Intraspecific trait variation and reversals of trait strategies across key climate gradients in native Hawaiian plants and non-native invaders

Background and AimsDisplacement of native plant species by non-native invaders may result from differences in their carbon economy, yet little is known regarding how variation in leaf traits influences native–invader dynamics across climate gradients. In Hawaii, one of the most heavily invaded biodiversity hotspots in the world, strong spatial variation in climate results from the complex topography, which underlies variation in traits that probably drives shifts in species interactions.MethodsUsing one of the most comprehensive trait data sets for Hawaii to date (91 species and four islands), we determined the extent and sources of variation (climate, species and species origin) in leaf traits, and used mixed models to examine differences between natives and non-native invasives.Key ResultsWe detected significant differences in trait means, such that invasives were more resource acquisitive than natives over most of the climate gradients. However, we also detected trait convergence and a rank reversal (natives more resource acquisitive than invasives) in a sub-set of conditions. There was significant intraspecific trait variation (ITV) in leaf traits of natives and invasives, although invasives expressed significantly greater ITV than natives in water loss and photosynthesis. Species accounted for more trait variation than did climate for invasives, while the reverse was true for natives. Incorporating this climate-driven trait variation significantly improved the fit of models that compared natives and invasives. Lastly, in invasives, ITV was most strongly explained by spatial heterogeneity in moisture, whereas solar energy explains more ITV in natives.ConclusionsOur results indicate that trait expression and ITV vary significantly between natives and invasives, and that this is mediated by climate. These findings suggest that although natives and invasives are functionally similar at the regional scale, invader success at local scales is contingent on climate.     

Functional traits but not environmental gradients explain seed weight in Mongolian plant species

• Seed weight varies by several orders of magnitude among vascular plant species. However, the importance of potential drivers such as environmental conditions and plant functional traits have rarely been assessed for a larger taxonomic sample.
• We collected seeds of 148 species from 237 sites spread across Mongolia and compared their weight among the major zonal vegetation types, taxonomic groups and a set of functional traits (growth form, dispersal mode, fruit type, storage organs and palatability).
• Seed weight strongly varied among all functional traits and taxonomic groups, but no differences among vegetation zones were detected.
• These results suggest a low impact of environmental conditions on the evolution of seed weight, contrasting the strong phylogenetic signal.

     

Individual‐level leaf trait variation and correlation across biological and spatial scales

Even with increasing interest in the ecological importance of intraspecific trait variation (ITV) for better understanding ecological processes, few studies have quantified ITV in seedlings and assessed constraints imposed by trade‐offs and correlations among individual‐level leaf traits. Estimating the amount and role of ITV in seedlings is important to understand tree recruitment and long‐term forest dynamics. We measured ten different size, economics, and whole leaf traits (lamina and petiole) for more than 2,800 seedlings (height ≥ 10 cm and diameter at breast height < 1 cm) in 283 seedling plots and then quantified the amount of ITV and trait correlations across two biological (intraspecific and interspecific) and spatial (within and among plots) scales. Finally, we explored the effects of trait variance and sample size on the strength of trait correlations. We found about 40% (6%–63%) variation in leaf‐level traits was explained by ITV across all traits. Lamina and petiole traits were correlated across biological and spatial scales, whereas leaf size traits (e.g., lamina area) were weakly correlated with economics traits (e.g., specific lamina area); lamina mass ratio was strongly related to the petiole length. Trait correlations varied among species, plots, and different scales but there was no evidence that the strength of trait relationships was stronger at broader than finer biological and spatial scales. While larger trait variance increased the strength of correlations, the sample size was the most important factor that was negatively related to the strength of trait correlations. Our results showed that a large amount of trait variation was explained by ITV, which highlighted the importance of considering ITV when using trait‐based approaches in seedling ecology. In addition, sample size was an important factor that influenced the strength of trait correlations, which suggests that comparing trait correlations across studies should consider the differences in sample size.     

Trait coordination and environmental filters shape functional trait distributions of forest understory herbs

1. Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.
2. To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ¹³C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.
3. Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.
4. Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.
5. Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     

Plant invasion alters trait composition and diversity across habitats

Increased globalization has accelerated the movement of species around the world. Many of these nonnative species have the potential to profoundly alter ecosystems. The mechanisms underpinning this impact are often poorly understood, and traits are often overlooked when trying to understand and predict the impacts of species invasions on communities. We conducted an observational field experiment in Canada's first National Urban Park, where we collected trait data for seven different functional traits (height, stem width, specific leaf area, leaf percent nitrogen, and leaf percent carbon) across an abundance gradient of the invasive Vincetoxicum rossicum in open meadow and understory habitats. We assessed invasion impacts on communities, and associated mechanisms, by examining three complementary functional trait measures: community‐weighted mean, range of trait values, and species’ distances to the invader in trait space. We found that V. rossicum invasion significantly altered the functional structure of herbaceous plant communities. In both habitats V. rossicum changed the community‐weighted means, causing invaded communities to become increasingly similar in their functional structure. In addition, V. rossicum also reduced the trait ranges for a majority of traits indicating that species are being deterministically excluded in invaded communities. Further, we observed different trends in the meadow and understory habitats: In the understory, resident species that were more similar to V. rossicum in multivariate trait space were excluded more, however this was not the case in the meadow habitat. This suggests that V. rossicum alters communities uniquely in each habitat, in part by creating a filter in which only certain resident species are able to persist. This filtering process causes a nonrandom reduction in species' abundances, which in turn would be expected to alter how the invaded ecosystems function. Using trait‐based frameworks leads to better understanding and prediction of invasion impacts. This novel framework can also be used in restoration practices to understand how invasion impacts communities and to reassemble communities after invasive species management.     

典型草原围封后羊草地上功能性状对枯落物累积的响应

植物功能性状反映植物适应环境变化过程中在不同器官形态与功能间的资源权衡与分配策略。典型草原围封后枯落物累积导致群落光照、热量和水分的重新分配并改变微环境特征。在此过程中植物地上功能性状将通过怎样的变化来适应新的环境,目前尚不清楚。2015—2017年每年8月对内蒙古地区3种典型草原共有物种羊草（Leymus chinensis）的植株、叶片和茎干功能性状进行了测量与分析。结果表明：枯落物累积显著增加了羊草的植株高度、单株重量、茎叶比和总叶面积;枯落物累积显著增加了羊草的叶片长度、叶片重量、单叶面积、节间长度和茎干重量,这些性状属于敏感性状;枯落物累积对羊草的叶片数量和节间数量无显著影响,相对而言,这些性状属于惰性性状;羊草的单株重量与植株高度、叶片重量呈极显著的正相关关系（P<0.0001）;羊草的植株高度与节间数量、节间长度呈极显著的正相关关系（P<0.0001）。本研究结果从植物地上功能性状的角度阐明了典型草原植物对环境变化的适应方式,可为围封草原的合理管理提供基础数据与理论依据。     

Leaf decomposition depends on nutritional trait values but increasing trait variability does not always increase decomposition efficiency

The mass ratio (MRH) and niche complementarity (NCH) hypotheses can explain how leaf trait composition drives decomposition, an ecosystem process linked to nutrient cycling and carbon sequestration. However, few studies have used an experiment designed to disentangle the role of the mechanisms proposed by these hypotheses. This is especially true regarding the role of leaf functional traits for decomposition rates in tropical ecosystems. Here, we quantified the biomass loss of 120 leaf mixtures assembled according to four quasi-orthogonal combinations of different mean trait values (community-weighted mean; CWM) and trait variability (functional diversity; FD) of three leaf functional traits (leaf nitrogen and leaf magnesium concentrations and specific leaf area). We found that CWM values of leaf nutritional traits were positively related to greater biomass loss. This supports the hypothesis that the mean trait values of leaf mixtures can drive biomass loss (MRH). However, contrary to NCH expectations, in some circumstances, increasing trait variability of leaf nutritional traits decreased biomass loss. Our results reinforce some previous evidence that, together, CWM and FD can explain leaf decomposition and highlight that the mean resource quality of leaf mixtures is a driver of biomass loss. Also, as previously reported for temperate ecosystems, trait variability does not always increase leaf decomposition in tropical ecosystems. Therefore, there is a need to consider simultaneously both MRH and NCH in future studies, using an appropriate design, keeping in mind that both mechanisms will always be present in any species mixture or combination.     

Coordination of photosynthetic traits across soil and climate gradients

“Least-cost theory” posits that C₃ plants should balance rates of photosynthetic water loss and carboxylation in relation to the relative acquisition and maintenance costs of resources required for these activities. Here we investigated the dependency of photosynthetic traits on climate and soil properties using a new Australia-wide trait dataset spanning 528 species from 67 sites. We tested the hypotheses that plants on relatively cold or dry sites, or on relatively more fertile sites, would typically operate at greater CO₂ drawdown (lower ratio of leaf internal to ambient CO₂, C_i:C_a) during light-saturated photosynthesis, and at higher leaf N per area (N_area) and higher carboxylation capacity (V_{cmax 25}) for a given rate of stomatal conductance to water vapour, g_sw. These results would be indicative of plants having relatively higher water costs than nutrient costs. In general, our hypotheses were supported. Soil total phosphorus (P) concentration and (more weakly) soil pH exerted positive effects on the N_area–g_sw and V_{cmax 25}–g_sw slopes, and negative effects on C_i:C_a. The P effect strengthened when the effect of climate was removed via partial regression. We observed similar trends with increasing soil cation exchange capacity and clay content, which affect soil nutrient availability, and found that soil properties explained similar amounts of variation in the focal traits as climate did. Although climate typically explained more trait variation than soil did, together they explained up to 52% of variation in the slope relationships and soil properties explained up to 30% of the variation in individual traits. Soils influenced photosynthetic traits as well as their coordination. In particular, the influence of soil P likely reflects the Australia's geologically ancient low-relief landscapes with highly leached soils. Least-cost theory provides a valuable framework for understanding trade-offs between resource costs and use in plants, including limiting soil nutrients.     

Intraspecific trait variation in three common grass species reveals finescale species adjustment to local environmental conditions

Aims Studies along environmental gradients have shown that intraspecific trait variation (ITV) may contribute considerably to community-level trait variation. However, we lack knowledge about how the extent of ITV varies on a local scale and whether a varying extent of ITV is related to differences in local environmental site and plant community characteristics.     

Microanatomical traits track climate gradients for a dominant C4 grass species across the Great Plains,USA

Background and AimsAndropogon gerardii is a highly productive C₄ grass species with a large geographic range throughout the North American Great Plains, a biome characterized by a variable temperate climate. Plant traits are often invoked to explain growth rates and competitive abilities within broad climate gradients. For example, plant competition models typically predict that species with large geographic ranges benefit from variation in traits underlying high growth potential. Here, we examined the relationship between climate variability and leaf-level traits in A. gerardii, emphasizing how leaf-level microanatomical traits serve as a mechanism that may underlie variation in commonly measured traits, such as specific leaf area (SLA).MethodsAndropogon gerardii leaves were collected in August 2017 from Cedar Creek Ecosystem Science Reserve (MN), Konza Prairie Biological Station (KS), Platte River Prairie (NE) and Rocky Mountain Research Station (SD). Leaves from ten individuals from each site were trimmed, stained and prepared for fluorescent confocal microscopy to analyse internal leaf anatomy. Leaf microanatomical data were compared with historical and growing season climate data extracted from PRISM spatial climate models.Key ResultsMicroanatomical traits displayed large variation within and across sites. According to AICc (Akaike’s information criterion adjusted for small sample sizes) selection scores, the interaction of mean precipitation and temperature for the 2017 growing season was the best predictor of variability for the anatomical and morphological traits measured here. Mesophyll area and bundle sheath thickness were directly correlated with mean temperature (annual and growing season). Tissues related to water-use strategies, such as bulliform cell and xylem area, were significantly correlated with one another.ConclusionsThe results indicate that (1) microanatomical trait variation exists within this broadly distributed grass species, (2) microanatomical trait variability appears likely to impact leaf-level carbon and water use strategies, and (3) microanatomical trait values vary across climate gradients, and may underlie variation in traits measured at larger ecological scales.     

不同生活型被子植物功能性状与基因组大小的关系

基因组大小在被子植物物种之间存在着巨大的变异, 但目前对不同生活型被子植物功能性状与基因组大小的关系缺乏统一的认识。本研究基于被子植物245科2,226属11,215个物种的基因组大小数据, 探讨了不同生活型物种种子重量、最大植株高度和叶片氮、磷含量4个功能性状与基因组大小之间的关系。结果表明, 被子植物最大植株高度和种子重量与基因组大小间的关系在草本和木本植物中存在显著差异。草本植物最大植株高度与基因组大小的关系不显著, 但种子重量与其呈极显著的正相关关系。木本植物最大植株高度与基因组大小显著负相关, 但种子重量与其关系不显著。木本植物叶片氮含量与基因组大小呈显著正相关, 但其他生活型植物的叶片氮、磷含量与基因组大小均无显著相关性。本研究表明被子植物功能性状与基因组大小的相关性在不同生活型间存在差异, 这为深入研究植物多种功能性状和植物生活型与基因组大小的权衡关系在植物演化和生态适应中的作用提供了重要依据。     

Leading trait dimensions in flood-tolerant plants

Background and AimsWhile trait-based approaches have provided critical insights into general plant functioning, we lack a comprehensive quantitative view on plant strategies in flooded conditions. Plants adapted to flooded conditions have specific traits (e.g. root porosity, low root/shoot ratio and shoot elongation) to cope with the environmental stressors including anoxic sediments, and the subsequent presence of phytotoxic compounds. In flooded habitats, plants also respond to potential nutrient and light limitations, e.g. through the expression of leaf economics traits and size-related traits, respectively. However, we do not know whether and how these trait dimensions are connected.MethodsBased on a trait dataset compiled on 131 plant species from 141 studies in flooded habitats, we quantitatively analysed how flooding-induced traits are positioned in relation to the other two dominant trait dimensions: leaf economics traits and size-related traits. We evaluated how these key trait components are expressed along wetness gradients, across habitat types and among plant life forms.Key ResultsWe found that flooding-induced traits constitute a trait dimension independent from leaf economics traits and size-related traits, indicating that there is no generic trade-off associated with flooding adaptations. Moreover, individual flooding-induced traits themselves are to a large extent decoupled from each other. These results suggest that adaptation to stressful environments, such as flooding, can be stressor specific without generic adverse effects on plant functioning (e.g. causing trade-offs on leaf economics traits).ConclusionsThe trait expression across multiple dimensions promotes plant adaptations and coexistence across multifaceted flooded environments. The decoupled trait dimensions, as related to different environmental drivers, also explain why ecosystem functioning (including, for example, methane emissions) are species and habitat specific. Thus, our results provide a backbone for applying trait-based approaches in wetland ecology by considering flooding-induced traits as an independent trait dimension.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.     

藏东木本植物群落功能性状分布与环境的关系

研究植物群落功能性状间的相关关系及其对环境变化的响应,能够有效揭示植物功能性状的权衡模式及其对环境的适应策略。藏东昌都地区位于横断山脉西北部,复杂气候地貌孕育了丰富的植物资源,是青藏高原森林灌丛生态系统主要组分和国际生物多样性保护的热点地区。以藏东森林灌丛群落优势木本植物为研究对象,在大量野外调查基础上,采用相关分析、主成分分析、线性回归和方差分析等方法,研究了该区域植物功能性状间的相关关系、功能性状对环境变量的响应规律以及功能性状的变异来源。结果表明：(1)藏东木本植物表现出适应高寒环境的性状权衡模式,即：比叶面积、叶体积较小而叶干物质含量较大,叶磷含量和叶钾含量协同变化;(2)海拔和气候变量共同驱动着藏东木本植物功能性状的变化,并且藏东木本植物倾向于采取“高投入—慢回报”提高御寒能力的保守型适应策略;(3)海拔是影响藏东植物功能性状变异最显著的环境变量,种间变异在藏东植物群落功能性状随环境变化中起主要作用。研究结果揭示了藏东木本植物功能性状的权衡模式及其对高寒环境的适应策略,有助于加深对藏东自然植物资源分布规律和生态功能的认识,为区域生态系统功能和生物多样性保护提供科学依据。