological Royal hosts:their signal of climate change or pre-existing adaptive strategy? Cell andEnvironment, galls (Hymenoptera : Cynipidae): betweenSimilarities gallscynipid and seeds.

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Accepted Article number Bercé, are shown inbold. traits genotypic and the data, mean values across alltrees of each provenance for the phenotypic decimal and longitude in latitude country), andname (site origin codes,provenance INRAprovenance digit showing 3 east their west to Table 1. This article This protectedis by Allcopyright. rights reserved. in Fig.1 in Fig.1 Site Site 17 17 16 15 14 13 12 11 10 20 20 19 18 9 8 7 6 5 4 3 2 1 Budburst, DBH, Budburst, Code 184 249 179 248 255 181 257 252 225 250 230 233 245 201 211 194 210 237 217 185 Summary ofthe 20 studied provenances of

La Haie Renaut Johanneskreuz Saint Germain Réno Valdieu Valdieu Réno Romersberg Horbylunde marienberg Spakensehl Provenance Provenance Wolfgang Wolfgang Blakeney Vachères Soudrain Prémery Cochem Kloster- Sycow Étangs Telavi Telavi Bercé Bercé name name Bolu Bolu Still and Form Germany Denmark Germany Germany Germany . Values for the provenance closest to the trial site, Forêt de Forêt site, trial the to closest the provenance Values for . Country Country Georgia Georgia France France Austria Turkey Poland France France France France France France France France France UK 45.47 45.47 31.67 17.93 16.57 Long Long (DD) 5.63 4.96 4.95 2.38 2.08 0.67 0.39 10.6 10.6 9.41 9.05 7.83 7.25 7.05 6.73 -2.5 3.6 3.6 degrees, altitude (Alt), the availability of of availability the altitude (Alt), degrees, 43.98 46.93 48.67 46.95 47.81 51.78 41.88 41.88 40.92 51.18 47.41 56.13 50.15 48.58 50.08 48.82 (DD) 47.2 47.2 48.9 48.5 52.8 52.8 49.4 Lat Lat Q. petraea, 1200 650 200 180 300 178 230 155 700 700 210 310 115 160 460 688 400 220 (m) Alt 60 60 76 80 80

Genotypic data data Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes No No No ordered ordered by longitude from Budburst Mean 3.69 3.69 1.77 1.55 1.24 1.71 1.65 1.59 1.08 3.79 3.79 1.58 1.42 3.23 0.54 0.88 1.61 1.03 1.55 1.86 1.06 2.4 2.4

Mean DBH 113 113 109 106 110 113 116 120 104 104 103 102 116 105 112 101 94 94 98 77 77 94 89 96

Mean Form 4.15 4.19 4.24 4.34 4.33 4.13 4.14 4.24 3.06 3.06 4.39 4.76 4.74 3.99 3.92 4.42 4.56 4.43 4.17 4.2 4.2 3.9 3.9

Accepted Article bearing galls, and theof ratio meangallsper shoot between 2008 and2009. shoots proportion of -the incidence shoot, and the per galls type,number of meaneach gall recorded types La Charnie. Petite show,at Columns for eachsurveytotal number year, of the Table 2. This article This protectedis by Allcopyright. rights reserved. quercusbaccarum Neuroterus Neuroterus anthracinus albipes Neuroterus quercusfolii Cynips Cynips longiventris divisa Cynips Andricus solitarius Andricus inflator Andricus glandulae Andricus fecundatrix Andricus callidoma Autumnasexual generation surveys quercusbaccarum Neuroterus numismalis Neuroterus Neuroterus anthracinus albipes Neuroterus Biorhiza pallid Andricus testaceipes Andricus inflator surveys generation sexual Spring Gall-type (speciesand generation) generation) Prevalence, incidence, and between-year variation in the abundance of the abundance of 20gall inthe between-yearPrevalence, incidence,variation and

a

123708 6.44 0.56 0.56 6.44 123708 0.037 0.93 0.64 12194 0.51 16.33 313507 2.38 45771 0.0085 0.01 197 0.000052 1 0.057 0.099 1901 0.000052 0.026 0.028 528 0.0001 2 44 0.0023 0.0001 0.002 83 0.0043 0.0037 334 0.00021 4 0.00021 5808 1154 14278 822 13 973 52 galls Total

0.017 0.24 0.048 0.59 0.034 0.00054 0.041 0.0022 shoot galls/ Mean

2008 2008 0.15 0.15 0.039 0.35 0.03 0.00046 0.0025 0.0018 Incidence 0.013 0.013 7834 7834 17486 7411 5828 10 2291 0 galls Total 94830 94830 12124 23820 27531 456 3 184 726 43 3 335 1103 45

0.33 0.33 0.73 0.31 0.24 0.00042 0.095 0 shoot galls/ Mean 3.95 3.95 0.51 0.99 1.15 0.019 0.00013 0.0077 0.03 0.0018 0.00013 0.014 0.046 0.0019 2009 2009 0.2 0.2 0.29 0.23 0.16 0.00042 0.067 0 Incidence 0.49 0.49 0.028 0.29 0.32 0.013 0.00013 0.0057 0.028 0.0012 0.00013 0.012 0.029 0.0018 1 : 1.4 1.4 1 : 15 1 : 1.91 : 8 1 : 1.31 : 2.3 1 : - 2008:2009 galls/shoot mean Ratio of 1.61 : 1.21 : 16 :1 2.11 : 2 1 : 2.5 1 : 13 :1 1 1 : 1:18 19 :1 2.3 1 : 2.5 1 : 9 1 :

Accepted Article N. numismalis N. anthracinus N. albipes A. testaceipes the differed from of theidentity models result the using Where pMCMC. assessed signfixed ofthose effect coefficients that di Table 3. This article This protectedis by Allcopyright. rights reserved. 0.05. 0.05, while allother results indicated ** as or and Benjamini &Yekutieli (2001)resulted adjustment pvaluesin of indicated as *top> ***=p<0.001, of positivefalse discovery for multiple rates **=p<0.01, follows: *=p<0.05, models, the identity result follow the the follow result identity models, Cynips divisaCynips A. solitarius glandulae A. A. fecundatrix N. quercusbaccarum C. quercusfolii N. anthracinus N. albipes N. quercusbaccarum N. numismalis Gallwasp species (generation) (Sex) (Asex) Summary ofGLMMs phenotypic for tree predictors,showing significance the and (Asex) (Asex) (Asex) (Sex) (Asex) (Sex) (Asex) (Sex) (Asex) (Asex)

(Sex) (Asex)

Year *** NS NS +* NS NS NS NS NS NS NS NS NS NS NS NS +* NS +** NS NS NS *** NS *** NS *** * S+** NS *** *** NS NS -* NS NS NS NS NS NS NS NS -* NS NS NS NS NS -*** NS +**/+* NS NS *** NS NS +*** NS *** NS NS *** *** NS NS -* NS NS NS NS NS NS NS NS -* NS NS *** *** NS +* NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS +* NS NS *** +* *** *** NS -*** -*** +* NS NS NS NS NS NS NS NS NS NS NS NS +* +* NS -*** -*** NS -*** -* NS NS *** NS *** NS *** NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS

Form 1- GeoF DBH / +*** +*** -* NS * NS *NS NS -* +*** +*** / *N SN NS NS NS NS NS +* testing following Benjamini & Hochberg (1995) ffered significantly Significance from zero. was ST *** in the table below remain significant at p < p at significant below remain table the *** in

result. Significance codes in the table are as are table inthe codes Significance result. NS=p>0.05, non-significant. Consideration burst Bud

burst Bud 2

burst: DBH Bud

burst: Form Bud

Form: DBH 1- GeoF

ST

Accepted Article (open circles). Vertical barsrepresent 95% confidenceintervals foreach mean. budburst in the spring the budburst in (c) and growth; vigorous more indicate (b)values indicate healthier trees; 2. Fig. byare indicated twotheir letter ISOcodes. provenances. Sitesare identified by inTable1. Countriescontributing number provenances 1. Fig. LegendsFigure This article This protectedis by Allcopyright. rights reserved. independence (i.e.or incorporate a1- identityindependence models), non- for account population circles) (black either donot that dataset 20-provenance the 3. Fig. intervals for each mean. confidence % barsrepresent 95 Vertical on gall abundance. effects provenance of modelling iterations, of number and sampling frequency describedthat previouslyfollow the for with a Poissonfitted distribution, and (b)w as fitted tree and plot, zone, soil provenance, werepackage modelledresponse variables and provenance with fitted as effecta fixed and Forêt deof shown.identitythe Bercé in Modelling MCMCglmm was conducted the R their position re In (c) de (Forêt site Bercé). trial genetic differentiationgenetic (estimated as Between-provenance variation in Between-provenance threevariation phenotypic (a) variables The proportionof in variation gall abundance attr Location of the Petite Charnie provenance trials and the 20 the selected study provenance Charnie trials and theof Petite Location . In (a) and (b) the position of provenances on the (b) theposition and on the provenancesIn of (a) Diameter at breast height(DBH) at Diameter GeoF Budburst Budburst ST ) from theprovenance closest tothe provenance random effects. Models of (a) Models randomand (c) were of (a) effects. ith a Gaussian distribution. Priors, distribution.burn-in, Priors, Gaussian ith a lates to the latitude of their site of origin withorigin of their site tothelatitude of lates date, in whichdate, in higher values indicate earlier ibuted to the effect of provenance provenance for of effect the to ibuted

GeoF ST

variance , in which higher valueswhich , in Form x-axis

covariance matrix , inwhich higher relates to their relates Accepted Article (c) (c)

and 95% confidence intervals from these dist these intervals from 95% confidence and predictor ( predictor of Distributions earlierbudburst. indicate 4. Fig. This article This protectedis by Allcopyright. rights reserved. x(mean coefficient couldnumismalis notbe producedinthis wayduetohigh random variances. effect (Forêtprovenance de Bercé). for Plots models theof asexual gallsgeneration of N. tree local most thefor trait the ofphenotypic value indicate themean lines vertical Budburst. Plots of significant relationships between gall abundance and (a) and(a) gall abundance between significant relationships of Plots x ) were the from derived 1000stored samples MCMC as: Higher i ) + 0.5x + 0.5x Form ∑ scores indicate healthier whiletrees, higher random effect variances effect random y ributions for each of the two studyyears. the of each ributions for at intervalsacross of range at the aphenotypic i ). Plotsshow themeans (boldlines)

y Form i = exp(intercept = Budburst ; (b) DBH scores scores

Dotted Dotted ; and i +

Accepted Article This article This protectedis by Allcopyright. rights reserved.

Accepted Article

This article This protectedis by Allcopyright. rights reserved.

Accepted Article This article This protectedis by Allcopyright. rights reserved.