Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. aei Troispoux Valerie Tysklind Symphonia Niklas the in traits history life syngameon. early role on the processes selective distributions: of species and adaptation local Microgeographic 95 ei ap 06,truhhrioy ies,lc frsucslk ae So ore 2007), Poorter & (Slot water like (Valen resources high of is mortality lack when disease, stages, herbivory, life early through in 2006), disappear Hampe (Moles will them & lifespan of Petit Most their 1975; over adults. become seeds never of will seeds millions produce may Trees Introduction taxa. related closely very the among in flow. ecotypes and different gene level the occasional of microgeographic despite maintenance sympatry the can the intimate part, stages at in in early even living explain, different very generalist, syngameon distributions, to the the a Symphonia contributes in at species also as seedlings filtering tree performance behaved and environmental differential of sp1 seeds that such Furthermore, of S. determinant indicating Conversely, performance ecotypes, key differential both strategy. a The of planted high-gain be Synthesis: distribution when high-risk known rates 4. a growth the environments. of matches fastest of the habitats specialist variety showed a a region. but provenance in with it’s habitat from or well adaptation, suggesting home habitat individuals performing its local own habitat, of outside their of home planted performance in signs its when better and showed in surviving penalised survival herbivory and strongly the and faster was evaluate growing globulifera growth, provenances to S. survival, and years Germination, 16 ecotypes 6 of seeds, bottomlands, 3. combinations of (510 in some experiment course provenances. exclusively transplantation the and almost reciprocal over ecotypes found A followed different globulifera sp1. and Symphonia Symphonia set-up Symphonia taxon/ecotype, ecotypes, shield, was generalist or Guiana conditions) more the taxa in undescribed In and sympatric variations yet syngameon. stage two on and a Symphonia distribution adult rely least and Neotropical germinate, species stages to at the tree never young with grow by determining very will questions in represented will at these all seeds are fraction survival studied at those have and minute matter We germination processes of a 2. these in most only variations do abundance? eventually and Although Do and heterogeneity? process? microgeographic produce. young, random to die they a adaptations many just seeds this germinate, of Is which number reproduce. those large Of the by will. characterised plenty are Trees 1. Abstract 2020 28, April 7 6 5 4 3 2 1 oayadMdligo ln n eeainArchitecture Vegetation and Plant AgroParisTech of Modelling and Botany ECOFOG Guyane la Universit´e de Mediterraneennes Forets des Ecologie Guyane IRMAR de forets des Ecologie 1 ai-ireEtienne Marie-Pierre , 1 an mrCazal Omer Saint , 5 2 oieBrousseau Louise , aoieScotti-Saintagne Caroline , 1 tal. et 6 rn Ferry Bruno , 04,yttevs aoiyo those of majority vast the yet 2004), 3 lxnr Tinaut Alexandra , 7 n vnScotti Ivan and , 4 , 3 Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. ieecsi lmt,si rpris optto,adpeaos n a eal otaeaata least at apart tease to ( able factor be one may than one more predators) cross bundle and that a RTEs competition, to designing properties, adaptation By differential soil in habitats. observe climate, constraint only between in can main differ genetic differences one The that that of processes. factors is evidence adaptation ecological RTEs no local from of obtained and with results plasticity plasticity the through phenotypic interpreting traits purely phenotypic to and due Eichhornconditions be 2005; may (Fang Stewart ecotypes influence & tree (Boshier between studies differences other in observed Palomar not were wide conditions, a for are Nagamitsu identified al. lifecycle, 2012; et been long Davies has the & parameters to Baltzer (Fine environmental due 2012; species seedlings particular tree in to tested of adaptation mostly range local adaptation, of local evidence on (Wadgymar RTEs diverse: two system from given conclusions The a trees, 2013). within pena- In traits (Anderson not different adaptation are for but coexist local may ’home’, the and at al. exclusive underpin better mutually may not perform orchestrating neutrality are populations are conditional processes some trade-offs however, then genetic always if, elsewhere, individuals then adaptation; ‘local’ lised conditions, local if and system: of populations the patterns across all the patterns for of individuals generality the ‘foreign’ on outperform depending better nuanced perform further ’home be populations (the can on elsewhere local based than when adaptation, provenance ’local of found own (the terms is ones in adaptation transplanted directly local than interpreted be expectations: can theoretical environments straightforward different in populations between eas hyalwosrigpromnevrac ietyars naryo niomna atr (Mor- factors environmental of distribution, phenotype array or an species across and larger directly an variation variance and are habitat RTEs between adaptive performance costs controlled. ris link observe observing implementation strictly a allow to less higher of they option are hypothesis with because conditions better the sometimes experimental test a as to albeit be parameters way conditions, may elegant of wild regeneration, estimation the natural in the in as occur variance in conditions sites they same between stress. as translocated the to processes are in response seedlings grow trait/survival or and and seeds divergence field whereby wild, about (RTE), the drawn Experiments Transplant in conclusions Reciprocal seedlings the by in experienced biases conditions possible replicate L´opez Pineda-Garc´ıa to exactly 2008; 2003; leading not nevertheless Markesteijn Kursar may & stresses & experimental (Poorter Villemereuil Queenborough (Lopez counter-intuitive (de 2008; proven give interest Markesteijn not of or & is traits Poorter in adaptation 2003; the signals however, Kursar adaptation compare & local to S´anchez-G´omez (Lopez Brousseau for possible & species 2006; make (Valladares or driven They 2020), habitats genetically performance different assess practice. Barton on to from common 2013; meant inferences is individuals stresses make of populations, environmental to in performance of way variations application straightforward response experimental (Baraloto a phenotypic the populations is without and or growth species with and tree survival among seedling factors differences environmental of the study of understanding The the to insufficient due and mainly power, trees, adaptation. statistical in local elusive low determining remain studies, adaptation long-term local identifying heterogeneity, and of environmental survive, phenotypes, between understand lack to and links to species than causal paramount of the die is However, distribution seedlings to Donohue spatial patterns. young chances 2006; distribution in higher phenotype conditions much Hampe and micro-environmental has species & local seedling to (Petit given adaptation selection any of that natural signals expect of must action one the though for (Donohue opportunity environment ample their to maladaptation and 2017). tal. et 06 Barton 2016; tal. et 07.Teedffrne npromnecmoet ie uvvl rwh n/rreproduction) and/or growth, survival, (i.e. components performance in differences These 2007). tal. et 04 aril iht21) nadto,RE aeas hw htsakmorphological stark that shown also have RTEs addition, In 2017); Pichot & Latreille 2014; tal. et tal. et 00,ppltos(Ram´ırez-Valiente populations 2020), 06.Teerslsso h opeiyo h eainhpbtenenvironmental between relationship the of complexity the show results These 2006). 00;wieteepce inl flclaatto o tlatsm environmental some least at to, adaptation local of signals expected the while 2020); tal. et 06 rgt20;Pizano 2007; Wright 2006; vs. oege’cniin n n ie ouainprom etri its in better performs population given any and condition) foreigner’ tal. et vs. 05 Pluess 2015; wy odto)(aek br 04.Lcladaptation Local 2004). Ebert & (Kawecki condition) away’ tal. et 2 00.Sc ihmraiyrtssol provide should rates mortality high Such 2010). tal. et tal. et tal. et tal. et 06 ahae rml 06 Rellstab 2016; Premoli & Mathiasen 2016; 05.Teaayi fcmo gardens, common of analysis The 2005). tal. et 09 Carsjens 2009; 01 les&Wbr21;Smith 2012; Weber & Pluess 2011; tal. et 06frarve) nsvrlcases several In review). a for 2016 tal. et 09 Queenborough 2009; 01.Cmo adn and gardens Common 2011). tal. et tal. et 04 Barton 2014; tal. et tal. et 06 Vizca´ıno-2006; 09 otest to 2009) 00.Even 2010). tal. et 2009) tal. et tal. et tal. et e.g. et Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. rwh ebvr eec,adsria,o obnto fteefcos r etrfrec ctp and explain ecotype to contribute each differences wild. for such the that better in argue are we observed habitats; considering factors, patterns other RTE, these distribution in germination, an of niche than that origin combination the established hypothesise of a We we habitat canopy. or its natural habitats, in survival, the provenance with under and ecotypes field defence, the two herbivory in the growth, region, provenance of and association habitat observed both the of spatial in causes heterogeneous higher the was thus globulifera mortality and S. Nevertheless, ecotypes, tell. two can the tions between flooding of and distribution drought to (Baraloto responses experiments siological garden common ( the shadehouse syngameon today Previous oak between covers the differentiation term of genetic members This The among 2019). species. than despite Petit different interfertile & are as (Suarez-Gonzalez (Cannon that classified unit representatives species evolutionary groups plant interbreeding related distinct inclusive closely believe morphologically most describe to of us to leading presence 1971) field, Grant the the 1917; in (Lotsy occurs century ecotypes Tysklind two XXth (N. the observed of been mixing have some adaptations that etc.) that with F2, suggesting (F1, associated data) variants individuals unpublished intermediate genetic – genetically show but, may habitats, and different (Degen paths to evolutionary (20-50m) different thus potential follow of ecotypes dispersal population pollen American South al. the and et African than than case so smaller less extreme although patches an mosaic constitutes al. in system et The distributed (Richardson known. are adaptation yet potentially, ecotypes not and Baraloto differentiation, is (; microgeographic determined roots of genetically prop or or plasticity pneumatophores phenotypic of presence the (H´erault bark, their of Schmitt texture the fruits, and Schmitt 2015; showed also by studies out pointed These was habitats (HT). as (SF) gradients, hilltops such flooded and across seasonally preferences slopes niche between on opposite gradient habitats display water Alli´e often surrounding widespread temporary species to other the congeneric tolerance all that from of and result gradient streams scale a along and local porosity at soil distribution of shield saturation Guiana (P´elissier water the saturation on prolonged has composition to community drainage of tolerance tree water mosaic the of in driving Steege complex factor Differences ter and ecological characteristics. variable 1991; main soil (Barthes highly a and a as identified show topography long Guiana, in been French variations construction in to the those linked for as microhabitats such essential forest, is species/phenotypes. Neotropical turn of Lowland distribution in spatial which heterogeneity, of spatial models predictive for of mechanisms the understanding (Wadgymarfor factors ecological covarying of subsets s’ctps,aefudi ypty fe ihitrige rws u r niomnal segregated, environmentally are but crowns, intermingled with often sympatry, in with found are ’ecotypes’, as as identified globulifera, S. Symphonia .globulifera S. 07,o vnta etoia ouainof population Neotropical than even or 2017), tal. et 04.Tetoeoye r eeial ieetae ae nncermcoaelts( microsatellites nuclear on based differentiated genetically are ecotypes two The 2004). tal. et ta.b nreview in b. al. et 21) yia xml spoie ytegenus the by provided is example typical A (2015). .sp1 S. edig eddt uvv etri Faesi h id(Baraloto wild the in areas SF in better survive to tended seedlings iepedars h etois n opotxno e neemndsau,currently status, undetermined yet of morpho-taxon a and Neotropics, the across widespread ta.a nreview in a. al. et 01.Wehrteemrhlgcldffrne r h rdc fevrnetlydriven environmentally of product the are differences morphological these Whether 2011). Symphonia prtsa ygmo nFec uaa ygmoswr endb oait uigthe during botanists by defined were Syngameons Guiana. French in syngameon a as operates en togylne oS aias while habitats, SF to linked strongly being hc skont cu nteGin hed dlso h w aa eerdhereafter referred taxa, two the of Adults shield. Guiana the in occur to known is which , tal. et tal. et sntepandb hs ln,a es sfra xeiet natfiilcondi- artificial in experiments as far as least at alone, these by explained not is ,a ela ieecsi aiu imtra rathih n rwhrates growth and height breast at diameter maximum in differences as well as ), 02.Alteesuisso httems tiigvrain nte species tree in variations striking most the that show studies these All 2002). 93 Sabatier 1993; ) h w ctpsaedffrnitdb h ieo hi evs flowers, leaves, their of size the by differentiated are ecotypes two The ;), F tal. et ST tal. et .globulifera S. SP)01 (Lang (SNPs)=0.13 97,alwn opsto h pce ln gradient a along species the position to allowing 1997), S 3 07.Sc yeo prahi hseteeyuseful extremely thus is approach of type Such 2017). . sp1 tal. et fe i ek fcnrle odn,adwild and flooding, controlled of weeks six after .sp1 S. Symphonia ( .globulifera S. F tal. et ST 07 ol o ipitdffrnilphy- differential pinpoint not could 2007) sfudo ohH n F(Alli´e SF and HT both on found is 018 Dc eet 08Tetwo 2008)The Heuertz & (Dick =0.138) 08,o hc a soeo h best the of one is oak which of 2018), tal. et fArcnoii,wt h species the with origin, African of tal. et Symphonia 2018). ( F ST tal. et 04,a re ftetwo the of trees as 2014), 03)(Torroba-Balmori =0.31) 07.T isc the dissect To 2007). ctpsi smaller is ecotypes F ST tal. et =0.086), tal. et 2007 Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. 08% 010)adte edighrioyatc ee a siae steaeaeo h percentage the 40-60%; of 20-40%; average (0-20%; the area as damaged estimated of as was percentage determined level was of attack Herbivory herbivory classes leafiness. seedling five seedling architectural then of was an of and one height As indicator 80-100%) measures. assigned Seedling two an 60-80%; was millimetres the leaves”, dead. in of leaf of mean found measured each the number was to but follows: as “total diameter estimated transplanted living Stem selected and seeds collar. we directions previously ungerminated the orthogonal trait, seedlings two and for in Individual for bud collar 0 1). apical the and and Table the at alive germinate, (Suppl. between 2012 centimetres found to in for seedlings yet measured except for and 2014, 1 until sites follows: 2009 field as from September recorded trans- ( in at was individual yearly collected survival each seedlings were then for herbivory) germinated and and Data as date diffe- traits, 2). plant transplanted growth-associated without (Table survival, and seed habitat year, shadehouse germination ungerminated and the transplantation, as region, in transplanted plantation germinated allocated those were ecotype, had or Individuals provenance, that garden. attri- their those each randomly in on between were slots rentiating depending those slots gardens within six positions different garden, random each to to In assigned slots. were plant ten-seedling than gardens mother twelve 44 other site the in Regeneration over understory Field arranged to undisturbed. distributed all was gardens. left were buted transplanting, garden was Individuals 12 to canopy hand. each by the follows: totalling Prior yearly removed; as wire. removed site, was chicken was D.B.H.) each seedlings with cm transplanted in herbivores Fig. 5 the 2; large conditions to (Table up from respectively HT (i.e. fenced “Habitat”, in vegetation and was region” three garden “Plantation Each and variables 1). the SF compose in habitat and three plantation habitat: and west) hs set eadn h hs fe emnto,ad“niiul o hs set eadn ohseed both regarding aspects ( those for confusion, Individuals for “seedling” avoid germination, “individual” To to and timing”. prior germination, phases. “germination in phase after seedling as the stored phase and stored regarding the was was aspects regarding ( not etc.) discussing seedlings aspects when or 3, as those “seed” year transplant recorded to 2, first of the refer year were moment in hereafter seeds 1, we considered the individual year were which at transplant, status at germinated of moment germination moment had life The in incorporated status”. seeds the differences were “transplant in Whether how variable collection groups the shadehouse, for traits. seed in and in among other an biases rates, spent of trends as how germination time analyses included germination for of of below be of analysis amount See must the comparisons about analyses. themselves into information traits all, rates growth-associated carrying of germination and turn secondly, First history its bias; time, analyses. in this flowering cofactor, in in to important differences according biases ecological interpreted time. as from transplantation be viewed originate at must seedlings, to be germinated likely the can the ( are and by in groups germinated they collection reached among spent had stage seed differences days developmental that the such the of on seeds Although in number as depending of This the well shadehouse, in proportion 2009. as the groups July the ground), in among and 2a), days differences May at (Fig. substantial 315 between shadehouse shadehouse introduced and common sites This a 27 field in dates. between into soil transplant spent transplantation 35- with seeds to tree, plates that prior mother germination meant each Campus polypropylene From Agronomic in ’eastern’ 1). sown Kourou the from (Table in and the times, “Ecotype” trees collected flowering mother and were five unequal region” from seeds and largely “Provenance 39 region variables to ’western’ seeds. the the due the in composing 2009, for ecotypes French region, both April sites to of sampled and belonging the part 2008 two trees among mother at September driest not nine habitats between were the SF which collected and respectively, ’west’, were regions, HT Seeds and ’west’ on and installed Guiana, the ’east’ gardens French the and experimental in in onto habitats sites transplanted rainfall contrasting field were highest two Seeds 1). the from (Fig. with trees Guiana mother ’east’, from patterns: seeds rainfall collect SF- to ecotypes: aimed associated design experimental The collection: data and design Experimental Methods and Materials Symphonia i.e. edo edig eetaslne ogresetbihda ahfil ie( site field each at established gardens to transplanted were seedling) or seed termiigsoswr sdfrohreprmns,adte he niiul per individuals three then and experiments), other for used were slots remaining (the .globulifera S. n HT- and .sp1; S. 4 n rmtobodrgoswt akdydifferent markedly with regions broad two from and i.e. tlatctldn mre from emerged cotyledons least at i.e. emnto ttsat status germination i.e. east e.g. vs. Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. iermdl(M n eeaie iermdl(L)wr diinlyue ots h oeta effects potential the test to used additionally were (GLM) model linear generalized and (LM) model analyses: Linear model (Hot- linear ‘party’ general package branch. and the each models with in Linear 2020) observations Team 2 Core of the (R minimum of environment a course statistical and the R over the value were average in single The herbivory horn run a average 6). were Therefore, year individual the analyses (i.e. each analysed. as All 5 giving were well age individual, as 2014 at the leafiness in is of and that measure life diameter, experiment. 2014, final the height, in over a in trait summarised having (RGR) each also for rate and value growth field discrete relative the unique yearly a in 1). has or material individual shade-house (Supplementary each field the the in in predictor whether seeds all as of and planted the rates, individuals For germination of shadehouse rates on germination ecotype the and on region variables provenance of trans- effects removing the and ( evaluated variables interest explanatory of as covariates variables the predictor ( the status all including plant analyses the individuals. repeated We ‘local’ (e.g. or variables ‘home’ the of (Strobl of of combinations power performance vote certain prediction of majority if of covariates identify a those drop identifying significant to on by a us based importance, ensue predictions variable removed, relative response when assessing obtain which, allows to methodology variables the Such predictor of forest. strongest of each whole with number the (Hothorn repeated reduced found with then a be variable is can and predictor process response The the and groups. covariates for two between the value in al. dependence response significant split and where the no best tested; herbivory) until divide recursively, data, the to is groups, and used status) found, the and transplant leafiness, is retained of and is growth, dependence habitat, effect classification significant survival, region, a binary plantation germination, if ecotype, recursive here then, region, by provenance (i.e. here achieved response (i.e. covariates is the This of predictor variables. independence the predictor variables among among on interactions interactions complex dependent and untangle (Cutler relationships responses. nonlinear (RGR), the expected where of data rate are each for growth suited on particularly effects relative are their experiment 2001) yearly the visualise of average graphically end and the the variables, at and individuals on 6) herbivory and (year leafiness, growth, survival, germination, explaining forest effects. Random of confusion the analyses: reduce forest named and Random also meaningful comparisons the means, such on least-squares make variables the to predictor on effects of based effects test general vs. a Linear find b) introduce to (Searle and we means used variables; adjusted specifically, were dependent interac- More (GLM) the of variables. on model structure effects dependent linear the their understand Generalized visualise of and random and graphically untangle significance a) model and importance, variables: biological covariates, variable predictor the the explore among studied extract to tions the to used on were data depending methods the performance forest to and survival, applied germination, were individual strategies analytical complementary Two 1). analyses: Table Data (Suppl. leaves its all of area damaged of 06.I a In 2006). aa’hbttado en ‘local’ being of and habitat ‘away’ tal. et admforest random 06 Strobl 2006; i.e. admforest, random ehd n lsicto re eeapidt vlaerltv aibeiprac in importance variable relative evaluate to applied were trees classification and methods edo edig oceki rnpatsau ofuddteaayi fteipc of impact the of analysis the confounded status transplant if check to seedling) or seed tal. et nlsso rwhpromneadhrioy nyidvdasgriae n2009, in germinated individuals only herbivory, and performance growth of analyses tal. et tal. et i.e. 07.Casfiaintesvsaiepeitv oeso epne significantly responses of models predictive visualize trees Classification 2007). 90,wihalw st opr h ffc fgoigi hi hm’habitat ‘home’ their in growing of effect the compare to us allows which 1980), 09.Cniinlifrneteswr logoni at ihan with party in grown also were trees inference Conditional 2009). rvnnergo,eoye lnainrgo,adhbtt.Frhroe we Furthermore, habitat). and region, plantation ecotype, region, provenance h bv lsicto rei efre nabosrpsbe ftedata the of subset bootstrap a on performed is tree classification above the vs. frinr nagvnhbtt hl vrgn o te potential other for averaging while habitat, given a in ‘foreigner’ 5 .globulifera S. nS)la osgicn improvement significant to lead SF) in admforest Random tal. et 07.I u ae tallows it case, our In 2007). ehd (Breiman methods α 0.05 = et Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. hr ( where for. accounted be should provenance and and collection ecotype the ( on effects: germination depends of genetic shadehouse probability were the the for of in account logit spent to The time effects ( the However, success only germination the region. in therefore, difference transplanted, the ( been of germination evaluation overall the and In shadehouse the in subscripts success bear Germination they that except identical, are case region here. plantation / region o provenance subscripts the with for case, formulas ecotype/habitat success the response for of formulas normal probability of a with as distribution expressed Bernoulli be a might herbivory) and μ leafiness, Ν( diameter, stem height, seedling a ( status and transplantation east better for of 1 advantage = comparative the has h approach power. relative LM/GLM greater denote population having our our We one possibly, (2004), of 2004), and, superiority design Ebert Ebert global unbalanced and & of with Kawecki and (Kawecki coping combinations”. of adaptation others deme-habitat local wording ‘allopatric’ all “true” and the “home of to ‘sympatric’ In “ effects the of the of means interactions). confounding the analysis environment While comparing separate x to the tantamount population is on for method rest test which with they contrasts adaptation, This described , local or as performance. provenance of individual approach, of for detection environment Our proxies vs. the the as taken in for design. traits planted strategies final on been itself, previous having unbalanced in of both environment, the effect different test of the a single in observing effect a synthetically the in at disentangle subsuming aims by to below, adaptation effects local other test possible to procedure ad-hoc ‘home an the define to importantly ( traits most on variables predictor different of ieo emnto.Tersos smdle hog emti itiuinadtelgln function. link log germination. the of and time the study distribution expected to the geometric used don’t compare a is we to approach through that GLM used modelled except a are is success, germination Means germination overall response Square of the The Least case study In germination. to shadehouse. of used the time is in means time the square for least account using approach same The ( ecotype of effect the ecotype: shadehouse, of the ( means in success square time germination in least in difference classical difference and the design on unbalanced ) of effects the correct To Where o rvnnergo and region provenance for o h g ftecniee niiul( individual considered the of age the for o T ota niiul rwn t’oe r pcfidb 1o 22), or 11 by specified are ’home’ at growing individuals that so HT, for 2 = wy n “local and away” εηορσα t t ´ steaeaetm pn nteshadehouse. the in spent time average the is ) tnsfrtetm pn nteshadehouse. the in spent time the for stands vs. σ , e h ctp ( ecotype the 2 wy,adte‘local the and away’, oi ( logit ), hl iayrsosslk h iehsoytat,griainadsria,aeepesdas expressed are survival, and germination traits, history life the like responses binary while o o west), for 2 = p vs. || d,G adj, s oege”eet n euetepeec fteeetfo lp oprsn ( comparisons slope from effect the of presence the deduce and effects foreigner” ftaslne sase or seed a as transplanted if 1 = e || for 1 = r e = ) o lnainrgo,a ecie bv,adwl o efrhrdescribed further be not will and above, described as region, plantation for r u logit vs. G o h lnainrgo ( region plantation the for .globulifera S. + oege’tsso aek n br 20) hl vrgn vrall over averaging while (2004), Ebert and Kawecki of tests foreigner’ i.e. α p G eok G G e a emnto,sria,got,acietr,adhrioy,and herbivory), and architecture, growth, survival, germination, G +
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(Fig. variance the germination the the of of in of timing analysis drivers time main The in the difference were 3). ecotype, the than for rather provenance accounting habitat, western when x ( for germination even ecotype overall identified of 2b), ecotypes was different analysis two shadehouse ( the (Fig. the the house of groups shade in among shadehouse the success other the in shadehouse germination time in average of the success the probability after germination at provenance lower of of time probability region the the of on compared depending we duration 2a), in (Fig. seedlings bias groups Symphonia the of survival overcome and To germination the on both transplant covariates in two analyses. observed of the the was field, Effects of tested the rest in effect germinated the main have for the that together Since seeds considering the merged when experiment). and were GLM the shadehouse statuses of using in germinated years observed have 6 was that the result seeds during same ( point, The germination strong some shade- field. at of indicated the the in success 1) germinated in global having material germinated seeds the (Supplementary having status: germination seeds transplant the of or of house success independently region the provenance seedlings. experiment the of of the vs. effects analysis of seeds tree end survival: classification the on The at status alive transplant still the the over and of herbivory 2009 Impact statistics average in Summary and germinated experiment leaves) individuals 2). 3. the (Table of those Table of gardens number in for end SF reported years total seeds. the are western 6 diameter, as At in the height, transplanted lowest of 1). were was (Table (i.e. course remainder survival experiment traits The and the growth-associated alive, of seedlings. of were end as seedlings the transplanted of by were 37% 61.2% and reached transplant germination of Overall time the at individual 510 of total A data: the of Description Results 2010), Weisberg & (Fox ‘car’ packages the with environment statistical 2009). R the (Hothorn in run ‘multcomp’ were analyses GLM All tal. et vs. Symphonia at months). 3 east: 08,‘men’(et 06 08,advsaie sn gpo2 (Wickham ‘ggplot2’ using visualized and 2018), 2016, (Lenth ‘emmeans’ 2008), i.e. i.e. hdhueadfil obnd niae htpoeac n plantation and provenance that indicated combined) field and shadehouse eefloe vrtecus f6yas ftee 61 a germinated had 36.1% these, Of years. 6 of course the over followed were htvrtetasln tts l h ed hthv germinated have that seeds the all status, transplant the whatever 8 i.e. .globulifera S. 9dy) significantly A days). 89 .sp1 S. oprdt all to compared ie ‘foreign’) (i.e. .sp1 S. Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. eandvr o ilteedo h xeiet(i.2) nsakcnrs,alfu atr provenance eastern four all contrast, stark Western In success. germination 2c). overall 100% (Fig. nearly had experiment east the the in western western of planted for habitat of end and germination germination ecotype the of that The other combinations till indicating field. all low 2c), very the than Fig. days remained in western (˜70%, of 89 while high experiment recovered after the relatively of shadehouse HT and was end the west 2 the Western of at the (Fig. patterns: germination in conditions germination overall However, unexpected controlled of 2b). the some (Fig. probability groups under exposed overall germination 5) and of shadehouse 4, probability estimated 3, the Fig. and Suppl. trees classification adaptation. The caused local exclusively of patterns not germination: the are of for traits Patterns coding measured genes the underlying that the revealing in complex; trade-offs however, genetic by are, suggest patterns traits The growth-associated 2004)). and ‘home’ life-history both in find individuals the yet ’local’ and in and found, that ’home’ ecotypes occasionally of two are performances hybrids our Superior way, genetic properties. ecological and a respective morphological in their where retain stands, trees; they neighbouring mature or of mixed in distribution underlying as the interpreted typically in Symphonia are of differences gradients ecological evidence interspecific Ram´ırez-Valiente along and observed performances 2007; plants seedling (Wright in in trees Differences recognised 2020). for widely accumulating are is significance Brousseau adaptive processes its adaptive and microgeographic differentiation genetic 0 Local age at x diameter provenance their 5h). ‘home’ to Discussion (Fig. their relative combination in plantation 3-5 grown x ages provenances Individuals for ‘away’ at in ages diameters 5c,g,k). individuals early larger to but the (Fig. significantly compared individuals, years in ‘away’ had outperformed later effects combination, individuals in significant, plantation ‘home’ near disappeared where leaves, or significance of x significant, the number total ecotype showed differences and ‘home’ analyses the diameter, that plantation height, their and x in combination, provenance grown habitats The individuals x 5a,b,e,f,i,j). of ecotype (Fig. ‘away’ 0) age in age with those at increase Figure to measures growth- view. compared contrasting reference the combination a ( to habitat of provide performances to compared design, growth comparison growth unbalanced yearly better relative a significantly for on the correct focused illustrates but to 5 results west used RF the were the in corroborated Means planted tions) ‘home’ individuals LS comparing with individuals where 4d), traits of analyses, height associated (Fig. the that Model east. diameter in on the and Linear RGR in impact The 3), the those an on node than had and thicker 4e region 3) and plantation 3, node taller Finally, 4a node being 5n). (Fig. that 4b 4g, HT 4a), than in (Fig. herbivory (Fig. (Fig. planted less habitats HT HT suffered SF in in and in than than 5i,j), planted SF 4e, SF in average growth- in (Fig. on leafier 7 leafier taller but were of 5e,f) individuals out 4c, that: 5 (Fig. in thinner RGR on globulifera average was effect S. and 5a,b), leaves, 4b, significant separating of herbivory, number (Fig. strongest relative total faster average the diameter, and height, leaves, had RGR of average ecotype number herbivory: trees, and seedlings traits classification Symphonia associated the of to traits growth-associated According on variables predictor of Effects Symphonia tal. et ygmo eaei pce-iewyrltv ohbttpeeecs ie htte a grow can they that given preferences, habitat to relative way species-like a in behave syngameon Fg g n.Hbttas a infiateet nfu rwhascae ris revealing traits, growth-associated four on effects significant had also Habitat 5n). 4g, (Fig. re aelclyaatdt ieetevrnetlcniin cosFec uaa We Guiana. French across conditions environmental different to adapted locally have trees 03 05 08 Carsjens 2018; 2015, 2013, vs. aa’ n ’local’ and ‘away’, vs. aa’( ‘away’ vs. frinr xmlso oa dpain( adaptation local of examples ’foreigner’ tal. et i.e. 04 Pluess 2014; ctp aia n rvnnexpatto combina- plantation x provenance and habitat x ecotype 9 e.g. .globulifera S. tal. et .sp1 S. egt imtr n N,a ela their as well as TNL, and diameter, height, .sp1 S. .sp1 S. 06 Rellstab 2016; nohrhbttrgo combinations habitat-region other in uee h es ebvr when herbivory least the suffered .globulifera S. from ed aeasgicnl lower significantly a have seeds .sp1 S. tal. et sensu Fg 4). (Fig. .sp1 S. rwfse n was and faster grew 06 Barton 2016; .sp1 S. aek Ebert & Kawecki lne nHT in planted .sp1 S. tal. et nwestern in 2009; tal. et grew Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. dpainars w aibe bt aia n einl,srsigteecc fteslcinpressures selection the in of of germination performance eastern and efficacy lower survival where the overall in stressing effect Variations an regional), regional ‘local habitats. and a a HT habitat of also eastern west. example (both in but double variables drier HT, two a the survival drier across constitutes in high to case adaptation better adaption relatively This cope tolerant. ecotype a drought to an suggestive with particularly able of is west be only the group therefore not may indicative in first western and 3). planted The only Guiana is group (Fig. French 2c). it Furthermore, only throughout habitat (Fig. hilltops the and rates in is ecotype common germination It of high adaptation. had regardless local ( also east specific groups the very survival of in high planted two individuals These eastern-provenance ecological and ( of survival HT, highest maintenance the the with and groups The Guiana French the in within distribution ecotypes between species of differences survival of of patterns predictor patterns explaining the the among potentially interactions explaining complex only reveal survival not and variables, germination combined of distribu- trees classification species The of patterns the understanding towards contributions tion: and survival of patterns Patterns survival in variance the explaining mechanisms the (Alli´e of adult one of rarity be could tree’s among which the observed we along vigour, membership species vigour in or individual variance status in ontogenetic shown as above variance been well where survival, has survival, predicting growth, variable (Aubry-Kientz tree expected important Neotropical and most the observed on was between effect life difference pervasive the a in have as to for vigour, habitat tree of Individual effect individual habitat. significant of any signal for performance detect that growth traits not how competitive did pinpointed growth-associated in trees in reduction classification decreases a the indicating and significant individuals traits, by outperformed growth-associated significantly dominated of habitat was terms home in their habitats in away individuals in how exposed analyses LSMeans The local traits: a growth-associated be of may Patterns time germination quicker a ( rate Conversely, (Dalling mortality ecotypes seed western in or overall. the differences that environment to indicating response harsher 3) in (Fig. a adaptation hypothesis, west such in the Supporting in are provenance-individuals 2014). Venable eastern plantations western & of in survival (Gremer reduced adaptation years observe several local we across a risk as mortality seedling emerged spreading have may of germination, germination for Symphonia delayed requirements for environmental advantages tighter evolutionary of or indicate germination may rapid of or study west populations our the among of adaptation genetic course local the the for and over evidence duration, as dormancy reported seed been in thaliana has Variation such variation, range. underpinning such cues for species for environmental conditions a and basis possible across timing differential vary best the may Such and the survival, conditions with average). seedling favourable germination for on in Matching paramount differences later cues. is to growth and year due seedling timing be 1 could germination combinations 2d: to ecotype-provenance adaptation (Fig. among germination local seeds of timing eastern and than rate success later significantly germinated seeds i.e. > tal. et 0 uvvla g ,cmae to compared 5, age at survival 50% Dnhe20;Postma 2009; (Donohue 05 .Shit nulse data). unpublished Schmitt, S. 2015; seilyin especially , .globulifera S. tal. et Symphonia tal. et 05.Vrac ngot efrac,sc sse in seen as such performance, growth in Variance 2015). 01 Postma 2011; .sp1¸ S. nH aias n h oegnrls itiuinof distribution generalist more the and habitats, HT in .sp1 S. .globulifera S. edig.Teosre ainei rwhi nedi codnewt the with accordance in indeed is growth in variance observed The seedlings. Symphonia tal. et samast oewt re niomn (Dalling environment drier a with cope to means a as hc infiatysprt rmalohr ntrso uvvl perhaps survival, of terms in others all from separate significantly which > 05 06.Smlry h einldffrne ngriainsuccess germination in differences regional the Similarly, 2016). 2015, 0 fe er)wr western-provenance were years) 6 after 50% Symphonia tal. et < ntees Fg d.Ahge edqisec rdrac level, dormancy or quiescence seed higher A 2d). (Fig. east the in 5 o h eto rustaslne ntewest). the in transplanted groups of rest the for 25% edig hnpatdi T ohpoal otiuigto contributing probably both HT; in planted when seedlings 2015) .sp1 S. syngameon. 10 ugsigacpct opromwl eadesof regardless well perform to capacity a suggesting , Symphonia e.g. ebvr,dsae raen)btenregions between ageing) or disease, herbivory, .sp1 S. niiul noreprmn,btalso but experiment, our in individuals .globulifera S. .globulifera S. r utemr copne with accompanied furthermore are .globulifera, S. vs. oeg’eiec flocal of evidence foreign’ .sp1 S. .sp1 S. nH.Cnesl,we Conversely, HT. in hnpatdi HT, in planted when a einterpreted be can lne nwestern in planted tal. et coshabitats across Symphonia Arabidopsis 01,and 2011), .sp1 S. .sp1 S. are in is Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. is hr iia atrsaeosre nRE ewe pce pcaiigi ihadlwherbivory low ( environment and herbivory high higher in a specializing to species exposed stu- between normally previous RTEs of are to in those which observed with species agreement are environments: in patterns pressure are similar analyses where herbivory Our dies, (Lamarre feeders ecotypes. SF leaf between in where strategies than different, potentially avoidance significantly HT 5e), are in (Fig. habitats abundant years environment.bulifera HT more 6 risky and are after a SF particular in diameters Guiana in growth French largest maximising in the strategy assemblages had a Arthropod and towards 5a) adaptation accordingly. (Fig. ecotype allocations an tallest habitats resource of the SF their indicative were in adapt living SF (Ferry therefore trees must in fall however, habitats seedlings tree habitats, SF HT through in than death specializing light SF. Species of to and access risk HT and between their fertility significantly double higher vary a herbivores have of habitats presence SF and community, presented microbiota microhabitats soil the and across covary floristic variables other many ( regime, water here in variations sharp (Baltzer an the with efficiency Beyond consistent use pattern water a improved show through results contrasted Our mediated the regime. potentially of hydric western effects influencing combined of factors the advantage soil to adaptive on patterns focus adaptation a detect with to habitats, designed adaptation: was setup local experimental of The signals the the them. behind between to occurs pressures contribute hybridisation Selective could occasional which when even ecotypes, ecotype between each of differences integrity ecological genetic the the of of preservation maintenance The in the habitats. sizes explain maximum SF larger helps to the of limited because specialist output habitat sp1 reproductive higher a potentially is a and it for rate availability that the habitat suggesting matches in herbivory), and limitation higher behaviour, apparent and generalist height, habitat in a Conversely,RGR suggests distribution. which environments, species other extant on penalisation no that has evidence find divergent we between Overall, quality. confusion potential stressful habitat the less in exemplifies and which a differences gardens, east, infer and the plantation we selection in east habitats gardens, for or in these ecotypes site individuals in between field survival survival of eastern affecting survival the significantly high in factors general environment other the of Given ‘home non-appearance germinated. of the once example east an ‘local constitutes the not individuals, This in to but eastern-provenance significantly. level adaptation, of drops regional local west survival the of the the at pattern in in as adaptation planted individuals tolerance, local ecotype, eastern drought either of of of poor regardless indicative alternatively, performance sites, or better field rainfalls, a western heavier to or confirms compared survival experiment sites the high (Miglia field in comparatively eastern history for with life account group trees’ not second the did The across we variables missed dryer environmental we in to stages related performance to be improved related could its conditional latter underlying The 1) basis gardens. either costs penalising genetic that without pressures the environment suggesting selective in one Wadgymar habitats, play in 2013; SF for advantage at (Anderson in cost performance conditions is a nor adaptive environments) conveys gardens an alternative adaptation eastern detect in an either not whereby in did (i.e. We survival neutrality lower ecotypes. two of the form between distributions contrasted .sp1 S. loigteceitneo oheoye.Tevrac njvnl efrac ntetohbtt also habitats two the in performance juvenile in variance The ecotypes. both of coexistence the allowing i.e. eadeso l te oaits n oetfor lowest and covariates, other all of regardless east ,eprecdrdcdhrioyi o ebvr niomns( environments herbivory low in herbivory reduced experienced ), vs. et SF west, vs. T:acs orsucssc slgtadsi uret,ters fdah the death, of risk the nutrients, soil and light as such resources to access HT): .sp1 S. .sp1 S. .sp1 S. otedis odtosicue nteeprmn ie etr HT), western (i.e. experiment the in included conditions driest the to a etrsria ntedis odtos uesls ebvr,and herbivory, less suffers conditions, driest the in survival better has nwte odtos uha hs on nS aiaso eastern or habitats SF in found those as such conditions, wetter in Symphonia tal. et .globulifera S. vs. oeg’a etr-rvnneidvdashv ihsurvival high a have individuals western-provenance as foreign’ .globulifera S. 07,o ht2 u xeietldsg i o atr the capture not did design experimental our 2) that or 2017), tal. et ngnrl eddntcpueeiec fdffrne in differences of evidence capture not did We general. in 11 stil eaie u fS ( SF of out penalised triple is .sp1 S. 00,cetn ihrs ihgi environment. high-gain high-risk a creating 2010), ol ecmestdb h atrautgrowth adult faster the by compensated be could tal. et nS aias ugsigdffrn predator different suggesting habitats, SF in tal. et 06.Hrioywshgetfor highest was Herbivory 2016). tal. et 2005). 2005). i.e. iia oS)cmae to compared SF) to similar i.e. .globulifera S. oe N,lower TNL, lower .sp1 S. .globulifera S. i.e. vs. similar than .glo- S. nthe in away’ S. Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. h c-vltoaypoessudrinn h iest n h pta tutrn fNorpcltree syngameons of Neotropical maintenance of the in structuring involved processes spatial the the of sympatry. understating and in our diversity furthering environmental the well mosaic as underpinning such communities the a suit processes to Overall, determine eco-evolutionary traits and geneflow. that differentiation life-history the small evolutionary different occasional filters the relatively with to despite even sympatry ecological lead that in heterogeneity other can and taxa suggest SF, stochastic the distinct and processes results exclusively HT of on our an selective between maintenance Furthermore, between those distribution, of the as habitats. being link tree such to across from differences, a adult environmental distribution way far and suggest of stages, significant life pattern hand, therefore a early species’ one results in at the Our occurring contribute adults on processes syngameon. nature, in that seedlings, coexistence the observed indicates the in of allowing and also survival thus, hand, maintenance and pattern and the attained, a growth is and biomass differential habitat), ecotypes adult x greater both if (ecotypes output for groups reproductive survival. higher such seedling in other results and phenology outgrew germination they Symphonia of habitats coevolution revealed SF of have in result We the environments. provenances, be natural between globulifera may foreign and and ( and ecotypes risks), constraints own environmental between herbivory known their differences with trait in match that growth-associated germination individuals differential and of governing when life-history mechanisms stages, cohorts ecological significant life the (Donohue of into early recognised survival insights during us been and given expressed has has differences mortality, experiment genetic RTE highest Our on the selection experience natural trees of importance critical The Conclusion: included. be should reproduction and growth, including atic comparisons multi-life-stage variables, the environmental in across play som taxa diffe- at related germination, regulation is of / mechanism performance as expression which and magnitude gene tell vival environmental and of to divergence to Miglia order hard genetic by reactions is same stressed about seedling it the information in ecotypes, in precise differences between of still to rences absence contribute probably the also is In compensate may mass ger- cues. to inheritance in seedling meant Epigenetic Differences is because ( mass. biases. effects developed resources, seed multiple Maternal we and from results. method suffer growth the analytical may affect seedling the results still their may While they of unbalances. them, meaning to the lead and successes tests mination our of power The study: the of of Limitations (Bryant rate compen- habitats herbivory to lower HT the compounds in explain toxic resources also and in would unpalatable limitation scenario of potential production a the for increasing sate by (HT) environment herbivory between occur also nimcoiladat-aaii opud (Cottet compounds anti-parasitic and anti-microbial atclryrc ngouiatoeE(Cottet E fack (Ondeyka globulixanthone organisms in other rich particularly in (Fine properties environments insecticidal low-herbivory have from species 2012). and Davies environment ‘home’ their tal. et 2002). edig eepnlsdi Thbtt ihrdcdgot n ihrhrioy however, herbivory, higher and growth reduced with habitats HT in penalised were seedlings ugsigthat suggesting , .globulifera S. .globulifera S. tal. et .globulifera S. 20) ogi opeesv nesadn fteeooia atr rvn sur- driving factors ecological the of understanding comprehensive a gain to (2005), ise r ihi eodr eaoie fthe of metabolites secondary in rich are tissues .globulifera S. ouain rmCmro n rnhGin ie o hi otn in content their for differ Guiana French and Cameroon from populations and .sp1. S. a pcaie opttv datg nS aias hc may which habitats, SF in advantage competitive specialized a has tal. et ne hshypothesis, this Under i.e. 04,wihhssrn nimcoilatvt (Nkeng- activity anti-microbial strong has which 2014), 12 tal. et .sp1 S. yrcrgms uretaalblt,dahrs,and risk, death availability, nutrient regimes, hydric e.g. Symphonia tal. et 07,sgetn htceia ieecsmay differences chemical that suggesting 2017), aenlpoiint ed)mysilinfluence still may seeds) to provision maternal nS aiascmae oH habitats. HT to compared habitats SF in 06 Wezeman 2006; tal. et oe utesorcmrhninof comprehension our furthers model .sp1 S. tal. et 95 Fine 1985; b a aeaatdt higher a to adapted have may Symphonia 00 ota& Postma 2010; sxnhn aiy nw to known family, is-xanthone tal. et tal. et tal. et 04 06 ate & Baltzer 2006; 2004, 05;telae are leaves the 2015); ytm ial,as Finally, system. 04 06.Such 2006). 2004, ge 2016). Agren ˚ S. Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. ate L aisS 21)Rifl esnlt n etpesr sdtriat ftoia respecies’ tree tropical of determinants as pressure pest and seasonality Rainfall (2012) distributions. SJ Davies JL, Baltzer ohe ,SeatJ(05 o oa slcl dniyn h cl faatv aito nah(Fraxinus ash in variation adaptive of scale the Identifying nursery. local? the is from local results How L.): (2005) excelsior J Stewart tolerance drought D, constrains Boshier adaptation Local tree. (2020) foundation T in Knight tropical AB, (Caesalpiniaceae) a Shiels species habitat in KF, Edwards Eperua and C, two Jones tolerance of KE, Barton distribution stress Guiana. the French water on of conditions Seasonal forests soil rain (2007) of the B Influence Ferry (1991) B L, Barthes genera. Blanc tree D, neotropical four Bonal within F, associations Morneau C, contrasting in Baraloto seedlings tree tropical among trade-offs Performance (2005) D microhabitats. Bonal DE, Goldberg Physiological C, trees: Baraloto transplantation. tropical reciprocal in specialization to Edaphic responses (2005) DFRP and Burslem correlates R, Nilus SC, Thomas JL, Baltzer dynamics. forest growth tropical coupling of model component individual-based key a joint is A vigor (2015) species tree H´erault, B on that J-J, change reveals Boreux mortality climate and V, of impacts Rossi estimate M, to Aubry-Kientz models niche using for framework distributions. A (2013) RP Anderson community. J Engel P´elissierAlli´e R, E, manuscript. the of acceptance upon database Bibliography: plant TRY the to submitted be will data The the wrote publication. MPE availability: for approval and Data final CSS, their gave IS, soil and expe- NT, with drafts RTE analyses; contributed previous the data BF to established the critically campaigns. SOC contributed conducted measurement authors and MPE field All VT and the manuscript. MC, NT coordinated characterisation. experiment. SOC habitat the and and VT designed LB, and AT, idea MC, the riment. conceived CSS and IS Guyane. Universit´e was contributions: de the thesis Author AT at Guyane, out Region carried were the theses and Both (BDI) “Investissement CEBA. CNRS grant Labex the and Recherche by FEDER financed la UE was ENERGIRAVI) de by (PO-FEDER Thesis financed Union greatly National MC have European support. Agence that the financial drafts and the for ANR-10-LABX-25-01) previous thank ref. on CEBA, comments kindly (Labex for We d’Avenir” Heuertz 2009-2014). of manuscript. (years Myriam species seedlings and the this master related the Schmitt AgroParisTech measure improved analysed Sylvain closely many helped to the within and that thank thankful divergence We module experiment are the thesis. Humides” We AT’s RTE Tropicales of “Forets of “Genetic the the part thesis from includes started students his also manuscript who of This part MC, trees”. in tropical to measurements of dedicated years posthumously first is manuscript This Acknowledgements: 5 2457–65. , LSONE PLoS clg n evolution and Ecology naso h e okAaeyo Sciences of Academy York New the of Annals Ecology , 86 , 10 tal. et 2461–2472. , ora fEcology of Journal 1–16. , 21)Praielclsaete-olhbttascaini rpclforest tropical a in association habitat tree-soil local-scale Pervasive (2015) eu ’clge(er tl Vie) la et (Terre d’Ecologie Revue , 2 oety nItrainlJunlo oetResearch Forest of Journal International An Forestry: 2682–94. , . Ecology 13 Ecology , 88 , 1297 478–489. , , 86 8–28. , 3063–3077. , , 46 303–320. , clg n evolution and Ecology , 78 135–143. , Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. ei iegnewti otnosppltos h aeo h etoia reEeu act (aubl.). falcata Eperua ge- tree microgeographic neotropical of the drivers of adaptive case and The ONE Neutral populations: (2015) continuous I within Scotti divergence C, netic Scotti-Saintagne M, Foll L, Brousseau (Fabaceae). Aubl. falcata Eperua tree Amazonian E Dreyer PVA, intrapopu- Fine trees. L, promotes forest Brousseau variability rain environmental tropical local from Highly example (2013) An I 1169–1179. traits: quantitative Scotti of J, divergence Cigna lation D, Bonal L, Brousseau Forests. Random (2001) L Breiman annC,PttR 21)Teoksnaen oeta h u fisparts. J its of Guzy sum Q, the Ngoc than Nguyen more syngameon: C, oak Carsjens The (2019) Defense. RJ Antiherbivore Petit Plant CH, and Cannon Availability Resource (1985) PD Coley 230 FS, Chapin JP, Bryant oet uhr(s) alV.A.Fn tl eoe n hli Coley. . D Phyllis and Mesones Amazonian forests. Italo I in Amazonian , Trees Mesones in Fine by plants ZJ, . Specialization A Miller Habitat PVA, . Promote V Fine Paul Herbivores ): (2004) s PD ( Coley Author Forests I, forest. waterlogged rain Mesones and tropical slopes PV, a on in Fine rates productivity treefall and Higher biomass (2010) stand V lower Freycon in L, result Blanc soils JD, Bontemps F, Morneau experiments. B, Ferry transplant a long-term in in pinaceae) herbivory Botany rigida, leaf of (Pinus of pines GA rates pitch Fox normal-stature determines DR, Taub species W, Seedling Fang (2006) SE forest. Hartley rain SG, Malaysian Postgermination Compton Germination, MP, (2010) Eichhorn CG Willis K, Ranges. Kovach L, Ecological 293–319. Burghardt Species R, histories. and Casas life Adaptation, de plant in Rubio Sciences link Biological K, missing B: Donohue the Society as Royal effects the Maternal of cycle: Transactions the phical Completing Symphonia (2009) K TRO- in Donohue WIDESPREAD inbreeding A OF biparental HISTORY , BIOGEOGRAPHIC SPECIES. and COMPLEX TREE THE dispersal PICAL (2008) M pollen Heuertz Limited CW, of Dick (2004) effects H Interacting Guiana. soil: Caron French in in E, survival globulifera Bandou Seed B, (2011) community. A Degen microbial Arnold soil Elizabeth the BJ, and Schutte dormancy predation, AS, Davis JW, Dalling KH 2783–2792. Beard , TC, Edwards DR, Cutler LC–HRMS. tandem by M globulifera Kritsanida Symphonia E, Amazonian Kouloura K, Cottet globulifera. Symphonia tree rainforest tropical Y Fromentin ancient G, responses. 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Seedlings: Associated Tree a Traits of Ecophysiological S´anchez-G´omez Energy (2006) F, and Valladares D Death, Life, an (1975) of Van populations L Valen Neotropical and African species. in tree structure tropical genetic ancient spatial fine-scale K affect adaptation Heer crohabitat KB, Budde P, Torroba-Balmori perspective. plant A introgression: Adaptive (2018) QCB random , Cronk for C, measure Lexer variable-importance A, conditional Suarez-Gonzalez new, package. A party on! the Party measures: in (2009) available importance A variable forests Zeileis a forest T, in Hothorn random C, factors in Strobl Bias soil (2007) solution. and T a types Hothorn and forest A, sources Zeileis rain illustrations, A-L, Tropical Boulesteix (1993) C, MJA Strobl Werger AM, Guyana. Polak in VG, area Jetteer watershed H, Steege ter P Turk performance. JA, Drought. 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Data may be preliminary. ei iegnewti otnosppltos h aeo h etoia reEeu act (aubl.). falcata Eperua ge- tree microgeographic neotropical of the drivers of adaptive case and The ONE Neutral populations: (2015) continuous I within Scotti divergence C, netic Scotti-Saintagne M, Foll L, Brousseau (Fabaceae). Aubl. falcata Eperua tree Amazonian E Dreyer PVA, intrapopu- Fine trees. L, promotes forest Brousseau variability rain environmental tropical local from Highly example (2013) An I 1169–1179. traits: quantitative Scotti of J, divergence Cigna lation D, (Fraxinus Bonal ash L, in Brousseau variation adaptive Forests. of Random scale (2001) the L Identifying Breiman nursery. local? the is from local results How L.): (2005) excelsior J Stewart tolerance drought D, constrains Boshier adaptation Local tree. 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Promote V Fine Paul Herbivores ): (2004) s PD ( Coley Author Forests I, forest. waterlogged rain Mesones and tropical slopes PV, a on in Fine rates productivity treefall and Higher biomass (2010) stand V lower Freycon in L, result Blanc soils JD, Bontemps F, Morneau experiments. B, Ferry transplant a long-term in in pinaceae) herbivory Botany rigida, leaf of (Pinus of pines GA rates pitch Fox normal-stature determines DR, Taub species W, Seedling Fang (2006) SE forest. Hartley rain SG, Malaysian Postgermination Compton Germination, MP, (2010) Eichhorn CG Willis K, Ranges. Kovach L, Ecological 293–319. Burghardt Species R, and Casas Adaptation, de Rubio K, Donohue eerhRpr eis8 eateto ttsisadMteais UWe,2004. Wien, framework. WU inference Mathematics, conditional and Statistics A Statistics Graphical partitioning: of and recursive Department tational Unbiased 8, (2006) Series models. A Report parametric Zeileis Research K, general Hornik in T, inference Hothorn Simultaneous (2008) P Westfall F, Journal Bretz T, Hothorn species. tree forest L rain Poorter B, Bachelot H´erault B, edig ffiepoeacso iu aaini:ptnilt eetvraini drought-tolerance. in variation detect to in potential deficit canariensis: water seasonally Pinus to of 23 tropical response provenances in morphological five and distribution of Physiological seedlings species (2009) tree L Gil explain Rodr´ıguez-CalcerradaL´opez J, tolerance R, flood Does habitats? (2003) Means. TA flooded Least-Squares Kursar aka Means, OR, Marginal Lopez Estimated emmeans: (2018) V. R Lenth The Means: Least-Squares (2016) 1–33. V. by R assessed Lenth populations. gradients fir elevational silver along in adaptation adaptation local and , of diversity lack Local-scale experiments: transplant (2017) reciprocal C robur). Pichot Q. AC, and Latreille petraea (Quercus oaks white European among L´eger P, V Abadie T, Lang forests. Amazonian contrasting PVA across Fine assemblages adaptation. H´erault B, local GPA, in issues Lamarre Conceptual (2004) D Ebert TJ, Kawecki 74 509–519. , 77. , , 50 , 93 346–363. , 1125–1133. , ln Speciation Plant Oecologia ora fToia Ecology Tropical of Journal ora fEcology of Journal nRCmaint ple ersin(ogeeBook) (Google Regression Applied to Companion R An tal. et clg Letters Ecology , Ecology 136 tal. et 21)Hg-ult Nsfo ei ein ihih nrgeso patterns introgression highlight regions genic from SNPs High-quality (2018) , oubaUiest rs,NwYr,N,USA. 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Weber Fagus AR, in Pluess scale regional the at drive C climate interactions Heiri to belowground A, trees. and Frank tropical Above- AR, of (2011) Pluess JW species Dalling cryptic A-H, two between Eom EA, segregation Herre habitat SA, forest. of Mangan dry C, differentiation tropical Pizano physiological a in and habitats Morphological wet (2011) 1547. and C dry Tree. between Tinoco-Ojanguren a seedlings H, Being Paz of Consequences Pineda-Garc´ıa F, Evolutionary Some Systematics (2006) and A Evolution, Hampe RJ, analysis. Petit correlation canonical through investigated Guiana French hydrolo- in and Ecology example occurrences An species constraints: tree between soil relationships gical Within-plot (2002) D Sabatier species. S, P´elissier fungal Dray R, non-sporulating a from bis-xanthone, novel a JP xanthonol, Polishook AW, pre- Dombrowski Three JG, Ondeyka E: globulifera. and Symphonia D of C, bark Globulixanthones root the (2002) from B , of properties Bodo antimicrobial disadvantage ZT, with Fomum a xanthones M, trees. nylated Meyer suggests densiflora) P, (Pinus Mkounga trial AE, pine transplant Nkengfack red reciprocal Japanese of A growth (2015) and Genomes survival & A in Kanazashi transfer seed K, northward Shimada T, Nagamitsu meta-analysis. A performance: lifetime. plant AA on per Agrawal individual RA, square Hufbauer per per WF, seeds not Morris more but produce Creek year, species per Salt Small-seeded canopy (2004) of M of Westoby metre DS, zone Falster MR, hybrid Leishman AT, Asteraceae) Moles tridentata: fitness. (Artemisia of sagebrush tracking multiple-year big 213–225. of mountain importance and the Canyon: Nothofagus basin tree the the WS of of Moore responses ED, expansion. Mcarthur plastic upward KJ, and rear-edge Miglia adaptive predict edge: experiment transplant the long-term on a Living to (2016) pumilio AC Premoli P, Mathiasen croisement. th´eorie du la S´eries de III quintessence Naturelles La (1917) JP Lotsy 61 181–187. , , 162 , 40 , ge 21)Erylf tgscnrbt togyt oa dpaini rbdpi thaliana. Arabidopsis in adaptation local to strongly contribute stages life Early (2016) J Agren ˚ 11 143–156. , 321–331. , 813. , , 3 rbdpi thaliana Arabidopsis 351–353. , ge 21)Se omnyccigadmraiydffrbtentolocally two between differ mortality and cycling dormancy locally Seed two (2016) between J differ Agren ˚ mortality and cycling dormancy Seed (2015) J Agren ˚ , 37 tal. et 187–214. , 21)Gnm-niomn soito td ugsslcladaptation local suggests study association Genome-environment (2016) tal. et tal. et Ecology 20)Nn-errcpoa rnpateprmn ntegardens the in experiment transplant reciprocal Nine-year (2005) . naso Botany of Annals 20)Drc n neatv ffcso nme n mutualists and enemies of effects interactive and Direct (2007) tal. et naso Botany of Annals , , 88 e Phytologist New 113 20)Ioainadisciia/nhlitcatvt of activity insecticidal/anthelmintic and Isolation (2006) 20 1021–1029. , 7590–7595. , ilgclJunlo h ina Society Linnean the of Journal Biological rhvsNelnassdsSine xce et Exactes Sciences des Neerlandaises Archives ora fEcology Of Journal , , 117 Ecology ln,Cl n Environment and Cell Plant, LSONE PLoS 117 , 210 249–256. , ora fAntibiotics of Journal mcv171. , 589–601. , , 92 , 47–56. , 7 nulRve fEcology, of Review Annual e33636. , Oecologia , 92 384–396. , Phytochemistry , 181 , reGenetics Tree 59 , 34 607–619. , 288–292. , 1536– , , Plant 86 , Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. privileged. Br T, adaptation. Wezeman local to contribute that processes 8 the Mediterranean evaluate BA to in Gould methods Survival DB, Early Lowry Ed,). and SM, Delzon, Wadgymar Emergence (S Aiton) Seedling pinaster in Role (Pinus Microenvironment The Pine and Gonz´alez-Mart´ınezMaritime (2014) Al´ıa Origin R, SC Population MA, quanti- Zavala of evolutionary B, for Revuelta-Eugercios methods models. Vizca´ıno-Palomar General N, mixed (2016) generalized M from Morrissey inference S, genetic Nakagawa tative H, Schielzeth P, Villemereuil Species. de Eleven Mediterranean in Trends in Interspecific versus Drought Responses with Individual Tree. Seedlings: Associated Tree a Traits of Ecophysiological S´anchez-G´omez Energy (2006) F, and Valladares D Death, Life, an (1975) of Van populations L Valen Neotropical and African species. in tree structure tropical genetic ancient spatial fine-scale K affect adaptation Heer crohabitat KB, Budde P, Torroba-Balmori perspective. plant A introgression: Adaptive (2018) QCB random , Cronk for C, measure Lexer variable-importance A, conditional Suarez-Gonzalez new, package. A party on! the Party measures: in (2009) available importance A variable forests Zeileis a forest T, in Hothorn random C, factors in Strobl Bias soil (2007) solution. and T a types Hothorn and forest A, sources Zeileis rain illustrations, A-L, Tropical Boulesteix (1993) C, MJA Strobl Werger AM, Guyana. Polak in VG, area Jetteer watershed H, Steege ter P Turk performance. JA, Drought. 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Core tree R tropical of survival Sciences habitat Biological on of B: neighbourhood scale-dependence Society Taxonomic biotic (2009) and R Valencia partitioning NC, niche Garwood DFRP, Burslem SA, Queenborough 14 738–749. , . p. ihrsett rsn n uueciai conditions. climatic future and present to respect with spp.) aua rdc Reports Product Natural rnsi clg n Evolution and Ecology in Trends ln n Soil and Plant s ,MsesK 21)Xnhn ies opudfml hc sbt omnand common both is which family compound A dimers: Xanthone (2015) KS Masters S, ¨ ase :Alnug n niomn o ttsia computing. statistical for environment and language A R: h mrcnStatistician American The ora fVgtto Science Vegetation of Journal , LSONE PLoS 352 , 276 tal. et tal. et 243–251. , , 4197–4205. , M bioinformatics BMC 32 tal. et Journal R tal. et 21)Sgaue flclaatto ncniaegnso as( oaks of genes candidate in adaptation local of Signatures (2016) 6–28. , tal. et 21)Si-eitdlclaatto lesseln uvvland survival seedling alters adaptation local Soil-mediated (2012) , 12 tal. et 19)TeIflec fSi oe raiaino h floristic the on Organization Cover Soil Of Influence The (1997) 1–23. , 21)Ietfigtresadaet fslcin innovative selection: of agents and targets Identifying (2017) , 21)Attdnlgains igorpi itr n mi- and history biogeographic gradients, Altitudinal (2017) 29 , 20)Euiaigterl fgntcditadnatural and drift genetic of role the Elucidating (2009) 1 165–176. , , 21 14–17. , 34 ln Ecology Plant LSONE PLoS Biotropica Genetics 216. , , , 4 8 705-716. , 25. , oeua Ecology Molecular , , , 204 7 9 , 259. , oeua Ecology Molecular e109132. , 131 ehd nEooyadEvolution and Ecology in Methods 1281–1294. , 81–108. , onainfrStatistical for Foundation R ln Biology Plant Biotropica rceig fteRoyal the of Proceedings , 18 3803–3815. , , 25 ilg Letters Biology , 5907–5924. , 39 , 8 683–690. , 688–697. , , Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. evs vrtecus fte6yas( years 6 ( the Height of 2014. course in the alive over still leaves) ( and leaves 2009 in of germinated number seedlings of herbivory and 3 Table ( experiment the tabulated. of end the at ( overall flooded seasonally ( garden indicated: also is garden ( Regina stations, ( temperatures maximum and and 2 Table ( experiment the of end the at ( overall germinated ( had collected seeds of of number type the as the well and as ) west), and east (e.g. indicated site 1 Table ponderosa. Pinus Tables: in soils serpentine to adaptation Local (2007) JW Wickham. Wright Hadley by ggplot2 (2009) H Wickham lv Y6 Alive longitude and : eircltaslnaineprmna adnifrain The information: garden experimental transplantation Reciprocal : umr ttsis( statistics Summary : Ni Symphonia latitude r lotabulated. also are ) ,tenme fsesgriae ttetm ftasln ( transplant of time the at germinated seeds of number the ), the , east habitat TNL and niiulifrain the information: individual lnainregion plantation longitude n rcuo( Iracoubo and ) ,adaeaehrioy(eryaeaeo ecnaeo aae rao l its all of area damaged of percentage of average (yearly herbivory average and ), Ni h ohrte a on:saoal odd( flooded seasonally found: was tree mother the Tmax min ,tenme fsesgriae ttetm ftasln ( transplant of time the at germinated seeds of number the ), hr h ohrte a on.Togeneral Two found. was tree mother the where imum, Go n average and ) Herb west ,adtenme fselnsaiea er6( 6 year at alive seedlings of number the and ), , mean habitat ecotype ave SF ,aeidctd(ee rne.Tehbtttp feach of type habitat The France). 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Data may be preliminary. lne ntesm aiasadrgo sterpoeac.Dse ie ersn rnpat between transplants lines represent Dotdashed lines habitats. within Dashed habitats. regions provenance. and between regions their transplants across as represent transplants lines region represent in Dotted difference and found regions. habitats morphologies edaphic within different same habitat the ( the the represent SF represent in trees and curves planted cartoon HT sigmoid Laussat on the black details rainfall: with The across R´egina). The associated Average SF; pluviometry and experiment. and of (Laussat HT transplantation map between sites Greyscale reciprocal plantation b) the two America; of South the Brousseau of of from north R´egina (reproduced location (2500mm/year), (3500mm/year) the and along Guiana Guiana French French of Location a) 1: Figure Figures: egahcllcto,puimty n eino eircltaslnaineprmna sites. experimental transplantation reciprocal of design and pluviometry, location, Geographical ..S sp1 S. i.e. nH and HT in 23 .globulifera S. tal et nS) oi ie ersn juveniles represent lines solid SF); in ,21 ihpriso) )Design c) permission). with 2015 ., Symphonia Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. iue2: Figure nlsso emnto ucs of success germination of Analysis Symphonia 24 ed.a ieseti h hdhuebefore shadehouse the in spent time a) seeds. Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. codn otevralssuidcvrae finterest: of covariates studied =West), variables the to according 3: Figure either ( east to either refer be can can Ecotypes plantation plantation. and ( habitat, provenance, ecotype, by by transplantation before by house gardens the to transplantation sp1 ;hbttcnb ihrHltp( Hilltop either be can habitat ); Plant lsicto reo emnto n uvvlof survival and germination of tree Classification Region lnigrgo ( region planting = E ecotype rws ( west or ) ecotype HT and rSaoal odd( flooded Seasonally or ) E W and =East; provenance ). provenance W 25 =West), Prov )Oealpoaiiyo emnto success germination of probability Overall c) . Symphonia )Poaiiyo emnto nteshade the in germination of Probability b) . Region Habitat SF ;Rgo fpoeac ( provenance of Region ); poeac ein( region =provenance ed n ueie fe years 5 after juveniles and seeds lnighbtt( habitat planting = S.globulifera ( glo E SF Prov =East; =Season- or ) )and .) S.sp1 W Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. .Idvda epne griain uvvl n otlt)wr aeoie as: categorised were experiment, mortality) the and of survival, end (germination, responses Individual ). nyidvdasaiei 04wr nlddfrmaue tae5 nyidvdaswt tlattoannual two least at with individuals Only 5. analyses. age included. RGR were at field) for measures or for included (shadowhouse included were 2009 were in measures 2014 germinated in individuals alive Only individuals experiment. Only the of course the over TNL ( tree mother the of ecotype =East; interest. 4: figure flooded; nally g 5, age ttlnme fleaves, of number =total RGR W Prov lsicto reaaye fjvnl hntpctat codn otesuidcvrae of covariates studied the to according traits phenotypic juvenile of analyses tree Classification =West), H Region HT rltv rwhrt nH; in rate growth =relative =hilltops), Habitat D1-D5 poeac ein( region =provenance glo Ecotype RGR eda er1t er5respectively, 5 year to 1 year at dead , lnighbtt( habitat planting = = S.globulifera TNL ctp ftemte re( tree mother the of ecotype = rltv rwhrt nTNL; in rate growth =relative D= E ; sp1 =East; imtea g , 5 age at diametre SF 26 = S.sp1 W Saonlyflooded; =Seasonnally =West), .Maue hntpctraits: phenotypic Measured ). 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Plantation log Y the and herbivory, on and are traits growth of ( leaves of 5: Figure rwn nter‘oe niomn eaiet h oprsnaedntdb rage rusgrowing Groups triangle. a by denoted are comparison the to relative environment ‘home’ their in growing ( flooded seasonally es qaemasyal nlsso rwhtat Hih ( (Height traits growth of analysis yearly means square Least TNL cdghklop.Ecotypes: (c,d,g,h,k,l,o,p). ) n ebvr ( herbivory and )), SF .Poeac:es ( east Provenance: ). Herb .globulifera S. of ) E n et( west and ) Symphonia ctp Habitat x Ecotype 27 ( glo W and ) seedlings. .Patto:es ( east Plantation: ). .sp1 S. Age ( abefijmn and (a,b,e,f,i,j,m,n) sp1 H ,Daee ( Diameter ), nyasi nteXae.Log axes. X the on in years in .Hbtt:hltp ( hilltops Habitats: ). E n et( west and ) D ,Ttlnumber Total ), rvnnex Provenance W HT .Groups ). and ) Posted on Authorea 16 Apr 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158705417.70708833 — This a preprint and has not been peer reviewed. Data may be preliminary. n‘wy niomnsrltv otecmaio r eoe ycrls infiac:nnsgicn ( non-significant Significance: circles. by denoted are comparison ( the 0.05 ), to relative environments ‘away’ in * ,00 ( 0.01 ), ** ,001( 0.001 ), *** ). 28 NS