Temporal intraspecific trait variability drives responses of functional diversity to interannual aridity variation in grasslands

Interannual climate variation alters functional diversity through intraspecific trait variability and species turnover. We examined these diversity elements in three types of grasslands in northern China, including two temperate steppes and an alpine meadow. We evaluated the differences in community‐weighted means (CWM) of plant traits and functional dispersion (FDis) between 2 years with contrasting aridity in the growing season. Four traits were measured: specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen concentration (LNC), and the maximum plant height (H). CWM for SLA of the alpine meadow increased in the dry year while that of the temperate steppe in Qinghai showed opposing trends. CWM of LDMC in two temperate steppes became higher and CWM of LNC in all grasslands became lower in the dry year. Compared with the wet year, FDis of LDMC in the alpine meadow and FDis of LNC in the temperate steppe in Qinghai decreased in the dry year. FDis of H was higher in the dry year for two temperate steppes. Only in the temperate steppe in Qinghai did the multi‐FDis of all traits experience a significant increase in the dry year. Most of the changes in CWM and FDis between 2 years were explained by intraspecific trait variation rather than shifts in species composition. This study highlights that temporal intraspecific trait variation contributes to functional responses to environmental changes. Our results also suggest it would be necessary to consider habitat types when modeling ecosystem responses to climate changes, as different grasslands showed different response patterns.     

去除干扰对内蒙古典型草原植物叶片功能属性的影响

以内蒙古典型草原为研究对象,选取放牧和割草、去除放牧、去除放牧和割草样地进行群落调查和叶片属性测量,比较分析各样地土壤性质、群落生产力及主要物种的比叶面积(SLA,Specific Leaf Area)、叶片干物质含量(LDMC,Leaf Dry Matter Content)、叶片氮含量(LNC,Leaf Nitrogen Concentration)在个体、功能群和群落水平对去除干扰的响应。结果表明,1)去除干扰处理在短期对土壤特性和群落生产力的影响不显著;2)多数物种在放牧和割草样地SLA较低,说明典型草原多数物种的SLA表现为放牧逃避;3)不同功能群植物叶片属性对去除干扰的响应不一致,去除放牧后,多年生杂类草的SLA和LDMC不受影响,但LNC变小;多年生禾草的SLA增加,而LDMC和LNC无显著变化。一年生植物在去除放牧和割草后,LNC显著增加。去除割草后,多年生禾草SLA减小,而多年生杂类草SLA、LNC增加,LDMC减小;4)在群落水平,放牧和割草样地由于较占优势的多年生禾草SLA较低,群落比叶面积最低,在去除放牧和割草样地,群落叶片氮含量显著增加;5)在内蒙古典型草原,LDMC能够很好地将多年生禾草和多年生杂类草区分,SLA在个体、功能群和群落水平均比LDMC敏感。     

退化草地物种种内和种间功能性状变异对放牧的响应

研究草地植物功能性状变异对放牧干扰的响应机制,有助于解析草地生态系统群落构建和功能多样性维持机制及植物对环境的适应及权衡策略。以科尔沁沙地退化草地为对象,研究围封和放牧草地物种多种功能性状(植株高度、根系长度、叶面积、根面积、叶片干物质含量、地上干物质含量、根系干物质含量、比叶面积、比根长和根冠比)变异特征及其对放牧干扰的响应机制。结果表明: 退化草地物种种间性状变异明显高于种内变异,种间性状变异对总体变异的贡献率占比高达70.2%～95.1%,而种内变异仅为4.9%～29.8%,但群落构建中物种的种内变异仍不可忽视。放牧草地物种种间性状变异低于围封草地,且放牧草地种内变异增加而种间变异减小。放牧导致不耐牧的优良禾本科牧草叶面积和叶片干物质含量下降而比根长增加,但耐牧的杂类草则通过增加叶面积和叶片干物质含量并降低比根长来提高在群落中的优势地位。退化草地对放牧响应较为敏感的功能性状有叶面积、叶片干物质含量、比根长和比叶面积。植物叶片性状和根系性状自身及彼此间均呈显著正相关,但放牧会增强根系性状的协同效应而减弱叶片性状的协同效应。说明放牧会驱动植物个体和种群功能性状权衡策略发生改变,进而起到调控植物群落结构和功能的作用。     

Intraspecific trait variation and allocation strategies of calcareous grassland species: Results from a restoration experiment

Intra- and interspecific trait variation express the response of plants dealing with different environmental conditions. We measured root and leaf traits on 14 species of calcareous grasslands in a restoration experiment. We aimed at identifying intraspecific differences in biomass allocation and functional plant traits under contrasting soil conditions by comparing plants growing in ancient grassland and two restored grasslands on ex-arable land, one of them with topsoil removal. Relative importance of trait variation within and among species, and among site was assessed by variance partitioning. Interspecific variation was more important than intraspecific variation, but the contribution of the latter to total variation was considerable, especially for specific leaf area. Changes in soil properties due to topsoil removal resulted in lower values of plant height, specific leaf area and specific root length compared to the control (ancient grassland). Soil fertility found in the treatment without top soil removal did not affect plant plasticity compared to the control. The study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species. We conclude that caution must be taken when using mean trait values for plastic species or when working with environmental gradients.     

柯-青冈常绿阔叶林优势树种叶片性状变异及适应策略

厘清叶片功能性状的变异及性状间的权衡关系，对揭示植物对环境变化的响应及适应策略具有重要意义。以中亚热带柯(Lithocarpus glaber)-青冈(Cyclobalanopsis glauca)常绿阔叶林为对象，测量了1 hm²固定监测样地内6个优势树种的叶面积(LA)、比叶面积(SLA)、干物质含量(LDMC)、叶片厚度(LT)、叶片碳(LC)、叶片氮(LN)、叶片磷(LP)含量和叶片碳氮比(LC ∶ LN)8个功能性状，采用多重比较、主成分分析(PCA)法分析了群落叶片功能性状的生活型、种内和种间变异及性状间关系。结果表明：(1)群落内叶片功能性状种内、种间差异显著，变异中等(CV： 0.02-0.59)，其结构性状的可塑性较化学性状保守，变异格局符合"性状空间变异分割假说"；针叶树种的LA、SLA显著低于阔叶树种，常绿树种的LC和LDMC最小，而落叶树种的SLA、LN和LP最大以及LT和LC ∶ LN最小。(2)群落叶片功能性状变异主要来源于生活型和种间变异，种内变异亦有显著贡献；生活型对多数性状的贡献率较大，其对LDMC、LC ∶ LN的贡献分别高达93.11%和91.76%；种间变异(LDMC除外)对结构性状的贡献率均高于化学性状；种内变异对LP的贡献率(23.66%)较种间变异高。(3)叶片性状之间多表现出显著相关关系，针叶树和阔叶树在PCA排序图中聚集于不同区域，叶经济型谱在柯-青冈群落中具有适用性。不同树种通过叶片结构、化学性状之间的权衡策略来适应环境变化，从而实现群落物种共存。结果可为理解森林群落物种的环境适应策略、预测群落动态变化和制定植被恢复措施提供科学依据。     

A community‐level test of the leaf‐height‐seed ecology strategy scheme in relation to grazing conditions

Question: Is the assumption of trait independence implied in Westoby's (1998) leaf‐height‐seed (LHS) ecology strategy scheme upheld in a Mediterranean grazing system dominated by annuals? Is the LHS approach applicable at the community level? Location: Northern Israel. Methods: LHS traits (specific leaf area [SLA], plant height and seed mass), and additional leaf traits (leaf dry matter content [LDMC], leaf area, and leaf content of nitrogen [LNC], carbon [LCC], and phosphorus [LPC]), were analyzed at the species and community levels. Treatments included manipulations of grazing intensity (moderate and heavy) and protection from grazing. We focused on species comprising 80% of biomass over all treatments, assuming that these species drive trait relationships and ecosystem processes. Results: At the species level, SLA and seed mass were negatively correlated, and plant height was positively correlated to LCC. SLA, seed mass, and LPC increased with protection from grazing. At the community level, redundancy analysis revealed one principal gradient of variation: SLA, correlated to grazing, versus seed mass and plant height, associated with protection from grazing. We divided community functional parameters into two groups according to grazing response: (1) plant height, seed mass, LDMC, and LCC, associated with protection from grazing, and (2) SLA, associated with grazing. Conclusions: The assumption of independence between LHS traits was not upheld at the species level in this Mediterranean grazing system. At the community level, the LHS approach captured most of the variation associated with protection from grazing, reflecting changes in dominance within the plant community.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

桂林岩溶石山青冈群落植物功能性状的种间和种内变异研究

研究植物功能性状的种间和种内变异,有助于揭示植物对环境的适应策略及其群落构建机制研究。以桂林岩溶石山青冈群落主要木本植物为研究对象,选取叶干物质含量（LDMC）、叶厚度（LT）、比叶面积（SLA）和木材密度（WD）4个功能性状,采用混合线性模型结合方差分解方法对其在种内和种间尺度的变异程度和相对贡献进行分析,同时运用零模型方法探讨驱动岩溶石山青冈群落构建的潜在机制。结果表明：（1） LDMC,LT,SLA和WD这4个性状在种内和种间水平上均存在不同程度的变异幅度,种间变异大于种内变异,但种内变异亦不容忽略。（2）零模型检验表明,将种内变异纳入考虑的环境过滤检测得到改善,基于个体性状值的尺度考虑群落构建机制研究是必要的。（3）对于桂林岩溶石山青冈群落,环境过滤作用确实是群落构建的重要驱动机制,与相似性限制作用和随机作用共同塑造了以青冈为优势物种的群落构建。     

Plant functional diversity mediates the effects of vegetation and soil properties on community-level plant nitrogen use in the restoration of semiarid sandy grassland

Species-rich plant communities use nitrogen (N) more efficiently in grassland ecosystems; however, the role of plant functional diversity in affecting community level plant N-use has received little attention. We examined plant N content, stock and N-use efficiency at community-level along a restoration gradient of sandy grassland (mobile dune, semi-fixed dune, fixed dune and grassland) in Horqin Sand Land, northern China. We used the functional trait-based approach to examine how plant functional diversity, reflected by the most abundant species’ traits (community-weighted mean, CWM) and the dispersion of functional trait values (FDis), affected N-use efficiency in sandy grassland restoration. We further used the structure equation model (SEM) to evaluate the direct or indirect effects of plant species richness, biomass, functional diversity and soil properties on community-level plant N-use efficiency. We found that plant biomass and its N stock increased following sandy grassland restoration, and there were lower plant N content and higher N-use efficiency in semi-fixed dune, fixed dune and grassland as compared with mobile dune. N-use efficiency was positively associated with plant species richness, biomass, CWM plant height, CWM leaf C:N, FDis and soil gradient, but SEM results showed that species richness, CWM leaf C:N, plant biomass and FDis controlled by soil properties were the main factors exerting direct effects. CWM plant height also had a positive effect on N-use efficiency through its indirect effect on plant biomass. Soil gradient increased N-use efficiency through an indirect effect on vegetation rather than a direct effect. Final SEM models based on different plant functional diversity explained over 74% of variances in N-use efficiency. Effects of plant functional diversity on N-use efficiency supported both the mass ratio hypothesis and the complementarity hypothesis. Our results clearly highlight the important role of plant functional diversity in mediating the effects of vegetation and soil properties on community level plant N-use in sandy grassland ecosystems.     

Interspecific competition alters leaf stoichiometry in 20 grassland species

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods.     

鼎湖山南亚热带常绿阔叶林叶功能性状沿群落垂直层次的种内变异

探究功能性状沿着环境梯度如何变化一直以来是基于性状的群落生态学的核心问题之一。尽管功能性状存在种内和种间变异, 但种内变异沿环境梯度如何变化仍有待探究。本文以鼎湖山南亚热带常绿阔叶林1.44 ha塔吊样地内16个树种的2,820个个体为研究对象, 探究4种叶功能性状(比叶面积、叶干物质含量、叶厚度和叶面积)沿群落垂直层次的种内变异。首先, 利用随机效应线性模型量化塔吊样地内的种内变异和种间变异; 其次, 利用Kmeans函数将森林的垂直层次划分为灌木层、亚冠层和林冠层, 并通过构建回归模型探究叶功能性状在群落垂直层次中的种内变异格局。最后, 应用混合线性模型和单因素方差分析的方法探究叶功能性状沿垂直层次的种内变异是否具有物种依赖性。结果表明: 在局域群落中, 并非所有叶功能性状的种内变异都低于种间变异; 叶功能性状在不同垂直层次的种内变异格局存在显著差异, 且种内变异与垂直范围呈正相关; 叶功能性状的种内变异具有较强的物种依赖性, 因此树种差异相对于小环境解释了更多的性状变异; 此外, 不同叶功能性状的种内变异沿垂直层次的变化趋势并不一致。本研究发现种内变异对于物种共存具有重要作用。     

Trait‐mediated community assembly: distinguishing the signatures of biotic and abiotic filters

Conflicting hypotheses predict how traits mediate species establishment and community assembly. Traits of newly establishing individuals are predicted to converge, or be more similar to the resident, preexisting community, when the biotic or abiotic environment favors a single best phenotype, but are predicted to diverge when trait differences reduce competitive interactions. We tested these competing hypotheses using transplant seedlings in an old‐field environment, and assessed the contribution of inter‐ and intra‐specific transplant trait variation to community‐level patterns. Using a soil moisture gradient and resident plant removals, we determined when traits of newly‐establishing plants converge or diverge from the resident community by calculating community weighted mean traits for transplant and resident communities. We saw evidence of environmentally‐ and competitively‐driven trait shifts that resulted in both trait convergence and divergence from the resident community, whose traits reflect the combined effects of both drivers. Leaf dry matter content (LDMC) of transplants diverged in the presence of competition, whereas plant height and stem‐specific density (SSD) showed the opposite pattern, converging with the resident community in their presence. Specific leaf area (SLA) shifted with competition but did not reflect resident community SLA. All transplant traits were influenced by soil moisture, often in an interaction with competition, indicating that the strength of convergence or divergence is contingent on the abiotic environment. Intraspecific differences in transplant traits among treatments were evident in three of four traits; intraspecific height and SLA trends mirrored transplant community‐level trends, whereas intraspecific shifts in SSD were distinct from community‐level trends. Our study shows competition between plant species may cause traits of newly establishing plants to converge with the resident community, as frequently as it selects for trait divergence. These opposing effects of competition suggest that it plays a pervasive role in both intraspecific and species‐level trait differences among communities.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

亚热带常绿落叶阔叶混交林植物功能性状的种间和种内变异

不同物种间的功能性状差异是自然生态系统中物种共存的基础, 而物种内个体间的性状变异对物种的共存和分布同样具有重要作用。本文以湖北星斗山自然保护区亚热带常绿落叶阔叶混交林内28种主要树种(通过物种多度排序获得, 其中常绿和落叶树种各14种)为研究对象, 探讨不同叶习性树种的4种功能性状(比叶面积、叶干物质含量、叶面积和比茎密度)在种间和种内的差异程度。结果表明: (1)常绿和落叶树种在4种功能性状上均存在显著差异, 常绿树种的比叶面积和叶面积显著低于落叶树种, 但叶干物质含量和比茎密度则显著高于落叶树种; (2)比叶面积的变化主要来源于叶习性(57.49%), 叶面积变化主要来源于种间(66.80%)和种内变异(27.52%), 叶干物质含量的变化主要来源于种间(38.12%)和种内(33.88%)变异, 但比茎密度的变化主要来源于种内变异(51.50%), 其次为种间变异(32.52%); (3)常绿和落叶树种种间水平的性状相关性可能掩盖各功能性状之间的相关性。种内变异能够显著影响群落间的植物功能性状差异, 但不同功能性状的种内变异程度存在差异。     

Functional organization of woody plant assemblages along precipitation and human disturbance gradients in a seasonally dry tropical forest

Chronic anthropogenic disturbances (CAD) and rainfall are important drivers of plant community assembly, but little is known about the role played by inter‐ and intraspecific trait variation as communities respond to these pervasive forces. Here, we examined the hypothesis that lower precipitation and higher CAD reduce both intra‐ and interspecific trait variation in Caatinga dry forests. We sampled woody plants across 15 plots along precipitation and CAD gradients and measured resource‐use traits. The effects of precipitation and CAD on RaoQ functional diversity were decomposed into species turnover and intraspecific variability. We used “T‐statistics” to assess the trait sorting from the regional pool to local communities (i.e., external filtering), and within‐community forces leading to low trait overlap (i.e., internal filtering) at individual and species levels. Intraspecific variability explained at least one‐third of the total trait variation and 46% of variation in multitrait diversity across communities. Increasing disturbance reduced multitrait diversity, while precipitation affected some particular traits, such as wood density. Overall, precipitation determined species sorting across communities, while disturbance relaxed internal filters, leading to higher trait overlap within communities due to higher intraspecific variability. Our results suggest that the woody Caatinga flora contains a substantial amount of both inter‐ and intraspecific trait variation. This variation is not randomly distributed within and across communities, but varies according to rainfall conditions and disturbance intensity. These findings reinforce the emerging idea that human disturbances can reorganize plant communities at multiple scales and highlight trait variability as a key biological asset for the resilience of dry forests.     

松嫩草地66种草本植物叶片性状特征

植物叶片功能性状及其相互关系越来越受到关注.以松嫩草地66种草本植物为研究对象,测量叶片干物质含量、比叶面积、叶片厚度、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量,检验性状间的相互关系,比较不同功能群(多年生根茎禾草,多年生丛生禾草,多年生杂类草,1年生或2年生草本)间性状的差异性.结果表明,叶片厚度变异系数最大,比叶面积、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量之间存在显著的正相关关系;叶片于物质含量与叶片磷含量没有显著的相关关系,与其它叶片性状呈显著的负相关关系;叶片厚度只与叶片干物质含量和比叶面积呈显著的负相关关系,与其它叶片性状不相关.叶片干物质含量、比叶面积、叶片厚度、叶片氮、磷含量在4个功能群间差异显著,叶绿素含量和类胡萝卜素含量在各个功能群间差异不显著;多年生根茎禾草和多年生丛生禾草叶片的7个性状差异不显著;多年生根茎禾草和多年生丛生禾草的叶片干物质含量高于多年生杂类草和1年生或2年生草本,其它性状小于这两个功能群.     

东灵山地区不同森林群落叶功能性状比较

植物功能性状(plant functional trait)是近年来生态学研究的热点.其中叶功能性状(leaf functional trait)与植株生物量和植物对资源的获得、利用及利用效率的关系最为密切.研究了东灵山地区叶功能性状之间的关系、叶功能性状与地形因子的关系,并对不同群落叶功能性状进行了比较.通过Pearson相关分析发现,叶干物质含量(LDMC)与比叶面积(SLA)、叶氮浓度(LNC)、叶磷浓度(LPC)、叶钾浓度(LKC)负相关;叶大小与叶厚度正相关;SLA与 LNC、LPC、LKC正相关;LNC与LPC、LKC正相关;LPC与LKC正相关.通过灰色关联度分析发现,对叶大小、LNC、LKC来讲,海拔是各项地形因子中的首要影响因子;对LDMC、叶厚度来讲,坡度对其影响最大;对SLA、LPC来讲,坡位是其首要影响因子.依据乔木层的SLA和LDMC将5种群落分成3类,第一类是黑桦林和山杨林,第二类是辽东栎林,第三类是胡桃楸林和糠椴林.群落的分类情况符合该地带性植被优势度类型的分类情况,LDMC和SLA是最能体现群落间差异的叶功能性状.     

Variations in leaf morphological and chemical traits in response to life stages,plant functional types,and habitat types in an old-growth temperate forest

Intraspecific leaf trait variations are becoming a topic of interest for many ecologists because individual-based traits are essentially the drivers of variations at the community level. Six coexisting major tree species in an old-growth temperate forest, Northeast China (i.e., Abies nephrolepis, Pinus koraiensis, Acer mono, Fraxinus mandshurica, Tilia amurensis, and Ulmus laciniata) were sampled, and three habitat types (i.e., Hab I: high soil organic carbon with a moderate slope; Hab II: low soil organic carbon with a gentle slope; and Hab III: low soil organic carbon with a strong slope) were used in the plot. We performed a two-way ANOVA to compare the specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen content (LNC), leaf phosphorus content (LPC), and leaf carbon content (LCC) between saplings (1 < DBH ≤ 5 cm) and adults (DBH ≥ 10 cm) and between habitat types within species. We simultaneously evaluated the effects of life stage, plant functional type, and habitat type on the six leaf traits. Our results showed that life stage and habitat type had varied influences on the leaf traits of the six species. Life stage was an important determinant for all leaf traits. Plant functional type was included in the best models for SLA, LNC, and LCC. Habitat type had a greater influence on LDMC than SLA. Meanwhile, habitat type had a greater influence on LNC and LPC than LCC. The correlation between leaf traits with local environmental factors varied across different plant functional types and life stages. We suggest conducting individual-based analyses of leaf trait variations according to plant functional type and life stage to understand the plant life strategies along an environmental gradient may improve understanding of the forest dynamics in an old-growth temperate forest.     

环境因子对海岛植物茎、叶功能性状的影响

植物功能性状与环境之间的关系是功能性状研究的重点,海岛作为独特的生态系统,其植物功能性状必然和大陆存在差异。为了明确海岛植被的生态适应机制,该文以平潭岛森林群落为研究对象,通过测定茎、叶10个功能性状,以及地形和土壤9个环境因子,探讨了植物功能性状之间的权衡关系,分析了环境因子对海岛植物功能性状的影响。结果表明:(1)比叶面积(SLA)与叶氮含量(LNC)、叶磷含量(LPC)呈正相关,与叶厚度(LT)、叶干物质含量(LDMC)、茎组织密度(STD)、叶碳含量(LCC)呈负相关; LDMC与LNC、茎氮含量(SNC)呈负相关; LT与STD呈正相关,与LNC和LPC呈负相关; LPC与LNC、SNC呈正相关;茎和叶C、N含量均呈正相关。(2)土壤有机质和TN是海岛植物功能性状的主要土壤影响因子。然而,由于土壤中磷含量的缺乏,LNC、茎磷含量(SPC)、SNC均与土壤全磷呈正相关; LDMC与土壤全氮呈正相关;STD与土壤有机质呈正相关; SLA随着土壤pH的增加而增加。(3)坡位和坡度是海岛植物功能性状的主要地形影响因子。SLA、SPC随着海拔上升而下降; STD、LDMC随着海拔和坡度增大而增大; LNC、LPC阴坡大于阳坡。该研究为海岛植被修复和重建提供了参考依据。     

Leaf trait variation captures climate differences but differs with species irrespective of functional group

Aims To clarify whether variation in leaf traits with climate differs with scale, i.e. across species and within a species, and to detect whether plant functional group affects species-specific response.Methods Leaf dry matter content (LDMC), specific leaf area (SLA), mass- and area-based leaf N (N mass, N area) and leaf P concentrations (P mass, P area) and leaf chlorophyll concentration (SPAD) were measured for 92 woody plant species in two botanical gardens in China. The two gardens share plant species in common but differ in climate. Leaf trait variation between the two gardens was examined via mean comparison at three scales: all species together, species grouped into plant functional groups and within a species. A meta-analysis was performed to summarize the species-specific responses.Important findings At the scale of all species together, LDMC, SLA, P mass and N mass were significantly lower in the dry-cold habitat than in the wet-warm one, whereas N area and SPAD showed an inverse pattern, indicating a significant environmental effect. The meta-analysis showed that the above-mentioned patterns persisted for SLA, N area and SPAD but not for the other variables at the species-specific scale, indicating that intraspecific variation affects the overall pattern of LDMC, P mass and N mass and P area. In terms of species-specific response, positive, negative or nonsignificant patterns were observed among the 92 species. Contrary to our prediction, species-specific responses within a functional group were not statistically more similar than those among functional groups. Our results indicated that leaf trait variation captured climatic difference yet species-specific responses were quite diverse irrespective of plant functional group, providing new insights for interpreting trait variability with climate.