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aia eo Duarte Melo Marina Authors FORESTS ENHANCES DIVERSITY TREE HIGH EDITOR: XIAOJUAN LIU This article isprotected rights by All copyright. reserved c which may lead to not been AcceptedThis article undergone has for buthas and accepted been publication review fullpeer [email protected] 4 Maynooth University, 3 13418 2 (ESALQ/USP)of SãoPaulo 1 H.; Bonat,W. J.L.;Potvin, C.; Stape, ArticleAuthors Brancalion Zuim Ferreira IA, M EoSl, F Eco&Sols, UMR CIRAD, A ite this articleite this as Department of Biological Sciences, “Luiz de Queiroz” College of Agriculture, University Agriculture, of College Queiroz” de “Luiz Sciences, Biological of Department Department of Department des A ddress: - 260.
(full names)(full through thecopyediting, andproofreadingprocess, typesetting, pagination names for citations for names 6
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2, 7 , Catherine Potvin Catherine , Mathematics and Statistics and Mathematics doi:
differences between this version anddifferences the version between this 1,2* Maynooth, Ireland. E Maynooth, Ireland. :
10.1111/1365 Rfe d Adae Moral Andrade de Rafael ,
. E : - Duarte, M. M. M. Duarte, mail address:mail [email protected] 8 , Wagner Hugo Bonat Hugo Wagner , - - 49 Mnplir Fac. E France. Montpellier, 34398 Brancalion, P.Brancalion, H.S. 2745.1366 -
, Maynooth University Maynooth , mail address:mail IH ITRETO I TROPICAL IN INTERCEPTION LIGHT ; Moral, R. A.; A.; R. Moral, ; ,
ã Dimas. São 9
3 Jans Guillemot Joannès ,
9 , José Luiz Stape Luiz José ,
iaiaa So Paulo São Piracicaba, Version of Record Version of Guillemot . Address: Logic House Logic Address: .
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al address: mail Zuim 10 aoie Isaac Caroline , Pedro H. S. H. Pedro , . Please , C. I. F.; I. C. , Brazil, ,
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Paulo(ESALQ/USP),São Brazil. Piracicaba, 6 Montpellier, France 5 This article isprotected rights by All copyright. reserved Dimas. Piracicaba,Dias, São Paulo,Brazil. 11. São 13418 Postal Code: Acceptedd Universidade Biológicas. Ciências de Departamento Florestal. Restauração e Ecologia de Laboratório author * Corresponding 11 [email protected] Paraíso do Altos 3780, Universitária, 10 19081. address postal caixa 100, Santos, dos Heráclito 9 Article Montréal, S3/27, 8 7
E Department of Forest of Department eatet f ilg, cil University McGill Biology, of Department eatet f Statistics of Department E c&os Ui Mnplir CRD IR, R, otele, uAr, F SupAgro, Montpellier, IRD, INRA, CIRAD, Montpellier, Univ Eco&Sols, Department of Forest Science, São Paulo State University (UNESP) University State Paulo São Science, Forest of Department - - mail address:mail [email protected] mail address:mail [email protected] :
[email protected] e São Paulo. Escola Superior de Agricultura “Luiz de Queiroz”. Av. Pádua Av. Queiroz”. de “Luiz Agricultura de Superior Escola Paulo. São e Canada, Canada, 1B1.H3A .
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Sciences, “Luiz de Queiroz” College of Agriculture, University of University Agriculture, of College Queiroz” de “Luiz Sciences, Marina MeloDuarte Fdrl nvriy f Paraná of University Federal ,
E - mail Btct, Brazil Botucatu, ,
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Address: Address: : [email protected] uiia Bai, 81531 Brazil, Curitiba, 25 v Dcer efed room Penfield, Docteur ave 1205
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Article type: Running title: E 19 +55 Telephone: 982324088 This article isprotected rights by All copyright. reserved Accepted Article - mail address:mail [email protected] Research article Research
Diversity enhances ligh enhances Diversity
t interception
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1. This article isprotected rights by All copyright. reserved Accepted cost designing time. and space light of Synthesis distribution oflightinterception. a promoted richness Tree season. dry the during pronounced more were and seasonally changed relationships despite 5 site, 20 from and Article3. Anhembi in high meters 5 to (up transects vertical and grids 2. Restoration Ecology. Ecological Complementarity; T BEF Keywords: mixtures). 18 to (monocultures in time and Here, forests. tropical by mitigation change climate enhance to importance foremost of is production
- species mixtures showed showed mixtures species
Understanding the Understanding Light interception increased from monocultu from increased interception Light We : intercepted intercepted ), inplots containing different richness combinations ofspecies. and levels High tree diversity (i.e., over a over (i.e., diversity tree High substantial differences differences substantial
assessed -
- to 114 to efficient large efficient
we investigated the effects of tree diversity on light interception over space over interception light on diversity tree of effects the investigated we
Di two versity by the canopy in canopy the by
- species necpe photosyntheti intercepted es variable less -
rpcl tree tropical species mixtures), and in Brazil Brazil in and mixtures), species er; idvriy n Eoytm ucinn Theory; Functioning Ecosystem and Biodiversity heory; - light interception relationship interception light processes -
scale tropical forest tropical scale restoration programs. mixtures in mixtures
iPAR
rcse; csse srie; ih partitioning; Light services; Ecosystem processes; eprl n sail ie bt hrzna a horizontal both (i.e. spatial and temporal observed
restored xeiet, oae i Panama in located experiments, that underlie the effects of tree diversity on primary on diversity tree of effects the underlie that
oprbe to comparable hundred species in species hundred ABSTRACT
Anhembi tropical forests through more throughforests tropical atv rdain ( radiation active c mns monocultures. amongst
(during the dry season) dry the (during
Sardinilla res to 5 to res the s –
should be critically consideredshould critically for be
Anhembi
mixture) maximizes the amount the maximizes mixture) best performing performing best -
species mixture species and up to 4 meters high in high meters 4 to up and
site (20 site i A) vr horizontal over PAR)
. – - At the At
even Diversity to 114 to monocultures, Sardinilla s in s d vertical) nd
capture in capture
Sardinilla Sardinilla
- species - iPAR
site
107 106 105 104 103 102 101 100 95 94 93 92 91 90 89 88 87 86 85 84 83 82 99 98 97 96
séis or a necpaã d lz eo osl e lrsa toias e dois em monocultur tropicais, florestas de Panamá no dossel localizados arbórea, diversidade pelo de experimentos luz de interceptação a sobre espécies mitigação é primária produção a sobre 1. This article isprotected rights by All copyright. reserved Palavras Accepted planejamento custo interc e biodiversidade entre relação em tropicais tempo. florestas no e espaço no de homogênea mais luz dosselde captura uma de pelo meio por restauração, interceptada luz de quantidade a maximizar Síntese forma homogênea mais pronunciada mais resultados nos diferença monoculturas grande da apesar luz, interceptar em de mistura a Sardinilla 3. deespécies. combinações em altura Articlede em altura de m 5 até (de verticais transectos de e horizontal espaço 2. 114 espécies).
: A alta diversidade arbórea arbórea diversidade alta A : Compreender os mecanismos que mecanismos os Compreender A interceptação de luz aumentou de monoculturas a misturas de misturas a monoculturas de aumentou luz de interceptação A do longo ao (iPAR) interceptada ativa fotossinteticamente radiação a Avaliamos -
chave: chave: e de mudanças climáticas. Neste trabalho, investigamos os efeitos do número de número do efeitos os investigamos trabalho, Neste climáticas. mudanças de entre as a misturas de 18 espécies) e no Brasil Brasil no e espécies) 18 de misturas a as cinco
qi estudadas aqui Anhembi
misturas de 20 a 114 espécies em espécies 114 a 20 de misturas Complementaridade; Ecologia de restauração; Processos ecológicos; ecológicos; Processos restauração; de Ecologia Complementaridade;
espécies mostrou iPAR equivalente à equivalente iPAR mostrou espécies - a saã seca. estação na eficiente de programas de restauração florestal em larga florestaleficiente programaslarga derestauração de em escala. temporal ( espacialmente e ns acls otno o dfrne nvi d rqea e riqueza de níveis diferentes com contando parcelas nas )
fundamental para maximizar o papel de florestas tropicais na tropicais florestas de papel o maximizar para fundamental A .
relaç ( mesmo superior a superior mesmo
iuz abra rmvu necpaã d lz de luz de interceptação promoveu arbórea Riqueza paã d luz de eptação ão
RESUMO nr dvriae iA vari iPAR e diversidade entre fundamentam
tanto verticalmente). horizontal quanto Anhembi
–
100 espécies em misturas) é capaz de capaz é misturas) em espécies 100
ee e levada ser deve área área s
os efeitos da diversidade diversidade da efeitos os das monoculturas mais eficientes eficientes mais monoculturas das
(n Anhembi a estação seca). Em Em seca). estação a –
área área
(com misturas de 20 a a 20 de misturas (com Sardinilla
Sardinilla peetds pelas apresentados m ot pr um para conta em ou cinco
nr estações entre e
espécies em espécies d (contendo e Sardinilla
até 4 m 4 até arbórea A
, ,
110 109 108
Funcionamento de Ecossistemas. de Funcionamento luz; de Partição ecossistêmicos; Serviços This article isprotected rights by All copyright. reserved Accepted Article
era E; era e idvriae e Biodiversidade de Teoria BEF; Teoria
119 118 117 116 115 114 113 112 111 135 134 133 132 131 130 129 128 127 126 125 124 123 122 121 120
2017; Ratcliffe 2017; forests to years grasslands on out carried historically ( functioning resource enhance can diversity species Introduction This article isprotected rights by All copyright. reserved Acceptedwas primary terrestrial, to aquatic Ramírez eco and richness et al. restoration forest in sequestration carbon maximizing for key is 2019) &Brancalion, (Chazdon al. et change climatemitigate to solution central re forest tropical because important particularly al. forests, tropical Consequently Article usual richness species tropical of diversity high the to compare not does that dozen) a than less (usually species tree of number limited a on based commonly 2018) , , 2013; Sapijanskas, 2013; , 2019) , on average 2.38 times higher than the average biomass of monocultures, and that half that and monocultures, of biomass average the than higher times 2.38 average on
producers ( producers B Current evidence suggests a positive but saturating relationship saturating but positive a suggests evidence Current t al. et remain , oiest ad csse fntoig BF ter peit ta increased that predicts theory (BEF) functioning ecosystem and iodiversity , as well to provide multiple provide to well as ,
the , oeu t l, 2001 al., et Loreau 2017) , et al. et
among Pti & oel, 08 Gamfeldt 2008; Gotelli, & (Potvin logical s oe f ih re pce rcns i te ucinn of functioning the in richness species tree high of role , 2017) , plant
I a meta a In . poorly adnl e a. (2011) al. et Cardinale y on i toia rsoain plantings restoration tropical in found ly
et al. the most species most the
s processes
and algae and , mainly in temperate biome temperate in mainly , , 2014) understood . Understanding the effects of tree diversity on forestgrowth diversity tree the effectsof Understanding . - analysis considering various types of ecosystems, from from ecosystems, of types various considering analysis . . xeiet crooaig hs hoy ae been have theory this corroborating Experiments ).
) (Cardinale (Cardinale , in mixtures in , (Balvanera (Balvanera
ecosystem se ecosystem - capture
and biomass and (Clarke (Clarke (Brancalion
on ta maximum that found
t al. et -
biomes biomes growth has been promoted worldwide as a as worldwide promoted been has growth tcmuiy ee ad otr ecosystem foster and level community at
containing increasing numbers of species of numbersincreasing containing et al. et t al. et 21; Clarke 2011; , rvices that underpin that rvices t al. et - , 2006) , s rich et al. et . Moreover, forest BEF studies are studies BEF forest Moreover, . 2017) , (Paquette
2013 ,
, 2019b; Lewis 2019b; , ecosystems on Earth Earth on ecosystems , and extended in the last 15 last the in extended and , Ti kolde a is gap knowledge This . (Potvin (Potvin Guerrero ; (Brancalion t al. et t al. et
biomass 2018) , et al. et 21; Guerrero 2017; ,
human wellbeing human et al. et between species between - , 2011; Hulvey 2011; , Ramírez Ramírez
hyperdiverse t al. et , 2019; Pugh 2019; , , civd by achieved
o t the to nor (Barlow 2018) , t al. et et - , . ,
144 143 142 141 140 139 138 137 136 161 160 159 158 157 156 155 154 153 152 151 150 149 148 147 146 145
al. (Meyer compositions species varying scales spatial large considering when function ecosystem species. 1.35 with reached be would yield this of This article isprotected rights by All copyright. reserved forests Accepted2019a) grass invasive of control the as such restoration, sequestration and forests diverse of functioning the understand to key therefore is interception light on diversity structure canopy forestin tropical acute particularly is interception light on effect diversity of evidence empirical of lack This Maire space and time across quantified directly been seldom have interception light forest on richness species tree of effects the Sapijanskas 2014; (Pretzsch, space canopy filling Guillemot Articletree in relationships BEF of al. et compared mixtures of productivity increased (i.e., overyielding productivity forest , 2017) , 2016; Schnabel 2016; , Understanding . al. et
et al. et s hn f ra iprac fr einn cost designing for importance great of then is . Forest structural complexity structural Forest
, 2013; Forrester & Albrecht, 2014; Sapijanskas 2014; Albrecht, & Forrester 2013; , t al. et , 2018) , how how . Moreover, light interception is linked to a number of key objectives of forest forest of objectives key of number a to linked is interception light Moreover, . 2020) , s, which typically display very high very typicallys, display tree which complex diversitymulti and (Laurans idvriy loss biodiversity
s and services and (Hardiman
is, however, still an unresolved issue unresolved an still however, is, et al
ih leaves with .
how eet tde showed studies Recent . ,2019) et al. et -
species mixtures species et al. et re diversity tree Siatv Vled 2005) Vellend, & (Srivastava 21; Trogisch 2014; , . This suggests that light Thissuggeststhat . al. et (Isbell , 2014; Juc 2014; , (Binkley wih iey eut i ehne lgt interception light enhanced in results likely which ,
can affect affect can 21; Gough 2011; ,
Siatv Vled 20; Cardinale 2005; Vellend, & (Srivastava has been has et al. et et al. et , 2011) , affect ker ker (Morin (Morin , 1 ,
vital et al. et
shown to be an important determinant of determinant important an be to shown ht ies forests diverse that s et al. et 992; Forrester 992;
, and long ,and (Rodrigues (Rodrigues atrs of patterns Levels of Levels
, 2015; Williams 2015; , clgcl processes ecological et al. et t al. et 2017) , - related interactions are key drivers key are interactions related , 2011; Forrester & Bauhus, 2016;Bauhus, Forrester& 2011; , 2019) , - (Cardinale (Cardinale
fetv large effective mlil eoytm functions ecosystem multiple , - et al. et term resilience term
. diversity required to saturate to required diversity et al. et Studying ih absorption light to monocultures to , 2014; Sercu 2014; , et al. et ad a prl explain partly may and , , 2009; Brancalion Brancalion 2009; , et al. et et al. et , 2012, 2018, 2019; le 2019; 2018, 2012, , r mr efcet in efficient more are
the effects of tree of effects the , 2017) , sc as such , - , 2012) , cl restoration scale (Brockerhoff et al. et et al. et . However, . ; , n tropical in ( especially Dănescu - 2011) , , 2017) , layered carbon et al. et et , , .
170 169 168 167 166 165 164 163 162 185 184 183 182 181 180 179 178 177 176 175 174 173 172 171
( the 4) canopy; the in light intercepted of variation vertical the 3) light intercepted of variation horizontal the 2) light; intercepted of tree eco the in expand to expected are which programs, This article isprotected rights by All copyright. reserved Acceptedexperi old natural, of diversity the on based was plots of level richness different BEF contains for It studies. forests.The tropical by interception light on diversity tree of influence the test to approaches methodological M and Material forestspecies), allowedlinking which BEF restoration. theorytotropical established species) 18 to (up mixtures and monocultures ArticleSardinilla investigation our mixtures) and monocultures in trees of volumes crown compared also we monocultures constituent their and species between system restoration (2021 diversity gradients diversity mental areas mental Study s Study
Here, Here, We included in this study two study this in included We , combinations of up to 18 species 18 to up of combinations n within and Panama in Anhembi in ites we ethods
based on two on based
explored explored (TreeDivNet, 2017) (TreeDivNet, Sardinilla
(Scherer years
in two tropical forests tropical two in
, Brazil with unprecedented high tree species richness (up to 114 to (up richness species tree high unprecedented with Brazil ) , in two d , intwo 5 te diffe the 5) ; - 2030). patterns of light interception through time and space across space and time through interception light of patterns - site, in Panama, site,in Lorenzen experiments: a 15 a experiments:
istinct plantations, istinct . The . sites that contained contained that sites rences t al. et temporal
(for this purpose, to support our conclusions, our support to purpose, this (for (Scherer , . .
consistsof Anhembi and a a and 2005) , S in in pecifically - year light interception between mixtures of of mixtures between interception light nine variation in canopy light interception light canopy in variation - current Lorenzen wih rsne is pce in species its presented which ,
-
site, in Brazil, in site, old five species in monocultures and in monocultures in and five speciesin
- an experiment an year
,
BEF experiment established in established experiment BEF experiments based on different different on based experiments we - ntd ain dcd on decade Nations United old quantified: 1) total amount total 1) quantified: - et al. et growth forests nearby the nearby forests growth
tree diversityexperimenttree , 2005) , holds
especially designedespecially
. We We . . Its maximum Its . an experiment an carried out carried
194 193 192 191 190 189 188 187 186 211 210 209 208 207 206 205 204 203 202 201 200 199 198 197 196 195
monsoon) climate climate monsoon) Forest Atlantic Brazilian diversity species functions, and processes ecosystem on level diversity tree unprecedented n does It functioning. forest tropical on levels diversity tree high very of effects assess to designed This article isprotected rights by All copyright. reserved 2003 in established was (HD) plantation” Accepted combination comb three species, two of combinations three monocultures, distinct five of replicates two thus: were plots effective died individuals all almost because f other 2009) Latin a in mixture), same the of replicates (six species different containing fast a plots of combinations (six three of combinations and species) each for replicates ( dimensions (Bignoniaceae A.DC. ex Bertero (Euphorbiaceae) excelsum Anacardium (Malvaceae) Article surveys field of ca hereafter both In times. different at started Methods . ot include species in monocultures, but its design allows for testing the effects of an an of effects the testing for allows design its but monocultures, in species include ot All All ive,
The S1 six
lled “main plantation”, established in 2001 in established plantation”, “main lled
semi for detailed information on this site) this on information detailed for 45 x 45 m 45 x 45
and species duringthedryseason shed leaves (Supporting Sardinilla ) slow ;
peetd i species six presented , -
deciduous. s on i high in found as oda alliodora Cordia
(Köppen Breo x ut) kes (Anacardiaceae) Skeels Kunth) ex (Bertero -
, 45 x 48 m or 48 x 48 m 48 x 48 or m 48 x 45 , - growing: experiment was established in Panama, Panama, in established was experiment rwn, slow a growing, ; Brancalion Brancalion nomto, Figure information, ntos f he seis ad i rpiae o a five a of replicates six and species, three of inations n o te pce ( species the of One - Geiger; Geiger;
Cedrela )
after plantation after - of them of – Ri & a. Oe (Boraginaceae) Oken Pav.) & (Ruiz iest rsoain projects restoration diversity et al.
n 24 in Kottek
- – odorata , 2018) rwn ad n nemdae pce) and species) intermediate an and growing
fast , spacing between seedlings was 3 x 3 m. The m. 3 x 3 was seedlings between spacing , patto ws 3 er old years 13 was (plantation xeietl lt of plots experimental -
growing: growing: . t al. et L. (Meliaceae) (Meliaceae) L.
S1) ) . , containing monocultures (12 plots: two plots: (12 monocultures containing , It consists It
establishment Te eefe cle “ih diversity “high called hereafter The . C
.
06 see 2006; (plantation was 15 years 15 was (plantation – alliodora -
square design ( design square C. uha seemannii Luehea
odorata of ws o cniee here considered not was ) and and two native tree plantations tree native two
ne a A (equatorial Am an under
(Kunert uprig information, supporting
a lat 0 pce, for species, 80 least (at being deciduousbeing the and
aeua rosea Tabebuia and prxmtl simil approximately Potvin & Dutilleul & Potvin
ua crepitans Hura raa Planch & Triana using using at onset of field field of onset at et al. et intermediate: ; , 2012) , old similar tree similar
(Bertol.) at onset at - species
f six of . Our .
L. ar
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subtropical, oceanic, hot summer) climate (Köeppen; (Köeppen; climate summer) hot oceanic, subtropical, ofeachtwo replicates group groups) called (hereafter combinations species different four contained 28 a within them of all ones, larger of subsets were pools species Smaller species. 18 and 9 6, level: richness each surveys This article isprotected rights by All copyright. reserved survey field our in phenology AcceptedLorenzi deciduous were 45.4% and evergreen were them of 19.8% them of 23.3% other deciduous(thewere 58.7% evergreen and were 2016 in individualsliving all of 18.0% them, Among species. 114 the of 94% for phenology s m 1.5 others inthe of repeated distribution spatial S1 (A replicates four with design randomized (see species tree native 114 and 58 20, yearsold wasnine (plantation restoration forest conservation biodiversity both Article for hotspot global leading region biogeographical threatened most leaves shed species of 50% seasonal by covered once
and
varied between evergreen and deciduous) and evergreen between varied ) S2 , in 24 18 24 in , pacing pacing et al. et The ee vrre ad 52.4% and evergreen were
and and , 2003) , Anhembi Methods S Methods -
8 idvdas e plot). per individuals 480 x 18 m plots organized in eight blocks, each one containing one plot of plot one containing one each blocks, eight in organized plots m 18 x
(Brancalion (Brancalion f pce ws admy eemnd o te is rpiae n then and replicate first the for determined randomly was species of .
Nevertheless,
experiment was established in southeastern Brazil, under Cfa (humidCfa under Brazil, southeasternin established was experiment - species pool species the same same the 2).
semi
(Ruiz The smaller species pools were subsets of the larger ones. larger the of subsets were pools species smaller The (Gandolfi at onset of fi of onset at s - ti ifrain a etrl tkn rm h literature. the from taken entirely was information this , t al. et deciduous forest deciduous - treatment. Seedlings were planted in 45 x 48 m plots (3 x mplots x48 in45 (3 treatment. planted Seedlings were Jaen &PotvinJaen 2011 it is important to stress that, stress to important is it
2019b) ,
( supporting information, supporting ee decidu were et al. et supporting information, supporting s
- eld sampling eld D) of each (see each of D) f h Alni Forest Atlantic the of , 2009) , Te xeiet a etbihd n 2006 in established was experiment The . e ee be o id nomto o leaf on information find to able were We . Within the 58 the Within . (Morellato & Haddad, 2000) Haddad, & (Morellato u. ihn h 114 the Within ous. . This forest type is part of part is type forest This . ) Alvares Alvares . )
using different tree richness levels:tree richness using different ihn h 20 the within
supporting information, information, supporting Luac, 2009) (Laurance,
Lrni 19, 98 2009; 1998, 1992, (Lorenzi, et al. et Table S1 Table Table S2 Table - species treatment, 15.3% treatment, species as we we as (Ribeiro
2013) ) - - did . The eight blocks eight The . pce treatment, species treatment, species tu tee were there thus , ), in a completely a in ), . This region was region This .
t al. et not assess leaf leaf assess not
n tropical and , where 30 where , one of the of one 2009) , Figures The a , -
245 244 243 242 241 240 239 238 262 261 260 259 258 257 256 255 254 253 252 251 250 249 248 247 246
study area. Thus,conditions. edaphic and climatic to according site, to site from vary may traits phenology leaf species, same A This article isprotected rights by All copyright. reserved Acceptedpart ofthe site this from Data R:FR light, diffuse characterized take to equipment different effic more measurements required which site, each of conditions climate specific age same et al. relationship the Nevertheless, b iPAR. from predicted commonly is it level, plantation tree the at aPAR measuring directly in difficulties practical to Due leaves. of surface abaxial (Weiss absorbed not was and canopy it through and canopy a reaches that PAR total Article between difference the simply is iPAR photosynthesis, for used be potentially can and leaf a enters actually that PAR the of fraction the is aPAR While (aPAR). PAR absorbed for proxy a as (iPAR, radiation etween them is variable, especially between early and late stages of vegetation of stages late and early between especially variable, is them etween og eiuu species, deciduous mong , 2000) et al. et Light i Light ih itreto ws hrceie uig necpe pooyteial active photosynthetically intercepted using characterized was interception Light t the At , iPAR and aPAR were expected to vary congruently. congruently. vary to expected were aPAR and iPAR ,
(Nouvellon . As tables 2004 , nterception assessmentnterception using all theall plots Sardinilla . However, Nouvellon )
measurements are a good proxy for proxy good a are measurements were converted from R:FR into iPAR for illustration, in figures in illustration, for iPAR into R:FR from converted were – the
nor t nor et al. et ai o red:far of ratio
iently this information would be more accurate if it were specific to theto itwereaccurateif specific morebe would informationthis site, , 2000) ,
e ih elce y h soil the by reflected light he compared inour compared durati et al. et all
–
analyses carried were (R:FR). outusing data raw lvld ke Pws U) K 10 esr a ue to used was sensor 110 SKR UK) (Powis, Skye leveled a and accurately and
but reflected by leaves and non and leaves by reflected but 2000) . It considers neither the amount of light tha light of amount the neither considers It . n n pretg o la ls my ay Ee fr the for Even vary. may loss leaf of percentage and on
at the at - e lgt spectra light red
experiment Anhembi , a t the t h aon o ti rdain ht passes that radiation this of amount the
site Sardinilla Sardinilla s
iPAR (Capers & Chazdon 2004) Chazdon & (Capers iPAR were closed were hc ol e bobdb the by absorbed be could which . (R:FR
At this site, we measured iPAR, measured we site, this At On the other hand, hand, other the On ) site, - ised f iPAR of instead , green parts of the canopy the of parts green
light interception was interception light - canopy forests t reached the reached t
(Nouvellon Under .
due to due and and of theof in .
271 270 269 268 267 266 265 264 263 288 287 286 285 284 283 282 281 280 279 278 277 276 275 274 273 272
10.5061/dryad.5mkkwh75q R:FR as plot, the of size grids m 6 x 6 by obtained plot, enco per points sample 49 at R:FR measured We cosine meas This article isprotected rights by All copyright. reserved deciduousness canopy to due level lowest its at be to expected Accepted were Measurements to equidistant grid m 6 x 3 a by determined regularly plot, per points 98 at ground, above meter one iPAR (see iPAR calculate and field open outside and inside PAR measure to other, each with calibrated USA), WA, (Pullman “ 6.76 = iPAR)] iP into R:FR of conversion and The 2016). (July August, season rainy the during PM, 2:15 to AM 9:50 from and ground above m 1.7 at interception light influence equally trees neighbor nearest four the to equidistant Articleillustration collection data for S4 Figure supporting see “ (hereafter the near points additional and plot the of part Sardinilla Sardinilla , which encompassed the entire plots (leaving out only two only out (leaving plots entire the encompassed which , mpassed the entire plot entire the mpassed ure R:FR in periods with ca. 100% diffuse incoming light. light. incoming diffuse 100% ca. with periods in R:FR ure ” - ,
corrected corrected At the At D iPAR next to next Sardinilla ataset - nomto, Fig information, – h fu nearest four the Anhembi R:FR regression”,R:FR DatasetS3 light
.5 RF (e spotn information, supporting (see R:FR * 6.85 the plantation, plantation, the - border HD taken S1 - collecting tip, in order to receive ra receive to order in tip, collecting , available in the Dryad Digital Repository, DOI DOI Repository, Digital Dryad the in available ,
supporting information’s Methods information’s supporting : horizontal : site, we we site,
s) at the peak of the dry season (August 2015), when iPAR was iPAR when 2015), (August season dry the of peak the at ) s
and 17 points per plot per points 17 and
in the in (leaving out only one or two rows of trees, depending on the on depending trees, of rows two or one only out (leaving ure used two leveled Decagon AccuPAR LP AccuPAR Decagon leveled two used without
planted main plantation plantation main
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8 sunny 2:2 and AM 10:00 between place 10.5061/dryad.5mkkwh75q levels (March season wet the horizontal season This article isprotected rights by All copyright. reserved Accepted Sardinilla 10.5061/dryad.5mkkwh75q iPAR horiz specific times ofthe year, thus measurements should colle ground above m 1 and Articleusing forest point sample 0 cm and and cm 0 ction time ction na R:FR ontal
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s h ohr etmtr usd the outside ceptometer other the as Dtst S, 2 S ad S5 and S4 S2, S1, Datasets , using the R:FR sensor (at sensor R:FR the using , (for all sections of this work) this of sections all (for , FigureS (dry and wet(dry and seasons) “ Anhemb
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10.5061/dryad.5mkkwh75q 2016) August and (July season rainy the during m, 5 to 1 of heights the from meter, every transects, vertical infor trees neighbor nearest four the to equidistant and plots the within plantation HD the of plot each in transects vertical four and plantation, This article isprotected rights by All copyright. reserved Accepteddata the of plantation HD the at m 0.20) (± 16.10 and plantation main the at m 0.08) (± 13.60 were trees living ceptometer. the to compared sensor, the on measurements ground take to difficult very it made which weeds, invasive of amount the of plots many in understory the 1) because site study each 10.5061/dr 2016 (February 10.5061/dryad.5mkkwh75q “ hereafter m 4 to 0 from transects, vertical the over meter S8) and S7 Fig. information, (supporting trees neighbor nearest Article four the to equidistant skyovercast 10.5061/dryad.5mkkwh75q DOI
were ain Fig mation,
level At the At Measurements were carried out at different heights over the vertical transects at at transects vertical the over heights different at out carried were Measurements the At
taken from the yad.5mkkwh75q .
n 2 i ws osbe o ec hge heights higher reach to possible was it 2) and Anhembi hratr “ hereafter , Anhembi Sardinilla ure Sardinilla , at the main (hereafter “ (hereafter main the at ,
S6 dy esn etcl iPAR vertical season dry : ).
site, we we site, botanical botanical site, we established six vertical transects in each plot of the main the of plot each in transects vertical six established we site,
sn a adr w ladder, a Using experiment and 8.20 (± 0.06) m at the at m 0.06) (± 8.20 and experiment ) ) “ (hereafter HD and ) , always from, always to2:00PM AM 10:00 season wet the during repeated were measurements The . Anhembi ) plantations ) established 12 systematically 12 established inventories carried
As a reference, reference, a As wt esn etcl iPAR vertical season wet : . Sardinilla lo using Also , always from 9:30 AM to 2:40 PM 2:40 to AM 9:30 from always , cnutd :R esrmns vr the over measurements R:FR conducted e
planted Sardinilla drn the during , - main: vertical main: mean heights (an heights mean
out in2016). lde, e esrd PR every iPAR measured we ladder, a i ec po o the of plot each in , -
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.
r sao (uut 2015; (August season dry
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Anhembi D
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site S7 ,
344 343 342 341 340 339 338 337 336 359 358 357 356 355 354 353 352 351 350 349 348 347 346 345
2016, February 2017 and January 2018 (hereafter “ (hereafter 2018 January and 2017 February 2016, whereas 2017, August February and 2015, February 2013, January 2012, January 2016 in were season wet in Junemeasurements 2015, August 2011, July in place took season 2 to 2011 from dry) and (wet seasons different transects vertical the as same (the plot This article isprotected rights by All copyright. reserved Accepted “ hereafter tr and crown between (difference depth crown determine to Sweden) (Långsele, two. by divided diameters four those of mean the from radius diagonals diameters: different four w (DME), measurer distance Sweden) (Långsele, Haglöf a Using plots. neighbor from influences minimize to center, plot the to closest five each in species per individuals two and monoculture per individuals five chose We combinations. distinct in species same the of differences growth plantation’s year. seasons, a months and within 10.5061/dryad.5mkkwh75q Article ground above meter 2018 March to 2017 April from year, 10.5061/dryad.5mkkwh75q DOI Lighttemporal variation Crown volume Crown assessment At the At At the At , using a compass a using Sardinilla species, both in monocultures and in the five the in and monocultures in both species, Sardinilla Anhembi Anhembi - an crown” main:
site, we measured iPAR one meter above ground at 12 points per points 12 at ground above meter one iPAR measured we site, site, in July 2016, we measured crown volume of each of themainof each of volume measuredcrown we 2016, inJuly site,
( hereafter hereafter the
) ( W itne t assess to intended We . see large
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upporting information, Fig. S9 Fig. information, upporting ).
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st e lo ok otl iA measurement iPAR monthly took also We D , at the same the at , “
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in Anhembi
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- species mixtures to account for account to mixtures species : seasonal iPAR”, D iPAR”, seasonal : per iPAR iPAR iPAR
- plot ). We calculated the crown the calculated We ). pce mxue te ones the mixture, species We used a Haglöf vertex Haglöf a used We ” ,
varied varied mentioned above mentioned D ataset ewe years, between
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versus mixture. The mixture. versus th at them compare to but volumes, crown of measurement that data overestima overlook we species, all radius) cylinder a as volume crown calculated This article isprotected rights by All copyright. reserved reasonable a was function link identity with model normal a bounded, not and continuous Acceptedt mean the by induced variation the separate covariates. with dispersion, and mean the of modeling simultaneous HD (McGLM) models 10.5061/dryad.5mkkwh75q DOI horizontal we richness, of levels different between varied plots within for for iPAR to converted were but units, original Sardinilla Articleshape for crowns. a my e cone fr ih ersos Bcue h dt fo the from data the Because regressors. with for accounted be may hat more intuitive comprehension and more comprehension and intuitive plantations 2 Lightinterception Dataanalyses * π * crown depth). crown * π *
t the At W were and ted R:FR e analyzed whether levels of richness enhanced light interception, for both the both for interception, light enhanced richness of levels whether analyzed e
the not available in this study. this in available not realis more Nevertheless, volumes. their ” and the “ the and ” Tee oes r ueu i ti cs, eas te alw o the for allow they because case, this in useful are models These . Sardinilla
Anhembi
(Bonat & Jørgensen, 2016; Bonat, 2018) Bonat, 2016; Jørgensen, & (Bonat refore,
ed
site
xeiet. For experiments. potential potential analyses were no were analyses
Sardinilla We We
bt mi n H plantations) HD and main (both acknowledge ) and and - variation HD: horizontal horizontal HD: easy comparisoneasy sites. among (Sapijanskas itd utvrae cov multivariate fitted O ur purpose here was not to provide an accurate an provide to not was here purpose ur - variance relationship from the extra the from relationship variance
t biased by the choice of choice the by biased t
the s that, by assuming a simple cylinder form for for form cylinder simple a assuming by that, results and and results
in crown shapes among species and and species among shapes crown in se
nlss :R aa eekp n their in kept were data R:FR analyses, et al. et R:FR tic models of crown shape crown of models tic , 2014) , graphical representation graphical
” datasets ” e species level, in monoculture in level, species e , separately for the main and main the for separately , ,
rac gnrlzd linear generalized ariance sd h “ the used (crown volume = (crown = volume (crown to analyze if mean R:FR mean if analyze to
(Datasets S1 and S2 and S1 (Datasets assuming
t s osbe to possible is It Sardinilla Sardinilla
- a cylinder a variability ,
to allow to ie are site require - main: likely ,
392 391 390 389 388 387 386 385 384 409 408 407 406 405 404 403 402 401 400 399 398 397 396 395 394 393
the density of individuals is a factor that can influence light interception. interception. light influence can that factor a is individuals of density the and plots some in mortality high was there since covariate, a as plot each within survival for dispersionparameters. It and w boththe mean plot the of grids horizontal of coordinates and survival of percentage richness, mean constant a with coupled assumption, This article isprotected rights by All copyright. reserved p dispersion Acceptedmean non the assumptions, moment second si model, mean a assumed and predictor, linear mean the for link logit a used we 1, and 0 between bounded is iPAR since However, 10.5061/dryad.5m “ “ datasets the using seasons, wet and dry measurementsF:RF iPAR) (andalso spatially beconsidered cannot influence likely variable a is interception light since covariates, include to important species). this from were individuals planted of 1/5 where mixtures, species 2 in lower ArticleFi information’s interception light lower present mor high no was intention our covariate, Anhembi - aine eainhp wees h extra the whereas relationship, variance At the At nce we are modeling continuous proportions. Hence, by making these first and first these making by Hence, proportions. continuous modeling are we nce iPAR” horizontal season wet : ta - ) lity, especially the ones the especially lity, species , as well as the interaction between coordinates x and y and x coordinates between interaction the as well as , rmtr wih depend which arameter, gure S10 gure Anhembi ( orthogonal orthogonal
kkwh75q mixtures, where 1/3 of planted individuals were were individuals planted of 1/3 where mixtures, in our models our in
: site, we also fit also we site,
it is possible to possible is it , aid ih ihes (20 richness with varied ), coordinates x and y and x coordinates - variance relationship analogous to the one for the binomial binomial the for one the to analogous relationship variance t pnlz, n u aaye, lt that plots analyses, our in penalize, to t
u t te isn individuals missing the to due that h codntsx n o ape points sample of y and x coordinates the ed
- gaussianity of the data data the of gaussianity contained ted
n h cvrae effects covariate the on - variance assumption. We included the effects ofeffects included the variance We assumption. note
McGLMs to analyze if iPAR means ( means iPAR if analyze to McGLMs Anhemb
that , in meters, in , - as important to include the percentage of thepercentage important toinclude as variability variability Cordia alliodora Cordia – , at the main plantation, main the at , : r sao hrzna iA” and iPAR” horizontal season dry i: each sample point sample each
- aaes 4 n S5 and S4 Datasets 58 ,
- relative to the to relative was
n 114 and was
d , which were expected to expected were which , pes cek supporting check (please
, oee va n extra an via modeled independent. y egbrod thus neighborhood, by
in the linear predictors linear the in e atd o study to wanted we accounted for by the the by for accounted C. alliodora, C.
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left corner corner left
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418 417 416 415 414 413 412 411 410 432 431 430 429 428 427 426 425 424 423 422 421 420 419
salsmn, at establishment, mortality High S10). Figure the at mortality higher presented levels Sardinilla included y and T This article isprotected rights by All copyright. reserved Acceptedtransects in variation light horizontal this work higher to corresponds parameter McGLMs, S5 and S4 and dry season horizontaliPAR” “ datasets: of dispersion and means analyze to section previous Article the95%confidenceobtain for intervals thetrue parameters. adequately diversity tree of level reatment
rm the from Vertical light Vertical variation Horizontallightvariation
o the For both At and McGLMs fitted the in effects of significance the assesstests to Wald used We in the linear predictors for the mean and dispersion parameters. dispersion and mean the for predictors linear the in measured) xeiet w ue pretg o sria a a oait, ic hg richness high since covariate, a as survival of percentage used we experiment,
Sardinilla DI 10.5061/dryad.5mkkwh75q DOI , adapted tothe conditions.area’s are are replicate poy f pta htrgniy n ih dsrbto ( hge dispersion higher (a distribution light in heterogeneity spatial of proxy a
top the Sardinilla Sardinilla Sardinilla - , we also used McGLMs to determine whether means and dispersion of of dispersion and means whether determine to McGLMs used also we , an hrzna RF” “ R:FR”, horizontal main: - fu replicates) (four
et onr f h plot the of corner left Anhembi Anhembi
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and and sites, we used the same model same the used we sites, : wet season horizontaliPAR” initial initial Anhembi o al species all not
i meters in , light variation variation light ) Sardinilla . at high richness levels levels richness high at :R r PR dispersion iPAR or F:FR td design study xeiet spotn information, (supporting experiment :R or R:FR - ad ecnae f uvvl were survival of percentage and ) D hrzna RF” “ R:FR”, horizontal HD: ih (:R or (F:FR light
across vial fr plantation for available included iPAR)
rhgnl distances orthogonal etcl pc ( space vertical tu representing thus ,
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described in the in described aft As done for the for done As S2, S1, Datasets aaees in parameters, er plantation plantation er i different in , Anhemb vertical
were x i:
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vertical space, at different richness levels) richness different at space, vertical heigh covariate) transect vertical dis and mean R:FR” vertical varied R:FR This article isprotected rights by All copyright. reserved q different including models, regression AcceptedS11 Dataset iPAR”, monthly random as year within replicate and richness species season, of effects 10.5061/dryad.5mkkwh75q DOI dispersion identity transect measurement species), 114 and 58, (20, level S9 and Anhemb Articlemeasurement measurement 9 (6, level R:FR” t
, , DOI 10.5061/dryad.5mkkwh75q DOI ,
f :R measurement R:FR of i: dry season vertical iPAR” and “ and iPAR” vertical season dry i: Temporal D For iPAR in different seasons and years and seasons different in iPAR For a variation iPAR vertical the For , height , ataset and and parameters. between , interaction between richness level and height and level richness between interaction , .
eso wr rcns lvl 1 2 3 n 5 pce) pretg o survival, of percentage species), 5 and 3 2, (1, level richness were persion ,
18 species), percentage of survival, of percentage species), 18
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ataset variationinterception inlight of R:FR measurement R:FR of identity , DOI 10.5061/dryad.5mkkwh75q DOI , richness levels and heights and levels richness
S6 , DOI 10.5061/dryad.5mkkwh75q DOI , ec vria tasc maue ws identified was measured transect vertical (each , DOI , r t ass i RF peetd itnt eair oe the over behaviors distinct presented R:FR if assess (to eplicate (four replicates of each treatment), each of replicates (four eplicate
) w fte mxd ea ersin oes icuig the including models, regression beta mixed fitted we , effects
10.5061/dryad.5mkkwh75q uadratic predictors over time per species richness species per time over predictors uadratic
. ) (1 to 5 m) and interaction between richness and and richness between interaction and m) 5 to (1 . For the HD the For . , Anhembi hi itrcin s ie efcs n ya and year and effects fixed as interaction their t the t
o mnhy PR vr n ya (“Anhembi: year one over iPAR monthly For we fit we Anhembi . For the main main the For .
ted
: wet season vertical iPAR” vertical season wet : ( vertical transect vertical “
Anhembi McGLMs including including McGLMs ) , effects accounted for were richness were for accounted effects , plantation site
: seasonal : of iPAR measurement iPAR of (in both dry and wet seasons, wet and dry both (in ) , the predictors of both the both of predictors the , plantation ), we also fitted mixed beta mixed fitted also we ),
identity (“ Sardinilla
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(“ , height height Sardinilla ,
- ,
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ataset s spatial as atasets f R:FR of
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S10
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465 464 463 462 461 460 459 458 478 477 476 475 474 473 472 471 470 469 468 467 466
10.5061/dryad.5mkkwh75q Sardinilla ofthe fixed significance effects using likelihood random as year within replicate of effects the and level This article isprotected rights by All copyright. reserved Accepted 2018 environment R the in out carried were analyses All variables. independent as level richness plot and species tree and variable dependent as volume 10.5061/dryad.5mkkwh75q 5 and iv) mixtures; species fivespeciesoftheother Between each mixture and treatments. three or two Amongst iii) species; three or two of combinations Be i) R:FR mean compare to contrasts specific established We level. confidence global the for correction Bonferroni with comparisons, multiple for level confidence 95% a at test Tukey used ArticleWe covariate. assumption. reasonable a was model normal a bounded, not and continuous was data the 1 Since species). and five of species combination three of combinations 3 species, two of combinations 3 monocultures, ) .
Species inmonocultures Species xmi ’s main main ’s - We carried out an analysis of variance to compare mean R:FR within within R:FR mean compare to variance of analysis an out carried We We also compared crown volume between the five the between volume crown compared also We species
plantation mixtures ) ) we mnclue; i Bten ah ooutr ad their and monoculture each Between ii) monocultures; tween
sn a two a using ewe dfeet pce cmoiin (2 ees 5 levels: (12 compositions species different between
(“ Sardinilla (“ Sardinilla
xtures We used percentage of survival within each plot as a as plot each within survival of percentage used We - an hrzna R:FR” horizontal main: -
- a AOA wt log with ANOVA, way an crown” main: - ratio (LR) tests nested models. for .
In main species main ,
oh ae, e sesd the assessed we cases, both
D ataset ,
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in monocultures in (R Core Team Core (R S12
S1 DOI ,
plot ,
ecies) ecies) DOI s
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486 485 484 483 482 481 480 479 501 500 499 498 497 496 495 494 493 492 491 490 489 488 487
plantation Results This article isprotected rights by All copyright. reserved increased) AcceptedS2 and “ forfit errors the estimated parameters using theMcGLMs of for ( other each richness tree with ( season (χ (χ Article( monocultures than light more intercepted significantly mixtures all and other, each from 2 than interception light higher P P P < 2 2 < HD the At <0.0001). = =1.54 0.0001; 0.05) (see 0.05) 108.07 ; , DOI 10.5061/dryad.5mkkwh75q DOI ,
Light Horizontal light variationlightHorizontal Sardinilla
iue 1 Figure , d.f.=2, , d.f.=2, re pce rcns ehne iA a bt sts A the At sites. both at iPAR enhanced richness species Tree Horizontal light light Horizontal supporting See the At ma iPAR mean , from monocultures to mixtures to monoculturesfrom , d.f.=2, d.f.=2, , Figure 1 Figure P
>0.05). >0.05). interception supporting information supporting - Anhembi B main: horizontal R:FR” and “ andR:FR” horizontal main: P (χ ). = A, 0.46). 2 In the the In P =
< supporting information’s supporting 57.91 .01 o te r ad χ and dry the for 0.0001
plantation
increased site, increasing levels of richness enhanced iPAR in both seasons both in iPAR enhanced richness of levels increasing site, information variation
r sao al ramns hwd infcnl dfeet iPAR different significantly showed treatments all season dry , d.f.=2, , -
or 3 or
, light interception was interception light , significantly significantly (assessed by the dispersion parameter of the M the of parameter dispersion the by (assessed P - , Figure ,
< species ( 0.0001), but 0.0001), al S4 Table )
at the at erae (thus decreased
mixture S1 Sardinilla Table S3). Five S3). Table ih pce rcns (χ richness species with 1 Sardinilla site Sardinilla fr siae ad soitd standard associated and estimates for ) ) 2 . In the wet season, iPAR also increased also iPAR season, wet the In . = 58 and 114 species did not differ from differ not did species 114 and 58 57.91
plots - HD: horizontal R:FR”horizontal HD:
not influenced by richness levels richness by influenced not
R:FR or iPAR or R:FR d.f.=2, ,
( evenness in iPAR distribution iPAR in evenness P ’s <
main - species 0.05) P plantation , which did not differ differ not did which , < Sardinilla 0.0001 for the wet wet the for 0.0001 mixtures .
2 =310.34 ,
D (χ site’s presented atasets 2 =258.41 d.f.=3, , cGLMs
main main
S1 ,
510 509 508 507 506 505 504 503 502 526 525 524 523 522 521 520 519 518 517 516 515 514 513 512 511
Thus spec 18 and 9 for lower was dispersion light horizontal horizontal ( mixtures 3 for intermediate monocultures, for higher d.f.=2, d.f.=3, This article isprotected rights by All copyright. reserved space filling Acceptedcanopy the of layers different over PAR of amounts higher interception) season wet the parameter mean the from independent variation, of source extra an represents which parameter, dispersion diversity) higher at space horizontal the over interception acc richness higher with variance their lower the 50%, from are data further the 50%, of proportion a at obtained variance maximum with data), proportion inter PAR Article( season rainy the in mixtures season dry the during levels richness all between the over horizontaldistribution space. of light interception , (Bonat & Jørgensen, 2016) Jørgensen, & (Bonat in Vertical lightvariationVertical P P < < At both At the At
P eto. ic w asmd qartc mean quadratic a assumed we Since ception. did not vary according to richness level ( level richness to according vary not did h H patto, high plantation, HD the 0.0001) and and 0.0001) 0.05 space in mixtures mixtures in space <0.05). <0.05).
for to20species 58species, (χ compared when with with Fgr 2 Figure ; Anhembi it did so, with lower dispersion values values dispersion lower with so, did it Sardinilla vegetation That means that means That from 6 from
site, horizontal variance in iPAR decreased with richness, differingrichness, with decreased iPAR in variance horizontal site, ). utd o lwr PR oiotl variance horizontal iPAR lower for ounted
n h main the In and and A the At . Figures 2, Figures - hn oprd to compared when
species to richer mixtures mixtures richer to species Anhembi . At the At .
light interception was more evenly distributed over the over distributed evenly more was interception light er Sardinilla
S12 plantation sites, higher richness levels were able able were levels richness higher sites, Anhembi diversity (McCullagh & Nelder, 1989) Nelder, (McCullagh& and and -
pce and species site’s main main site’s
S13 site, during the dry season, the dispersion the season, dry the during site, hrzna dseso ws significantly was dispersion horizontal ,
was χ , monocultures.
), following the same pattern of mean mean of pattern same the following ), 2= and and 0.9364, d.f.=2, P=0.63) d.f.=2, 0.9364, also - ies than for 6 species ( species 6 for than ies . Moreover, we also modeled the modeled also we Moreover, . for (typical relationship variance
l bt the but all n h HD the in ( thus
2
= , thus , plantation soitd with associated 2 for lower 6.7155, higher higher n h HD the In being
d.f.=2, d.f.=2, 58 plantation iA a different at iPAR ,
. Therefore, stands .
evenness - (more
more efficient in efficient more - o 114 to
n 5 and P =0.03). =0.03). , but during but , to intercept to more plantation even
P (χ
- <0.050
- f light of 2 species species =8.08
even light
). , ,
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10.5061/dryad.5mkkwh75q of monocultures. to compared canopy, the of layers (χ ( species (χ mo from differing significantly richness, with increased heights This article isprotected rights by All copyright. reserved interact Accepted(χ species 114 and 58 containing plots duringthedryseasoncanopy, evenly more light intercept could levels diversity 10.5061/dryad.5mkkwh75q the for fit McGLM iPAR of dispersion decreasing showed richness Increasing height and species of number the ( level confidence 95% a at level richness every (χ richness with increased significantly richness (χ either levels vertica HD: P Article(χ richness plantation more light (χ mixtures to = 2 2 =4.1200 =77.2180 0.1585) the o bten egt n rcns lvl (χ level richness and height between ion Figure 3 Figure During the wet season, in season, wet the During the At cL ft for fit McGLM I te HD the In . , 2
d. , = and its dispersion its and , d.f.=3, , R:FR” l evenly 5.5849, d.f.=2, 5.5849, f.=3, 2 =68.1179 ). Effects of interaction between richness and height were not significant significant not were height and richness between interaction of Effects ). Anhembi ,
, n ifrn lyr o vegetation, of layers different in P
P = D < 2 .5, hc mas ht itrs anand ihr PR n all in iPAR higher maintained mixtures that means which 0.25), =1.4887 ataset
0.0001) but, between mixtures, increasing only from three to five to three from only increasing mixtures, between but, 0.0001) plantation , d.f.=3, d.f.=3, ,
Anhemb ; )
ie i te r sao, PR vr h vria transects vertical the over iPAR season, dry the in site,
P over
χ . the = (dispersion parameter of the McGLM fit for the “ the for fit McGLM the of parameter (dispersion S7 2
= 0.0613) or by its interaction with height (χ height with interaction its by or 0.0613) , d.f.=2, , d.f.=2,
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2 = f measurements of 39.6361, d.f.=2,P<0.000139.6361, 2 P = = 725, d.f.=2, 97.2058, 0.4750). , iPAR over the vertical transects was higher in higher was transects vertical the over iPAR , ) - , an vria R:FR” vertical main:
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in different layers of vegetation of layers different in 2 Dispersion effects Dispersion = P
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Dtst S6 Dataset , (dispersion parameter parameter (dispersion i nt ay among vary not did , ifrn between differing ), ataset 2 = 3.6841, d.f.=2, 3.6841, ( )
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the 1 ( level richness the for fit McGLM the of parameter to according differ significantly not did effects Dispersion This article isprotected rights by All copyright. reserved Acceptedfor except monocultures, best their to more species some that indicates This mixtures. species two as well as interception, plantation, main ( 114 study, the of beginning 114 general, plots thatricher meaning ( time over behavior quadratic a and P=0.0154), d.f.=2, (LR=12.9574, level richness and time between Articleaverage mixtures the P<0.001). d.f.=2, (LR=17.2, dynamics seasonal iPAR Figure4 0.5061/dryad.5mkkwh75q
difference between wet and dry seasons for 20 for seasons dry and wet between difference Anhembi PAR
Temporal variation Temporal ininterceptionlight Species inmonoculturesSpecies mixturesand ( . In general, general, In . Figure4 ) There was a significant interaction effect of season and richness level on iPAR at iPAR on level richness and season of effect interaction significant a was There Analyzing different compositions within each richness level, level, richness each within compositions different Analyzing year, one over monthly iPAR Analyzing .
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Discussion (F 5) ( mixtures other to and monocultures to compared values, iPAR highest the have to This article isprotected rights by All copyright. reserved Anhembi Accepted not but species, Jucker 2007; Bagchi, affect can designs result for aBEF experiment. unprecedented an is season, dry which inthe interception, light tosaturate able was still not 2005) Vellend, Therefore, c choice roles. species and design different experimental play can they which in assemblage, community a in others ( processes responsi ecosystem were species dozen a than less instance, ( levels richness low at place take to reported commonly increases diversity species as saturate functions ecosystem in interception light Article levels. species) 114 to (58 high 2014) dynamics forest and production . 1,110
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) highlight that one must be careful when concluding about about concluding when careful be must one that highlight ) We ther We (Canham (Canham . Our results (iPAR at at (iPAR results Our . forests tree diversity, both at low (one to five species) and very and species) five to (one low at both diversity, tree
efore report substantial effects substantial report efore negig restoration undergoing an influence results of experiments of results influence an et al. et -
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were the the were between differences the when season, rainy the during place took site diversity functional reflect necessarily not (Bruelheide does diversity taxonomic thus and traits differentpresentfunctional may species setsof DistinctBEF. saturation in species of levels This article isprotected rights by All copyright. reserved Accepted in Guillemot than mixtures in higher was branches 2016) (Guillemot likely richness (Sapijanskas canopy the throughout spaces empty occupy Forrester light of use outputs traits Articleal. al. less 2015) , 2009) , horizontal variation of iPAR. The duration and intensity of deciduousnes of intensity and duration The iPAR. of variation horizontal . In fact, at the at fact, In . (Ruiz cneune f bte ocpto o te aoy y re rnhs n leaves and branches tree by canopy the of occupation better a of consequence a . In addition, these traits can show plasticity in plasticity show can traits these addition, In . Tree diversityeffects Tree on the h Tree diversityeffects Tree on the v were associated with more with associated were , crown shapes and arrange andshapes ,crown ihr re ihes also richness tree Higher to related is levels richness higher at interception light increased that report We lowest got speed growth , t al. et et al. - t al. et (Ruiz an Pti, 2011) Potvin, & Jaen et al. et t al. et 2018) , , 2020) a 2020) , , 2014) , t the - 2014) , Jaen & Potvin, 2011; For 2011; Potvin, & Jaen Sardinilla . Tes f itnt pce my neses ter rws n better and crowns their intersperse may species distinct of Trees . Anhembi Anhembi
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ments spatially spatially loe te aoy o necp lgt more light intercept to canopy the allowed erticalvariation orizontalvariation (Pretzsch, 2014; Sapijanskas 2014; (Pretzsch, rester & Albrecht, 2014; Sapijanskas 2014; Albrecht, & rester even
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shown that biomass allocation to allocation biomass that shown in lightinterception in n cuily eemn mixture determine crucially and
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, 2014; Forrester & Bauhus, 2016) Bauhus, & Forrester 2014; , ’s main main ’s (Wright (Wright single
et al. et s ely cue f ihr ih itreto o nt A the At not. or interception light higher of cause a really is
, 2014; Jucker 2014; , . According to According . lower at higher richness levels. This is This levels. richness higher at lower year all all plantation t al. et measured utpe ca multiple emporalvariationinterception inlight Siatv & eln, 2005) Vellend, & (Srivastava . . S When different species are combined, the diversity the combined, are species different When pecies 2009) , n individual in contain a , ’ more
ih iest plots diversity high et al. et Peooia dfeecs mn species, among differences Phenological . contributions to contributions egt. oevr vria dseso of dispersion vertical Moreover, heights. Sapijanskas et al. (2014), however, however, (2014), al. et Sapijanskas ies show that individuals can receive more receive can individuals that show ies oy layers nopy , 2015) , eiuu neighbors deciduous even , and thus, and interception, light
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pce ae epnil fr ihr csse fu ecosystem higher for responsible are species in pathways patterns phenology This article isprotected rights by All copyright. reserved more holding of advantage the Nevertheless, performers. good are monocultures Acceptedsome as This comparabl decrease level richness same the of compositions different between dispersion the with while increases richness, interception light mean that states which theory, BEF with well agreed the levels richness same the of mixtures species processes ecosystem influenced by suggested which shift to linked be reduction competitive of consequence a probably monocultures, in than volumes crown larger developed mixtures interception light to Article weighting measured easily no is there of amount the out single cannot we effects complementarity and selection separate to possible not performances species single from expected a Sardinilla Sardinilla specific may lead to ask whether high diversity is necessary to maintain ecosystem functions, ecosystem maintain to necessary is diversity high whether ask to lead may suggest Toward s ln seis diversity species Plant : by e to the best monocultures best the to e (Wright (Wright ecosystem function site effect selection s
Sapijanskas s that complementarity effect complementarity that
. a mechanistic understandingamechanistic lightinterceptionindiverse of forests
At s
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this site this (Grossiord biomass 2009) , (Forrester & Bauhus, & (Forrester t al. et s , average iPAR across species richness levels (see levels richness species across iPAR average , ,
where
; allocation and and allocation et al. (Naeem, 2016) (Naeem,
or . 2014) , can t al. et T by e five he , 2014)
light light a single species is responsible for playing a major role major a playing for responsible is species single a , as shown by other BEF works BEF other by shown as , coefficient coefficient complementarity effect complementarity influence 2013) , . intercepted by each by intercepted n diin to addition In .
s - (Loreau & He & (Loreau
pce mixtures species occur in mixture in occur performed differently in intercepting light intercepting in differently performed . , since different monocultures and different and monocultures different since ,
branching
2016) oee, e vdne that evidenced we However, to represent species represent to nctioning ecos . This crown plasticity was shown to shown was plasticity crown This . ystem functioning functioning ystem
pattern ctor, 2001) ctor, , species oprn t wa wud be would what to comparing s s , species of the mixtures the of species , at least to some extent (as extent some to least at ,
showed where o n light interception, since interception, light n
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species that it though even monocultures, the of any by outperformed ones c Cardinale 2009; multifunctionality keep to ability their is species This article isprotected rights by All copyright. reserved Accepted forest approaches restoration sequestration. overyielding biomass (Brancalion (Rodrigues projects, grass exotic invasion control help can light, more intercepting by diversity, High consequences. process this in role important plays composition species that and interception, light and occupation canopy regarding 2016) Bauhus, & space over distribution overall Article light interception light increases which enhancestime, and vertically) and (horizontally Diversity levels. diversity high very at even nann hge dvriy il iey rsn bte oeal ucinn ta ls diverse less than functioning overall better present likely will diversity higher ontaining (Aerts & Honnay, 2011) Honnay, & (Aerts Conclusion increase interception light increases, diversity when that, conclude We u to due t al. et had contrastinghad abilities
t al. et species High tree richness levels should be critically treerichnessHigh should levels critically considered for tropical be designing 2009) ,
et al. et . We gathered evidence that mixtures have advantages over monocultures over advantages have mixtures that evidence gathered We . 2012) , the s : implications tropical: for forestrestoration , a a ,
, 2012) , (Sapijanskas diinl costs additional ,
I as pooe cnp pooytei, hc cn eut in result can which photosynthesis, canopy promotes also It . ao concern major which can can which
and functional redundancy functional and . In fact, in this study, the five the study, this in fact, In . to foster
(Naeem, 2016) (Naeem, et al. et
in inte jeopardiz , 2014; Guillemot 2014; , carbon carbon soitd with associated
o mn large many for rcepting rcepting light. e . sequestration sequestration
Our findings may lead to various practical various to lead may findings Our their (Hector & Bagchi, 2007; Wright Wright 2007; Bagchi, & (Hector overall overall herbicide
(Hooper et al. et - scale tropical forest restoration restoration forest tropical scale
- in a changing climate. changing in a species mixture was in no case no in was mixture species iaca vaiiy n success and viability financial , 2020) , contained
application application et al. et ,
and enhanced carbon enhanced and , 2005) , i te ae plot, same the in , . Ecosystems . n weeding and (Forrester s
s well, as t al. et
by ,
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Paulo’s Experimental Station of Forest Sciences Sciences Forest of Station Experimental Paulo’s collection for ( Laboratoty and design experiment Acknowledgements This article isprotected rights by All copyright. reserved Accepted DOI Repository, Digital Dryad the inhttps://doi.org/10.5061/dryad.5mkkwh75q included were manuscript this in discussed statement: availability Data improvements tothis work. acknowledge supportWe from theIN 2013/50718 grant# (FAPESP, Foundation Personnel Education Higher ( Development p his diligent Article Mancilla Lady and Monteza José Mendes Teixeira Carlos João especially feasible, experiments our made staff. and facilities their We are grateful to Robin Chazdon and E and Chazdon Robin to grateful are We e ial tak dtr ad nnmu rvees ht ae ugse great suggested have that reviewers anonymous and editors thank finally We Technological and Scientific for Council National Brazilian thank also We . We thank the Smithsonian Tropical Research Institute and the University of Sãoof University the Instituteand Research SmithsonianTropical the thank . We ep n il work, field in help LASTROP roofreading ofthis article. Nq gat 166177/2014 grant# CNPq,
We very We
) on
and equipment use equipment
much appreciate appreciate much from the Forest Ecology and Restoration Laboratory ( Laboratory Restoration and Ecology Forest the from
especially Cps gat BX 7030/15 BEX grant# (Capes,
a dt ue fr nlss n rsls ipae and displayed results and analyses for used data Raw ( Sardinilla -
SYLVA French network. French SYLVA as Bc Sorrini Bech Taísi ,
and - . site 5
- and 2018/18416 and 8), all staff that maintained the study sites and sites study the maintained that staff all
to colleagues from colleagues to ) .
We sincerely thank Alistair Campbell for Campbell Alistair thank sincerely We duardo Moré de Mattos for their help on help their Mattosfor de Moré duardo oriain o te mrvmn of Improvement the for Coordination –
neb fr rvdn u wt their with us providing for Anhembi
n Lí Bradn, o data for Bernardini, Luís and - ) n So al Rese Paulo São and 0) - 2
) for funding this project. this funding for ) the Tropical Silviculture Tropical the
( Anhembi site LERF )
arch and and )
728 727 726 725 724 723 722 721 729
a first draft of the manuscript and PHSB, JG,PHSB, and manuscript the of firstdraft a data out carried ( Author This article isprotected rights by All copyright. reserved Accepted Article final version. Sardinilla Sardinilla s’ MMD, PHSBMMD,
contribution and and
Anhembi Anhembi
analyses. MMD, PHSB, RAM, CIFZ and CIFZ RAM, PHSB, MMD,analyses.
and epciey. MMD respectively).
CP designed the designed CP
research. CP
RAM and CP contributed substantially to theto substantially contributed CP and RAM n CF cletd aa RM and RAM data. collected CIFZ and
and JG interpreted data. MMD wrote MMD data. interpreted JG
JLS designed treeplantations designed JLS
WHB
737 736 735 734 733 732 731 730 752 751 750 749 748 747 746 745 744 743 742 741 740 739 738
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Ratcliffe S, Wirth C, Jucker T, van der Plas F, Scherer F, Plas der van T, Jucker C, Wirth S, Ratcliffe Team Core R This article isprotected rights by All copyright. reserved AcceptedScherer M Loreau N,Kunert PotvinC, Paquette A, J, Sapijanskas Ruiz Rodrigues Article MM Hirota FJ, Ponzoni AC, Martensen JP, Metzger MC, Ribeiro csse fntoig eain i Erpa frss eed n environmental on depend forests European in relations functioning ecosystem B, Ohse H, Bruelheide R, Benavides E, Allan Vienna, Computing, Statistical Austria. of Sciences Academy global in regrowth forest of Role oet iest ad ucin Tmeae n Bra Sses Berlin/Heidelberg: Systems. Boreal and Temperate Function: and Diversity Forest b functional E Schulze G, Reynolds differences. capture light enhances forest. natural a and plantation a in diversity functional and species Comparing diversity? tree Forest. Conservation Atlantic Brazilian the in experience of years 30 forests: diversity forest remaining the is how and distributed? conservation. for Implications left, is much How Forest: Atlantic Brazilian context. - Jaen MC, Potvin C Potvin MC, Jaen - Lorenzen M, Potvin C, Koricheva J, Schmid B, Hector A, Bornik Z, Z, Bornik A, Hector B, Schmid J, Koricheva C, Potvin M, Lorenzen
The New phytologist R Lm RF Gnof S Nv AG Nave S, Gandolfi RAF, Lima RR, Ecology Letters .
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Lorenzen M, Körner C, SchulzeE C, Körner M, Lorenzen https://www.R . 2009
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Sercu BK, Baeten L, van Coillie F, Martel A, Lens L, Verheyen K, Bonte D Bonte K, Verheyen L, Lens A, Martel F, Coillie van L, Baeten BK, Sercu C Potvin J, Bauhus CA, Nock A, Fichtner A, Dănescu JA, Schwarz F, Schnabel This article isprotected rights by All copyright. reserved Accepted E Bernhardt L, Jackson K, Engelhardt JM, Bullock A, Symstad J, Wright Cavender A, Paquette LJ, Williams P Coppin I, Jonckheere GJ, Smith F, Baret M, Weiss Castro F, Buscot S, Both JA, Blum J, Bauhus A, Schuldt S, Trogisch ArticleTreeDivNet M Vellend DS, Srivastava mature temperate forests.mature temperate li affect diversity and identity species tree How experiment. 2019 Springer results? In: Naeem S, Bunker DE, Hector A, Loreau M, Perrings C, eds. Biodiversity, eds. C, Perrings M, Loreau A, Hector DE, Bunker S, Naeem In: results? improve approach integrated an will function: ecosystem and biodiversity Restoring mixtures. Ecology Evolution and species in overyielding explains crowns tree in complementarity sampling. determina (LAI) index area leaf situ 10674. multifunctionality. forest assessing comprehensively for D, Eichenberg W, Durka D, Chesters http://www.treedivnet.ugent.be/experiments.html. December 2017].[accessed 11 to conservation? Dies f rdciiy n is eprl tblt i a rpcl re diversity tree tropical a in stability temporal its and productivity of Drivers .
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Figures This article isprotected rights by All copyright. reserved eachmeasured richness within level. 2016 April, and March Accepted in measurements P the at levels points (black iPAR = within measured. plots all represent the of part are which 2001) in (established 2 monocultures, season rainy the Mean Figure 1. Article < Cedrela 0.0001 iPAR each
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in distinct colors) over horizontal measurements grids, at at grids, measurements horizontal over colors) distinct in
Anhembi
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site (Anhembi, and Brazil) during thedry(C) (D) wet seasons.
site’s
en necpe photosyntheticall intercepted Mean an A ad ih iest (B) diversity high and (A) main ad 5 cniec itras sae aes maue fr different for measured areas) (shaded intervals confidence 95% and ) n legends in , t diffe at ), plantations atv rdain iA, ons in points (iPAR, radiation active y et egt wti te aoy at canopy, the within heights rent
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This article isprotected rights by All copyright. reserved Accepted fromthe fivespecies the main Here, visualization. their better for only iPAR (HD) to diversity converted were values Significant and data means. between difference treatments. both in present is species of a when shown magnitude to related confidence 95% at contrasts is color of Intensity column) in treatment five Ls) + 4) Hc + and (Co T4 Ls), + Co + (Ae T3 Tr), + Ls + (Ae T2 combinations: species three rosea. Tabebuia monocultures: 1) Treatments: column. the excelsum in Anacardium treatment the from and row the in treatment the from iPAR mean between difference the represent cells the within Numbers Sardinilla Figure 5. Article constituent
site’s main plantation (Sardinilla, Panama, e Panama, (Sardinilla, plantation main site’s
- species combination: A. A. combination: species ; 2) two species combinations: T1 (Co + Hc), T5 (Hc + Tr), T6 (Ae + Tr); 3)Tr); + (Ae T6 Tr), + (Hc T5 Hc), + (Co T1 combinations: species two 2) ; Comparisons of contrasts between different treatments’ mean iPAR mean treatments’ different between contrasts of Comparisons plant monocultures, and HD HD and monocultures, , Co = Co , ation are colored in green, while negative differences are colored in purple. in colored are differences negative while green, in colored are a nt osdrd sne e atd o opr mxue to mixtures compare to wanted we since considered, not was Cedrela level are in bold and contain an asterisk. an contain and bold in are level plantation
odorata oiie differences Positive . , Hc = Hc , plantation
Hura crepitans Hura
counted on a larger species pool than pool species larger a on counted 2016 in measured 2001), in stablished Analyses were performed using R:FR using performed were Analyses
(between treatment in row and and row in treatment (between , Ls = = Ls , Luehea seemannii Luehea Differences are only are Differences h high the
, Tr = = Tr , at the at e = Ae .