Congruency analysis of species ranking based on leaf traits: which traits are the more reliable?

Nine leaf traits (area, fresh weight, dry weight, volume, density, thickness, specific leaf area (SLA), dry matter content (LDMC), leaf nitrogen content (LNC)) from ten plant species at eight sites in southern mediterranean France were investigated in order to assess their variability along a climatic gradient and their ranking congruency power. After examination of trait correlation patterns, we reduced the nine initial leaf traits to four traits, representative of three correlation groups: allometric traits (dry weight), functional traits (SLA and dry matter percentage) and Leaf Thickness. We analysed the variability of these four leaf traits at species and site level. We observed that between species variation (between 64.5 for SLA and 91% for LDMC) is higher than within species variation. Allowing a good congruency of species ranking assessed by spearman rank correlation () and a good reallocation of individuals to species by discriminant analysis. A site level variability (between 0.7% for Dry weight and 6.9% for SLA) was identified and environmental parameters (altitude, temperature, precipitation, nitrogen, pH) were considered as probable control factors. We found significant correlation between SLA, LDMC and the average minimum temperature (respectively r=0.87 and r=-0,9) and no correlation for the other traits or environmental parameters. Furthermore, we conclude that two leaf traits appear to be central in describing species: specific leaf area (SLA), percentage of dry matter (LDMC. While, SLA and LDMC are strongly correlated, LDMC appears to be less variable than SLA. According to our results the Dry Matter Content (or its reversal Leaf Water Content) appears the best leaf trait to be quantified for plant functional screening. Leaf thickness appeared to be rather uncorrelated with other leaf traits and show no environmental contingency; its variability could not have been explained in this study. Further studies should focus on this trait. This revised version was published online in June 2006 with corrections to the Cover Date.     

Contrasting effects of plant inter‐ and intraspecific variation on community trait responses to restoration of a sandy grassland ecosystem

Changes in plant community traits along an environmental gradient are caused by interspecific and intraspecific trait variation. However, little is known about the role of interspecific and intraspecific trait variation in plant community responses to the restoration of a sandy grassland ecosystem. We measured five functional traits of 34 species along a restoration gradient of sandy grassland (mobile dune, semi‐fixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China. We examined how community‐level traits varied with habitat changes and soil gradients using both abundance‐weighted and non‐weighted averages of trait values. We quantified the relative contribution of inter‐ and intraspecific trait variation in specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), and plant height to the community response to habitat changes in the restoration of sandy grassland. We found that five weighted community‐average traits varied significantly with habitat changes. Along the soil gradient in the restoration of sandy grassland, plant height, SLA, LDMC, and LCC increased, while LNC decreased. For all traits, there was a greater contribution of interspecific variation to community response in regard to habitat changes relative to that of intraspecific variation. The relative contribution of the interspecific variation effect of an abundance‐weighted trait was greater than that of a non‐weighted trait with regard to all traits except LDMC. A community‐level trait response to habitat changes was due largely to species turnover. Though the intraspecific shift plays a small role in community trait response to habitat changes, it has an effect on plant coexistence and the maintenance of herbaceous plants in sandy grassland habitats. The context dependency of positive and negative covariation between inter‐ and intraspecific variation further suggests that both effects of inter‐ and intraspecific variation on a community trait should be considered when understanding a plant community response to environmental changes in sandy grassland ecosystems.     

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.     

Trait‐abundance relation in response to nutrient addition in a Tibetan alpine meadow: The importance of species trade‐off in resource conservation and acquisition

In competition‐dominated communities, traits promoting resource conservation and competitive ability are expected to have an important influence on species relative abundance (SRA). Yet, few studies have tested the trait‐abundance relations in the line of species trade‐off in resource conservation versus acquisition, indicating by multiple traits coordination. We measured SRA and key functional traits involving leaf economic spectrum (SLA, specific leaf area; LDMC, leaf dry matter content; LCC, leaf carbon concentration; LNC, leaf nitrogen concentration; LPC, leaf phosphorus concentration; Hs, mature height) for ten common species in all plots subjected to addition of nitrogen fertilizer (N), phosphorus fertilizer (P), or both of them (NP) in a Tibetan alpine meadow. We test whether SRA is positively related with traits promoting plant resource conservation, while negatively correlated with traits promoting plant growth and resource acquisition. We found that species were primarily differentiated along a trade‐off axis involving traits promoting nutrient acquisition and fast growth (e.g., LPC and SLA) versus traits promoting resource conservation and competition ability (e.g., large LDMC). We further found that SRA was positively correlated with plant height, LDMC, and LCC, but negatively associated with SLA and leaf nutrient concentration irrespective of fertilization. A stronger positive height‐SRA was found in NP‐fertilized plots than in other plots, while negative correlations between SRA and SLA and LPC were found in N or P fertilized plots. The results indicate that species trade‐off in nutrient acquisition and resource conservation was a key driver of SRA in competition‐dominated communities following fertilization, with the linkage between SRA and traits depending on plant competition for specific soil nutrient and/or light availability. The results highlight the importance of competitive exclusion in plant community assembly following fertilization and suggest that abundant species in local communities become dominated at expense of growth while infrequent species hold an advantage in fast growth and dispersals to neighbor meta‐communities.     

Trait–performance relationships of grassland plant species differ between common garden and field conditions

The way functional traits affect growth of plant species may be highly context‐specific. We asked which combinations of trait values are advantageous under field conditions in managed grasslands as compared to conditions without competition and land‐use. In a two‐year field experiment, we recorded the performance of 93 species transplanted into German grassland communities differing in land‐use intensity and into a common garden, where species grew unaffected by land‐use under favorable conditions regarding soil, water, and space. The plants’ performance was characterized by two independent dimensions (relative growth rates (RGR) of height and leaf length vs. aboveground biomass and survival) that were differently related to the eight focal key traits in our study (leaf dry matter content (LDMC), specific leaf area (SLA), height, leaf anatomy, leaf persistence, leaf distribution, vegetative reproduction, and physical defense). We applied multivariate procrustes analyses to test for the correspondence of the optimal trait–performance relationships between field and common garden conditions. RGRs were species‐specific and species ranks of RGRs in the field, and the common garden were significantly correlated. Different traits explained the performance in the field and the common garden; for example, leaf anatomy traits explained species performance only in the field, whereas plant height was found to be only important in the common garden. The ability to reproduce vegetatively, having leaves that are summer‐persistent and with high leaf dry matter content (LDMC) were traits of major importance under both settings, albeit the magnitude of their influence differed slightly between the field and the common garden experiment. All optimal models included interactions between traits, pointing out the necessity to analyze traits in combination. The differences between field and common garden clearly demonstrate context dependency of trait‐based growth models, which results in limited transferability of favorable trait combinations between different environmental settings.     

Drought responses,phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites

• Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade‐off between drought survival and phenotypic plasticity.
• Throughout the summer we measured physiological traits (photosynthesis – A_max, stomatal conductance – g_s, transpiration – E, leaf water potential – ψl) and structural traits (specific leaf area – SLA, leaf density – LD, leaf dry matter content – LDMC, leaf relative water content – LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north‐ versus south‐facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period.
• We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade‐off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period.
• Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade‐off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.

     

An intraspecific application of the leaf‐height‐seed ecology strategy scheme to forest herbs along a latitudinal gradient

We applied the leaf‐height‐seed (LHS) ecology strategy scheme (a combination of three ecologically important traits: specific leaf area (SLA), seed mass and plant height) intraspecifically to two widespread European forest herbs along a latitudinal gradient. The aims of this study were to quantify LHS trait variation, disentangle the environmental factors affecting these traits and compare the within‐species LHS trait relationships with latitude to previously established cross‐species comparisons. We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900–2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4% increase in plant height with every degree northward shift (almost a two‐fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three‐factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring‐flowering Anemone was affected more by temperature, whereas early‐summer flowering Milium was affected more by local and other latitude‐related factors. Finally, previously reported cross‐species correlations between LHS traits and latitude were generally unsupported by our within‐species approach.     

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.     

Is there coordination of leaf and fine root traits at local scales? A test in temperate forest swamps

Examining the coordination of leaf and fine root traits not only aids a better understanding of plant ecological strategies from a whole‐plant perspective, but also helps improve the prediction of belowground properties from aboveground traits. The relationships between leaf and fine root traits have been extensively explored at global and regional scales, but remain unclear at local scales. Here, we measured six pairs of analogous leaf and fine root traits related to resource economy and organ size for coexisting dominant and subordinate vascular plants at three successional stages of temperate forest swamps in Lingfeng National Nature Reserve in the Greater Hinggan Mountains, NE China. Leaf and fine root traits related to resource acquisition (e.g., specific leaf area [SLA], leaf N, leaf P, root water content, and root P) decreased with succession. Overall, we found strong linear relationships between leaf dry matter content (LDMC) and root water content, and between leaf and root C, N, and P concentrations, but only weak correlations were observed between leaf area and root diameter, and between SLA and specific root length (SRL). The strong relationships between LDMC and root water content and between leaf and root C, N, and P held at the early and late stages, but disappeared at the middle stage. Besides, C and P of leaves were significantly correlated with those of roots for woody plants, while strong linkages existed between LDMC and root water content and between leaf N and root N for herbaceous species. These results provided evidence for the existence of strong coordination between leaf and root traits at the local scale. Meanwhile, the leaf–root trait relationships could be modulated by successional stage and growth form, indicating the complexity of coordination of aboveground and belowground traits at the local scale.     

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.     

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.     

Plasticity in above‐ and belowground resource acquisition traits in response to single and multiple environmental factors in three tree species

Functional trait plasticity is a major component of plant adjustment to environmental stresses. Here, we explore how multiple local environmental gradients in resources required by plants (light, water, and nutrients) and soil disturbance together influence the direction and amplitude of intraspecific changes in leaf and fine root traits that facilitate capture of these resources. We measured population‐level analogous above‐ and belowground traits related to resource acquisition, i.e. “specific leaf area”–“specific root length” (SLA–SRL), and leaf and root N, P, and dry matter content (DMC), on three dominant understory tree species with contrasting carbon and nutrient economics across 15 plots in a temperate forest influenced by burrowing seabirds. We observed similar responses of the three species to the same single environmental influences, but partially species‐specific responses to combinations of influences. The strength of intraspecific above‐ and belowground trait responses appeared unrelated to species resource acquisition strategy. Finally, most analogous leaf and root traits (SLA vs. SRL, and leaf versus root P and DMC) were controlled by contrasting environmental influences. The decoupled responses of above‐ and belowground traits to these multiple environmental factors together with partially species‐specific adjustments suggest complex responses of plant communities to environmental changes, and potentially contrasting feedbacks of plant traits with ecosystem properties. We demonstrate that despite the growing evidence for broadly consistent resource‐acquisition strategies at the whole plant level among species, plants also show partially decoupled, finely tuned strategies between above‐ and belowground parts at the intraspecific level in response to their environment. This decoupling within species suggests a need for many species‐centred ecological theories on how plants respond to their environments (e.g. competitive/stress‐tolerant/ruderal and response‐effect trait frameworks) to be adapted to account for distinct plant‐environment interactions among distinct individuals of the same species and parts of the same individual.     

黄土高原子午岭不同森林群落叶功能性状

对黄土高原子午岭地区5种主要森林群落中重要值>0.1的物种叶功能性状进行了比较,结果表明:1)叶干物质含量(LDMC)与比叶面积(SLA)呈负相关;SLA与叶厚度(LT)呈负相关,但与叶氮含量(LNC)和叶磷含量(LPC)呈正相关;LT与LNC、LPC和叶钾含量(LKC)呈负相关;LNC、LPC和LKC三者之间均呈正相关。2)坡位是影响叶大小(LS)、LT、LNC和LKC的主要地形因子,海拔对LPC和SLA的影响最大,而坡向则是影响LDMC的主要因子。因此,叶功能性状之间所呈现的特征及其对立地条件的适应都表现出了植物的生态策略。3)各群落间的乔木层叶功能性状均有显著差异,灌木层的叶功能性状无显著差异,而草本层除了LDMC无显著差异(P>0.05)外,其他叶功能性状均有显著差异,说明群落的叶功能性状的大小取决于群落内物种的叶功能性状及其重要值。     

Plant traits link hypothesis about resource-use and response to herbivory

Grazing by large herbivores, in interplay with environmental productivity, is a key driver of the composition of the vegetation with important consequences on the ecosystem and, consequently, for land management. We tested the predictions of the resource availability – resource–acquisition theory by assessing the extent to which community averages of plant traits, known to be related to plant growth, competitive ability and response to grazing were correlated with resource gradients within local (200 km²) geographical ranges. Second, we assessed the applicability of the same set of plant traits to make inferences on ecological effects of grazing by sheep in alpine ecosystems in Norway, using a data set consisting of 16 sites in central Norway. We estimated grazing intensity by free-ranging sheep based on GPS telemetry, soil properties, plant species composition and species traits i.e. specific leaf area (SLA), leaf dry matter content (LDMC), leaf size and plant height. Soil fertility and the interaction between soil fertility and grazing, but not grazing intensity alone, were significantly related to plant species and traits composition. Generally, average SLA showed lower correspondence with soil fertility and grazing than the other traits. Leaf size and plant height were lowest at sites with high grazing intensity and in sites with low fertility, and increased with soil fertility in little and moderately grazed sites, but declined at high fertility sites when grazing was intense. LDMC showed the opposite trend. Grazing intensity was more related to the variability in plant composition and average plant traits when environmental productivity was high. Our results therefore are indicative of a convergence of responses to grazing and nutrient limitation.     

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.     

Morphological and functional traits of herbaceous plants with different functional types in the European Northeast

We aimed to identify marker traits indicating the functional types of plants in the European Northeast. We try to answer the following questions. Which ecological factors make the largest contribution to identifying the functional types of plants in the North and can CO₂-exchange and related traits be used as markers? The data were collected from 1000-km latitudinal gradient across middle, north, and far north boreal forests in the east border of Europe. Comparative analysis of 102 species from 36 plant families enabled us to determine the marker traits indicating plant functional types. Competitor species have maximal plant height, comparatively low leaf dry matter content (LDMC), and accumulate high amounts of nitrogen in leaves. These species also have comparatively high photosynthetic and respiration rates. Ruderal species have low values of LDMC, and maximal photosynthetic rate, respiration rate, and photosynthetic nitrogen-use efficiency (PNUE). Slow-growing stress tolerators have a low photosynthetic rate, low respiration rate, and low levels of nitrogen and PNUE. The specific leaf area (SLA) of these plants shows a highly significant correlation with the light regime. In the boreal zone, SLA was found to be more closely related to light availability than to the plant functional type, indicating that SLA is unsuitable for use as a marker trait. We found strong correlations between plant height, respiration rate, and photosynthetic activity and soil nutrition according to Ellenberg values. Soil mineral element contents and acidity were found to have a significant influence on the functional types of plants.     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.     

Intraspecific trait variability shapes leaf trait response to altered fire regimes

Background and AimsUnderstanding impacts of altered disturbance regimes on community structure and function is a key goal for community ecology. Functional traits link species composition to ecosystem functioning. Changes in the distribution of functional traits at community scales in response to disturbance can be driven not only by shifts in species composition, but also by shifts in intraspecific trait values. Understanding the relative importance of these two processes has important implications for predicting community responses to altered disturbance regimes.MethodsWe experimentally manipulated fire return intervals in replicated blocks of a fire-adapted, longleaf pine (Pinus palustris) ecosystem in North Carolina, USA and measured specific leaf area (SLA), leaf dry matter content (LDMC) and compositional responses along a lowland to upland gradient over a 4 year period. Plots were burned between zero and four times. Using a trait-based approach, we simulate hypothetical scenarios which allow species presence, abundance or trait values to vary over time and compare these with observed traits to understand the relative contributions of each of these three processes to observed trait patterns at the study site. We addressed the following questions. (1) How do changes in the fire regime affect community composition, structure and community-level trait responses? (2) Are these effects consistent across a gradient of fire intensity? (3) What are the relative contributions of species turnover, changes in abundance and changes in intraspecific trait values to observed changes in community-weighted mean (CWM) traits in response to altered fire regime?Key ResultsWe found strong evidence that altered fire return interval impacted understorey plant communities. The number of fires a plot experienced significantly affected the magnitude of its compositional change and shifted the ecotone boundary separating shrub-dominated lowland areas from grass-dominated upland areas, with suppression sites (0 burns) experiencing an upland shift and annual burn sites a lowland shift. We found significant effects of burn regimes on the CWM of SLA, and that observed shifts in both SLA and LDMC were driven primarily by intraspecific changes in trait values.ConclusionsIn a fire-adapted ecosystem, increased fire frequency altered community composition and structure of the ecosystem through changes in the position of the shrub line. We also found that plant traits responded directionally to increased fire frequency, with SLA decreasing in response to fire frequency across the environmental gradient. For both SLA and LDMC, nearly all of the observed changes in CWM traits were driven by intraspecific variation.     

Leaf trait co‐variation,response and effect in a chronosequence

Question: Is there any generality in terms of leaf trait correlations and the multiple role of leaf traits (response to and/or effect on) during secondary succession? Location: A secondary successional sere was sampled at four different ages since abandonment from several years to nearly 150 years on the Loess Plateau of northwestern China. Method: Specific leaf area (SLA), leaf mass per area (LMA), leaf nitrogen (N_mass, N_area), leaf phosphorus (P_mass, P_area) and leaf dry matter content (LDMC) were measured for all species recorded in the successional sere. Above‐ground net primary productivity (ANPP) and specific rate of litter mass loss (SRLML) were measured as surrogates for ecosystem properties. Soil total carbon (C) and nitrogen (N) were measured in each stage. Leaf traits were related to ecosystem properties and soil nutrient gradients, respectively. Results: LMA is correlated with N_area and P_area' and negatively with N_mass. Correlation between N_area and P_area was higher than between N_mass and P_mass. At the community level, field age, community hierarchy and their interaction explain 64.4 ‐ 93.5% of the variation in leaf traits. At the species level, field age explains 22.4 ‐ 45.5% of the variation in leaf traits (excl. P_area) while plant functional group has a significant effect only for N_mass. LDMC is correlated with ANPP and negatively with SRLML; P_mass is correlated with SRLML. Conclusions: Mean values of LMA, N_mass and N_area are close to the worldwide means, suggesting that large‐scale climate has a profound effect on leaf mass and leaf nitrogen allocation, while environmental gradients represented by succession have little influence on leaf‐trait values. Correlations between leaf traits, such as LMA‐N_area, LMA‐P_area and LMA‐N_mass shown in previous studies, are confirmed here. Although none of the leaf traits is proved to be both a response trait and an effect trait independent of time scale and community hierarchy, mass‐based leaf N is likely a sensitive response trait to soil C and N gradients. In addition, LDMC can be a marker for ANPP and SRLML, while mass‐based leaf P can be a marker for SRLML.