紫茎泽兰和飞机草的形态和光合特性对磷营养的响应

比较研究了紫茎泽兰和飞机草的生长、形态、生物量分配和光合特性对磷营养的可塑性反应,及与其入侵性的关系.结果表明,两种入侵植物对磷营养变化表现出很强的可塑性和适应性.低磷时,两种植物的根生物量比增大,利于养分吸收;高磷时,两种植物的比叶面积、最大净光合速率、光饱和点、单位面积的叶绿素和类胡萝卜素含量增大,同化面积和同化能力增强,利于碳积累.相比之下,紫茎泽兰对磷的适应性更强.随供磷量的增加,紫茎泽兰和飞机草的相对生长速率、总生物量、株高、分枝数、叶面积指数和最大净光合速率均显著增大,过量磷素对上述参数抑制较小,表明两种入侵植物偏好较高的磷环境,土壤磷含量升高有利于其入侵,并在高磷时,通过增大株高、分枝数和叶面积指数荫蔽排挤本地种.在本地种基本停止生长的干季,紫茎泽兰和飞机草仍维持较高的相对生长速率.这也与其入侵性密切相关.     

西双版纳外来入侵植物及其共存种叶片氮、磷化学计量特征

诊断外来植物和本地种的资源利用方式是入侵生态学研究的热点问题。叶片氮(N)、磷(P)含量和化学计量特征可以反映受侵地区植物的N、P吸收能力和限制状况, 为把握外来植物的入侵能力、为本地植物共存或消失的机制提供基础科学依据。该研究以我国西南地区典型外来入侵植物(飞机草(Chromolaena odorata)、紫茎泽兰(又叫破坏草, Ageratina adenophora))及其共存的本地植物为对象, 探讨不同入侵条件下入侵种及其共存的本地植物的N、P利用策略。在云南西双版纳孔明山研究区, 调查了无入侵条件下和飞机草与紫茎泽兰的不同入侵程度(按入侵种生物量比例划分)下的物种数量和生物量, 分析了主要植物的叶片N、P含量和N:P值。结果显示: 尽管群落地上生物量随飞机草和紫茎泽兰入侵程度增加而增加, 但本地植物种数随飞机草和紫茎泽兰入侵程度增加而显著减少。飞机草与紫茎泽兰的叶片N、P平均含量无显著差异, 但均显著高于无入侵条件下的本地植物以及与其共存的本地植物。两种入侵植物的N、P含量均随它们所占样方总生物量的比例增大而升高, 本地植物N含量也有相似的变化趋势。当对比入侵和无入侵两类样方的同种植物N、P和N:P变化时发现多数本地种叶片P含量呈降低趋势, N含量和N:P呈升高趋势。根据叶片N、P绝对含量和N:P值及其随入侵的变化规律, 推测入侵可能提高了植物的N可利用性, 但本地植物仍然受N限制; 入侵植物N:P值总体小于10与其具有相对于N吸收的较高的P吸收能力有关。该研究揭示了西南典型外来入侵植物具有较强的N、P吸收富集能力。     

氮肥和种植密度对紫茎泽兰生长和竞争的影响

紫茎泽兰（Ageratina adenophora）是我国西南热带亚热带地区危害最严重的外来入侵植物之一。本研究在野外设置不同氮肥供给水平和种植密度下的紫茎泽兰、拔毒散（Sida szechuensis）、伏生臂形草（Brachiaria decumbens）和非洲狗尾草（Setaria anceps）的单种和紫茎泽兰与其他3种植物混种的人工群落,从植被-土壤相互作用的角度分析紫茎泽兰的入侵机制。结果表明：单种时4种受试植物均表现了对氮营养的响应,随氮肥水平的升高,根冠比降低,比叶面积升高;通过比较总生物量增幅,氮响应能力非洲狗尾草>伏生臂形草>紫茎泽兰>拔毒散。混种时受试植物与紫茎泽兰的生物量比和株高也表明非洲狗尾草的竞争能力>伏生臂形草>紫茎泽兰>拔毒散;与拔毒散混种的紫茎泽兰以及与 紫茎泽兰混种的伏生臂形草和非洲狗尾草的种内竞争随种植密度的增大而增大,导致种间竞争能力下降,在高种植密度下对混种物种的抑制作用减弱。紫茎泽兰在氮响应能力、竞争能力和表型可塑性方面均强于本地种,可能与其入侵机制有关;牧草的氮响应能力和竞争能力强于紫茎泽兰,其中非洲狗尾草又强于伏生臂形草,可能更适宜于入侵地的替代控制。     

人工群落中苗期紫茎泽兰的化感作用和对光环境的适应

在有、无活性炭的条件下分别构建紫茎泽兰（Ageratinaa denophora）与4种受体植物——飞机草（Chromolaena odorata）、鬼针草（Bidenspilosa）、胜红蓟（Ageratum conyzoides）和兰花菊三七（Gynura sp.）混种的人工群落,研究了群落中紫茎泽兰的化感作用和对群落光环境的适应,探讨了化感作用和光适应特性与其入侵性的关系。结果表明,活性炭处理对4种受体植物的生长、生理特性影响不显著,说明苗期紫茎泽兰化感作用不明显,推测入侵初期化感作用不是紫茎泽兰排挤本地种的主要原因。4种受体植物可以通过化感作用对紫茎泽兰产生某些影响,但群落的光环境对其影响更大。紫茎泽兰能很好地适应群落中不同的光环境。苗期紫茎泽兰处于群落下层,叶片受光指数低,此时它能长期忍耐并缓慢生长;随着叶片受光指数的升高,其最大净光合速率、超氧化物歧化酶活性、叶绿素a／b比、总生物量、总叶面积、地茎、叶片数和分支数升高,比叶面积和比茎长降低,这有利于它维持叶片能量平衡并导致对邻近植物的严重遮荫。紫茎泽兰强的光适应能力、强光下对其它物种的遮荫效应与其入侵性密切相关。     

氮、磷养分对飞机草营养器官表型可塑性的影响

采用温室盆栽试验研究了不同氮、磷水平对入侵植物飞机草(Chromolaena odorata)营养器官表型可塑性的影响。结果表明:随着氮、磷水平的上升,飞机草的分枝数量、分枝长度、叶片数、总叶面积、总生物量以及茎、叶器官生物量显著增加。飞机草的根生物量比、根冠比随着氮、磷水平的升高显著下降;茎生物量比在供氮(磷)量达0.05 g·kg-1时显著增加,之后保持稳定;叶生物量比随氮水平的增加先降后升,但其受磷水平变化的影响较小。叶面积比、叶根比、比叶面积和平均相对生长速率随着氮、磷水平的上升显著增加,但叶面积比、叶根比和比叶面积在供磷量≥0.05 g·kg-1时的差异不明显。飞机草的分枝数量、分枝长度、叶片数、总叶面积、根生物量比、根冠比、叶根比以及茎、叶与植株总生物量等指标的可塑性指数较高,并且对氮素的响应更强。表明氮、磷水平能够显著影响飞机草的植株生长,飞机草亦能够通过植株形态、结构以及生物量积累与分配的调整来适应多变的养分环境,并表现出较高的可塑性。     

光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响

 比较研究了不同光强下生长的（透光率分别为12.5%、36%、50%、100%）两种入侵性不同的外来种——紫茎泽兰（Eupatorium adenophorum）和兰花菊三七（Gynura sp.）的生物量分配、叶片形态和生长特性。结果表明: 1）两种植物叶片形态对光环境的反应相似。弱光下比叶面积（SLA）、平均单叶面积（MLS）和叶面积比（LAR）较大,随着光强的升高,SLA、MLS、LAR和叶根比（LARMR）降低。2）100%光强下紫茎泽兰叶生物量比（LMR）、叶重分数（LMF）和叶面积指数高于低光强下的值,也高于兰花菊三七,支持结构生物量比（SBR）则相反。强光下紫茎泽兰叶片自遮荫严重,这可能是其表现入侵性的重要原因之一;兰花菊三七分枝较多,避免了叶片自遮荫,较多的分枝利于种子形成对其入侵有利。3）随生长环境光强的升高,两种植物的净同化速率（NAR）、相对生长速率（RGR）和生长对NAR的响应系数均升高（但100%光强下兰花菊三七RGR降低）,平均叶面积比（LARm）和生长对LARm的响应系数均降低,但不同光强下LARm对生长的影响始终大于NAR。4）随着光强的减弱,两种植物都增加高度以截获更多光能,但它们的生物量分配策略不同,紫茎泽兰根生物量比（RMR）降低,SBR增大,而兰花菊三七SBR降低,RMR增大。紫茎泽兰的生物量分配策略更好的反应了弱光环境中的资源变化情况。结论：紫茎泽兰对光环境的适应能力强于兰花菊三七。     

两种入侵植物与三种本地植物根系特征的比较研究

以菊科2种入侵植物飞机草(Chromolaena odorata(L.)R.M.King&H.Rob)和紫茎泽兰(Eupatorium adenophorum Speng)以及生活型相似的3种本地植物异叶泽兰(Eupatorium heterophyllum DC.)、佩兰(Eupatorium fortune Turcz.)和白头婆(Eupatorium japonicum Thunb.)为研究对象,以单种为对照,分析5种植物的株高、生物量以及细根形态和结构特征对混种(竞争)的响应。结果显示,在单种情况下,2种入侵植物的株高和地上生物量比3种本地植物高。2种入侵植物的株高和地上生物量在单种和混种下无显著差异,但在混种下3种本地植物的株高、地上生物量、根生物量、细根长度、细根表面积和体积与单种相比显著降低。在混种情况下,2种入侵植物的根组织密度显著降低,但比根长和比根面积仅在紫茎泽兰中显著增加。5种植物在单种和混种下的株高与细根长度、根生物量呈显著正相关,地上生物量与细根体积、表面积呈显著正相关。主成分分析结果表明入侵植物位于株高和地上生物量更高的一端。说明在与本地植物竞争的过程中,飞机草和紫茎泽兰能通过调整根系碳投资策略实现更强的竞争力。     

四种不同生活型植物幼苗对喀斯特生境干旱的生理生态适应性

喀斯特石漠化是我国西南喀斯特地区最严重的生态环境问题, 生境干旱是限制该地区植物生长的主要因素之一, 掌握喀斯特植被不同演替阶段不同生活型植物对干旱胁迫的适应策略有助于提高植被恢复的成功率。通过人工模拟4种干旱强度, 测定叶片水势、气体交换、叶绿素荧光、光合色素含量、渗透调节物质浓度、抗氧化酶活性以及生物量, 研究了喀斯特地区4种不同生活型植物幼苗对干旱胁迫的适应策略。这4种植物为常绿灌木火棘(Pyracantha fortuneana)、落叶灌木小果蔷薇(Rosa cymosa)、常绿乔木猴樟(Cinnamomum bodinieri)和落叶乔木圆果化香树(Platycarya longipes)。结果表明: 随着干旱程度的加深, 4种植物幼苗的叶片水势、光合能力、叶绿素含量、生物量增长、叶重比(LMR)、叶面积比(LAR)和比叶面积(SLA)逐渐下降, 而热耗散(NPQ)、类胡萝卜素与叶绿素含量比值、丙二醛含量和根重比(RMR)逐渐上升; 圆果化香树和猴樟的水分利用效率(A_n/g_s)、渗透调节物质浓度和抗氧化酶活性呈先升高后降低的趋势, 而火棘和小果蔷薇的A_n/g_s、脯氨酸含量和超氧化物歧化酶活性呈上升趋势。严重干旱下, 火棘和小果蔷薇幼苗的叶片水势和叶绿素含量下降较少, 具有较高的光合能力和生物量增长, 这主要是由于它们具有较低的SLA和LAR、较高的NPQ和A_n/g_s以及较高的渗透调节能力和抗氧化保护能力。中度干旱下, 猴樟幼苗叶片水势下降很少, LMR和LAR也较高, 脯氨酸含量和抗氧化酶活性非常高。但在严重干旱下, 其叶片水势、LMR、LAR和生物量增长大幅度下降, 最大光化学效率和光合速率也非常低, 渗透调节能力与抗氧化酶活性大幅度下降至正常水平以下。水分好的条件下, 圆果化香树幼苗具有较高的RMR以吸收充足的水分, 具有较高的LAR和叶绿素含量, 保证了生物量的大量积累。然而, 干旱胁迫致使其生物量大幅度下降, 主要是由于LMR、LAR、气体交换和叶绿素含量的大量下降以减少蒸腾面积、水分散失和对光能的吸收。研究结果表明, 火棘、小果蔷薇和猴樟幼苗主要采用耐旱策略, 其中猴樟抗严重干旱的能力较弱; 圆果化香树幼苗对干旱胁迫更为敏感, 主要采取避旱策略。     

氮碳添加和丛枝菌根对外来入侵植物豚草的影响

通过温室控制试验,研究了氮碳素添加及丛枝菌根(AM)对外来入侵植物豚草生长的影响.结果表明： 氮素添加对豚草的形态指标、生物量及其分配特征和生长速率均没有显著影响,却使豚草地上器官和根的氮含量显著增加;碳素添加增加了土壤中的有效氮含量,促使豚草增加营养吸收器官的生物量分配以缓解养分(氮素)胁迫,导致分枝数、总叶面积、比叶面积和叶生物量比明显降低,总生物量显著减少.豚草与AM真菌共生增强了其适应土壤氮素变化的能力,增加了比叶面积,提高了豚草的资源获取能力,其作用在土壤有效氮含量低时更加明显.AM真菌对豚草适应低氮生境有着重要意义.     

四川攀西地区云南松群落物种多样性和谱系多样性对紫茎泽兰入侵的影响

外来植物入侵严重威胁着入侵地本土植物多样性和生态系统功能, 认识外来物种的入侵机制有助于提高对入侵植物的防控能力。本文以攀西地区云南松(Pinus yunnanensis)林下外来入侵植物紫茎泽兰(Ageratina adenophora)为研究对象, 基于大量野外群落调查, 从群落可入侵性入手, 分析了环境因子和群落物种多样性、谱系多样性等群落生态学特征对紫茎泽兰入侵的影响。结果表明: 海拔、坡向和火烧强度等环境因子和冠层郁闭度、灌木层盖度等生物因子对研究区紫茎泽兰入侵强度没有显著影响(P > 0.05); 但群落灌木层物种多样性和草本层组分种与紫茎泽兰的亲缘关系显著影响紫茎泽兰的入侵强度(P < 0.05), 说明灌木层对光照等环境资源的占用和草本层(同层)物种对相似资源的竞争能够在很大程度上抑制紫茎泽兰的入侵。     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Net Assimilation Rate Determines the Growth Rates of 14 Species of Subtropical Forest Trees

Growth rates are of fundamental importance for plants, as individual size affects myriad ecological processes. We determined the factors that generate variation in RGR among 14 species of trees and shrubs that are abundant in subtropical Chinese forests. We grew seedlings for two years at four light levels in a shade-house experiment. We monitored the growth of every juvenile plant every two weeks. After one and two years, we destructively harvested individuals and measured their functional traits and gas-exchange rates. After calculating individual biomass trajectories, we estimated relative growth rates using nonlinear growth functions. We decomposed the variance in log(RGR) to evaluate the relationships of RGR with its components: specific leaf area (SLA), net assimilation rate (NAR) and leaf mass ratio (LMR). We found that variation in NAR was the primary determinant of variation in RGR at all light levels, whereas SLA and LMR made smaller contributions. Furthermore, NAR was strongly and positively associated with area-based photosynthetic rate and leaf nitrogen content. Photosynthetic rate and leaf nitrogen concentration can, therefore, be good predictors of growth in woody species.     

Relative growth rate in phylogenetically related deciduous and evergreen woody species

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR2SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.     

Contribution of root growth responses to leaf traits and relative growth rate of Populus alba under different water-table conditions

Water-table depth variations alter root growth response and may affect whole-plant growth in arid and semi-arid regions. We examined how root biomass allocation and root morphological traits affect the leaf physiological and morphological traits and whole-plant growth of Populus alba growing under different water tables. We exposed 1-year-old P. alba cuttings to contrasting soil–water conditions via water table changes in a greenhouse for 90 days. We examined relationships among net assimilation rate (NAR) and other growth components obtained from our published data for trees harvested every 30 days. Strongly negative correlations were found between RMR and root morphological traits. Root mass ratio had a strong negative relationship with LMR, and proportion of fine-root biomass per total root biomass was positively correlated with SLA and NAR. Both NAR and leaf area ratio were important determinants of variation in relative growth rate (RGR). Leaf mass ratio (LMR) and specific leaf area (SLA) were positively correlated with RGR; the correlation was stronger in the case of LMR. Along a water-table gradient, negative relationships between root growth responses are likely to indirectly influence RGR through changes in NAR, LMR, and SLA.     

Seedling growth strategies in Bauhinia species: comparing lianas and trees

BACKGROUND AND AIMS: Lianas are expected to differ from trees in their growth strategies. As a result these two groups of woody species will have different spatial distributions: lianas are more common in high light environments. This study determines the differences in growth patterns, biomass allocation and leaf traits in five closely related liana and tree species of the genus Bauhinia. METHODS: Seedlings of two light-demanding lianas (Bauhinia tenuiflora and B. claviflora), one shade-tolerant liana (B. aurea), and two light-demanding trees (B. purpurea and B. monandra) were grown in a shadehouse at 25% of full sunlight. A range of physiological, morphological and biomass parameters at the leaf and whole plant level were compared among these five species. KEY RESULTS: The two light-demanding liana species had higher relative growth rate (RGR), allocated more biomass to leaf production [higher leaf mass fraction (LMF) and higher leaf area ratio (LAR)] and stem mass fraction (SMF), and less biomass to the roots [root mass fraction (RMF)] than the two tree species. The shade-tolerant liana had the lowest RGR of all five species, and had a higher RMF, lower SMF and similar LMF than the two light-demanding liana species. The two light-demanding lianas had lower photosynthetic rates per unit area (A(area)) and similar photosynthetic rates per unit mass (A(mass)) than the trees. Across species, RGR was positively related to SLA, but not to LAR and A(area). CONCLUSIONS: It is concluded that the faster growth of light-demanding lianas compared with light-demanding trees is based on morphological parameters (SLA, LMF and LAR), and cannot be attributed to higher photosynthetic rates at the leaf level. The shade-tolerant liana exhibited a slow-growth strategy, compared with the light-demanding species.     

Traits of an invasive grass conferring an early growth advantage over native grasses

Aims Invasive species often have higher relative growth rates (RGR) than their native counterparts. Nutrient use efficiency, total leaf area and specific leaf area (SLA) are traits that may confer RGR differences between natives and invasives, but trait differences are less prominent when the invasive species belongs to the same plant functional type as the dominant native species. Here, we test if traits displayed soon after germination confer an early size advantage. Specifically, we predicted that invasive species seedlings grow faster than the natives because they lack trade-offs that more strongly constrain the growth of native species.Methods We quantified plant morphological and physiological traits and RGR during early seedling growth at high and low nutrient levels in three dominant perennial native C₄ grasses: Panicum virgatum L. (switchgrass), Schizachyrium scoparium (Michx.) Nash (little bluestem) and Andropogon gerardii Vitman (big bluestem); and a perennial C₄ exotic invasive grass, Sorghum halepense (L.) Pers. (Johnsongrass).Important findings After 2 weeks of growth, Johnsongrass seedlings had greater biomass, SLA and photosynthetic nitrogen use efficiency, but lower leaf N concentrations (% leaf N) and root:shoot ratio than natives. As growth continued, Johnsongrass more quickly produced larger and thicker leaves than the natives, which dampened the growth advantage past the first 2 to 3 weeks of growth. Investment in carbon gain appears to be the best explanation for the early growth advantage of Johnsongrass. In natives, growth was constrained by an apparent trade-off between allocation to root biomass, which reduced SLA, and production of leaves with high N content, which increased carbon gain. In Johnsongrass, root:shoot ratio did not interact with other traits, and % leaf N was decoupled from RGR as a result of a trade-off between the positive indirect association of % leaf N with RGR and the negative direct association of % leaf N with RGR.     

Relative growth rate and biomass allocation in ten woody species with different leaf longevity using phylogenetic independent contrasts (PICs)

In this study, we compare the relative growth rate (RGR) and biomass allocation of 10 woody species (5 deciduous and 5 evergreen) from the Mediterranean region using phylogenetic independent contrasts (PICs) to test if these two functional groups differ in these traits. In general, the results were similar when using PICs or without taking into account phylogenetic relations. Deciduous species had a higher RGR than evergreen species, due to the higher net assimilation rate (NAR). Deciduous species had a higher specific leaf area (SLA) but a lower leaf mass ratio (LMR), resulting in a similar LAR for deciduous and evergreen species (LAR = SLA x LMR). In some cases, the use of PICs revealed patterns that would not have appeared if phylogeny had been overlooked. For example, there was no significant correlation between RGR and final dry mass (after 4 months of growth) but PICs revealed that there was a positive relation between these two variables in all deciduous-evergreen pairs. In general, RGR decreased with time and this temporal variation was due primarily to NAR variations (r = 0.79, p < 0.01), and also to variations in LAR (r = 0.69, p < 0.05). Considering the phylogeny, the only variable constantly different for all deciduous-evergreen pairs was SLA. This result, and the fact that SLA was the best correlated variable with RGR (r = 0.81, p < 0.01), reinforce the value of SLA as a variable closely associated to growth and to the functional groups (deciduous vs. evergreen).     

Shade tolerance and herbivory are associated with RGR of tree species via different functional traits

• Relative growth rate (RGR) plays an important role in plant adaptation to the light environment through the growth potential/survival trade‐off. RGR is a complex trait with physiological and biomass allocation components. It has been argued that herbivory may influence the evolution of plant strategies to cope with the light environment, but little is known about the relation between susceptibility to herbivores and growth‐related functional traits.
• Here, we examined in 11 evergreen tree species from a temperate rainforest the association between growth‐related functional traits and (i) species’ shade‐tolerance, and (ii) herbivory rate in the field. We aimed at elucidating the differential linkage of shade and herbivory with RGR via growth‐related functional traits.
• We found that RGR was associated negatively with shade‐tolerance and positively with herbivory rate. However, herbivory rate and shade‐tolerance were not significantly related. RGR was determined mainly by photosynthetic rate (A_max) and specific leaf area (SLA). Results suggest that shade tolerance and herbivore resistance do not covary with the same functional traits. Whereas shade‐tolerance was strongly related to A_max and to a lesser extent to leaf mass ratio (LMR) and dark respiration (R_d), herbivory rate was closely related to allocation traits (SLA and LMR) and slightly associated with protein content.
• The effects of low light on RGR would be mediated by A_max, while the effects of herbivory on RGR would be mediated by SLA. Our findings suggest that shade and herbivores may differentially contribute to shape RGR of tree species through their effects on different resource‐uptake functional traits.